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Functional brain systems and personality dynamics

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Functional brain systems and personality dynamics
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Lindquist, David, 1944-
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English
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ix, 188 leaves : ill. ; 28 cm.

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Amygdala ( jstor )
Cognitive psychology ( jstor )
Emotional expression ( jstor )
Hemispheres ( jstor )
Hippocampus ( jstor )
Lesions ( jstor )
Memory ( jstor )
Mental stimulation ( jstor )
Personality psychology ( jstor )
Psychology ( jstor )
Dissertations, Academic -- Psychology -- UF
Personality ( lcsh )
Psychology thesis Ph. D
Psychology, Pathological ( lcsh )
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bibliography ( marcgt )
theses ( marcgt )
non-fiction ( marcgt )

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Thesis:
Thesis (Ph. D.)--University of Florida, 1985.
Bibliography:
Includes bibliographical references (leaves 168-187).
Additional Physical Form:
Also available online.
General Note:
Typescript.
General Note:
Vita.
Statement of Responsibility:
by David Lindquist.

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University of Florida
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Copyright [name of dissertation author]. Permission granted to the University of Florida to digitize, archive and distribute this item for non-profit research and educational purposes. Any reuse of this item in excess of fair use or other copyright exemptions requires permission of the copyright holder.
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FUNCTIONAL BRAIN SYSTEMS AND PERSONALITY DYNAMICS











By

DAVID LINDQUIST




















A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN
PARTIAL FULFILLMENT OF THE REQUIREMENTS
FOR THE DEGREE OF DOCTOR OF PHILOSOPHY


UNIVERSITY OF FLORIDA 1985






















































Copyright 1985

by

David Lindquist














ACKNOWLEDGEMENTS

I am deeply grateful to Max and Ruth Lindquist for their patience and the opportunities they gave me. My debt to my undergraduate mentor, Dr. Carol Van Hartesveldt, who taught me about science and let me make my own mistakes, is also gratefully acknowledged.

I want to thank my doctoral committee members, Doctors Grater,

Morgan, Schauble, Suchman and Ziller, who were also my most respected graduate teachers. Special thanks are due to David Suchman, who first encouraged me in this project, and to my chairman, Harry Grater, who shepherded me through the crises with grace and allowed me to

keep my dignity.

The help of Dr. Bill Froming in the design of this study, and of Mssrs. Denny Gies, Bill Baxter and Ted Shaw (of the North Florida

Evaluation and Treatment Center) and of Ms. Janet Despard (of Mental Health Services, Inc.) in facilitating its execution, is much appreciated. I am especially grateful to Ms. Cheryl Shaw, who tyDed the manuscript, and without whose friendship and organizational help this project would not have been completed.

Warmest heartfelt gratitude goes to my dear friends Gabriel Rodriguez, Marshall and Laura Knudson, and David Kurtzman, whose love sustained me through these difficult years.

I can never properly express my thanks to Ruth Lindquist, to whom this piece of work is lovingly dedicated.


iii














TABLE OF CONTENTS


ACKNOWLEDGEMENTS ................................................. iii

LIST OF FIGURES .................................................. vii

ABSTRACT ........................................................ viii

CHAPTER

I INTRODUCTION ......................................... 1

II THE PHYSIOLOGICAL SUBSTRATE OF PERSONALITY:
A SELECTIVE REVIEW OF THE LITERATURE ................. 7

Review of Basic Brain Anatomy and Organization ....... 8
Cortical Mechanisms ............................... 8
Subcortical Systems .............................. 12
The Interpretation of Neurology Literature ....... 17
Human Consciousness ................................. 18
Right versus Left ................................ 18
Language and the Left Hemisphere ................. 20
Aphasia: Anatomy and Syndromes .................. 21
Language, Symbolism, and Meaning ................. 24
Human Consciousness, Self-Awareness,
and Thought ...................................... 26
Discussion ............................. 35
Cortical Mobilization: Attention, Arousal,
and Activation ...................................... 39
The Reticular Activating System and
Tonic Arousal ................................. ... 39
Phasic Control Systems: The Frontal Lobes
and Thalamus ..................................... 41
Discussion ....................................... 44
Motivation: Emotion and Affect ..................... 45
Amygdala Circuits and the Prefrontal Lobes ....... 46 Affective Expression ............................. 48
Discussion ....................................... 50
Memory Functions .................................... 51
Human Amnesia Syndromes .......................... 57
Memory and the Neocortex ......................... 63
Discussion ....................................... 68
The Limbic System, RAS, and Memory ............... 70
Papez Circuit and Memory ......................... 74
Fornix ........................................ 75


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Mammillary bodies and mammillothalamic tract. .80 Cingulate cortex .......................... 81
Discussion................................... 83
Interfaces and Interactions of the Monitoring,
Motivating, and Mobilization Systems.............. 89
The Biochemistry of Emotion, Motivation,
and Learning................................. 92
Emotion, Amygdala Circuits and Memory........9 Discussion.................................. 103
Biochemical and Electrophysiological Aspects
of Cortical Mobilization Processes............ 104
The Functional Elements of the Personality
Structure ..................................... 108
The Problem-Solving/Response-Generating
System ..................................... 109
The Memory System ........................... 110
The Motivating System........................ 112
The Mobilization System...................... 114
Learning and Memory: Animal Studies ...........115
A Functional Meta-system........................ 120
Lateralized Mobilization Processes............ 125
Asymmetrical reaction time to laterally
presented stimuli......................... 125
Altered GSR following unilateral brain
injury................................... 126
Asymmetrical biochemical and electrophysiological processes ................... 127
Bilateral Interaction in Emotion and Cognition. .128
Mental Health and Psychopatholgy ................ 136
The Psychopathological Correlates of
Unilateral Temporal Lobe Epilepsy............. 139
Discussion: Hyper- and Hypo-dominance
Spectrum Disorders .......................... 141
Schizophrenia and the Affective Disorders .......143
Anxiety Disorders, Obsessive-Compulsive
Illness, and Paranoia........................ 145
Sociopathy and Hysteria...................... 147

III METHOD ........................................ 151

Subjects.................................... 152
Instruments................................. 153
Procedure................................... 154
Hypotheses.................................. 155

IV RESULTS ....................................... 156

Left versus Right Hemisphere Cognitive
Functioning Between Groups ................... 156
Overall Performance on the Tests Sensitive to
Right versus Left Hemisphere Cognitive
Functioning................................. 157

v









Differences in MMPI Scores ................... 157
Age........................................... 159

V DISCUSSION...................................... 160

APPENDICES

A HYPO- AND HYPER-DOMINANCE SPECTRUM DISORDERS BY
DSM III DIAGNOSTIC CLASSIFICATION .................165

B INFORMED CONSENT STATEMENT....................... 166

C REVISED SCORING CRITERIA FOR THE STREET (1931)
GESTALT COMPLETION TEST.......................... 167

BIBLIOGRAPHY ................................................. 168

BIOGRAPHICAL SKETCH........................................... 188



































vi














LIST OF FIGURES


FIGURES

1 Cytoarchitectural map of the lateral and medial
surfaces of the human cerebral cortex, with numbers
representing the areas of Brodman .................. 10

2 Partially schematized representation of the
limbic system .................................... 13

3 The position of Papez's circuit within a
larger cortical circuit ........................... 86

4 Schematic representation of the proposed model of
the physiological substrate of personality ..........111




























vii














Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy

FUNCTIONAL BRAIN SYSTEMS AND PERSONALITY DYNAMICS

By

David Lindquist

May 1985
Chairman: Dr. Harry Grater
Major Department: Psychology

A heuristic model of the physiological substrate of personality

structure was developed from a review of recent neurological and physiological psychology literature. The formulation of the model was based on assumptions that the phylogenetic transition from instinctive responding to self-determined behavior required the evolution of automatic neural mechanisms to insure that the organism would monitor the environment for significant stimuli, be motivated to respond in the presence of those stimuli, and mobilize the appropriate psychological operations to determine the form of that response. Functional elements of these basic mechanisms were identified and a functional meta-system was outlined which would organize the elements and optimize the -utilization of lateralized cognitive processes in the interest of assuring the emission of adaptive behavior.

The proposed model suggests that certain psychodiagnostic entities might be classified as hyper- and hypo-dominance spectrum disorders viii










based on the form of dysfunction within the meta-system. The ability of the model to predict membership in diagnostic categories was tested by assigning 42 adult psychiatric inpatient and outpatient subjects to hyper-dominant and hypo-dominant groups by diagnosis, according to the constructs of the model, and comparing performance on instruments shown to be sensitive to right and left cerebral hemisphere dysfunction. The Street Gestalt Completion Test and the Object Assembly, Similarities, and Information Subtests of the Wechsler Adult Intelligence Scale-Revised were administered to each subject. An abbreviated form of the Minnesota Multiphasic Personality Inventory (MMPI) was used to compare symptomology between groups.

Significant between-group differences (p < 0.001) in the ratios of test scores sensitive to right versus left cognitive functioning were found in the predicted directions, while the groups did not differ in overall performance on the instruments. Significant differences (p< 0.005) in the ratios of selected MMPI clinical scales, in the predicted direction, provided further support for the hypothesized relationship between lateralized cognitive functioning, symptomology, and diagnosis.

It was concluded that the proposed model provides tenable and potentially useful operational definitions of personality functions and psychopathology. Results were discussed in terms of their imrplications for psychotherapeutic interventions and additional methods to test the validity of the model.


ix














CHAPTER I
INTRODUCTION


The various schools of psychotherapy agree that the practitioner's task is to facilitate change in the client. They agree on little else. There is, as yet, no consensus regarding the two major issues in psychotherapy: what is to be changed, and how that change is to be brought about (Strupp, 1978). Strong opinions about both of these basic problems are available; data based conclusions are not. ProDonents of fundamentally different viewpoints debate acrimoniously (Eysenck, 1974); yet verifiable differences in outcome between theorybased forms of intervention are rare (Bergin & Lambert, 1978).

This unsatisfactory state of affairs in applied psychology has unfortunate consequences for practitioners and their clients alike.

The problem has been ascribed by Watson to the "preparadigmatic"

state of the discipline of psychology. According to Watson "psychology has not experienced anything comparable to what atomic theory has done for biology, what laws of motion have done for physics. Either psychology"s first paradigm has not been discovered yet or it has not been recognized for what it is" (Watson, 1967, p. 53). Hanson stated the problem succinctly, "The issue is not theory using, but theory finding" (1965, p. 3).

The two major forces in psychological thought, psychoanalysis and behaviorism, have encountered significant problems. Theories


1





2



based on the former have been criticized as untestable and therefore unscientific (Eysenck, 1970). The latter movement lost impetus with the discovery by Olds and Milner (1954) of "pleasure" centers in the brain. This revelation undermined the basic assumption of the learning theorists that behavior could be explained simply by defining the rules governing stimulus-response relationships.

Successful scientific theories are built on paradigms that

describe the fundamental properties and mechanics of their subject. The fundamental units of personality are networks of neurons in the brain. Sigmund Freud (1948) attempted to relate mental structures to anatomical locations but was forced to abandon his effort because the neurology of the time was not adequate. Instead he and subsequent theorists were forced to base their models on suppositions about the products of the personality processes. As noted above, the results have been less than satisfactory. The science of neurology has made significant progress in the interim and a large amount of useful information has accumulated. These data have been virtually ignored by the discipline of psychology. The integration of neurological data and psychological theory may provide a basis for a useful paradigm for the psychotherapist. The present work is intended as a step toward such an integration.

The purpose of this study is to develop and test a heuristic,

model of personality function, based on an understanding of its physiological substrate, with the ultimate goal of inDroving the effectiveness of psychotherapeutic interventions. Such a model

should identify the basic elements and processes of the personality





3


structure and describe the ways in which these interact to produce psychological health and psychopathology. Such a model might lead to new operational definitions of psychological phenomena which, in turn, may suggest new intervention points and methods.

The purpose of a theory is to integrate known facts within a

single framework and account for them in terms of a small number of interrelated concepts. Existing theories suffer from a lack of integration. The discipline of psychology has hindered such integration by institutionalizing a tradition of subspecializatiOn: theories of attention, perception, cognition, motivation, emotion, behavior, etc. are developed in relative isolation. The failure of individual theorists to understand and to appreciate the interrelatedness of these various personality operations may account for the inadequacy of psychological theory in general. It is assumed here that the component parts of the personality can only be properly understood in the context of their relationship to the whole; that more will be gained from a gross (but comprehensive and testable) description of the personality infrastructure than from a detailed examination of a single personality operation.

Rather than subdivide the personality structure on the basis of notions of what it should do; it might be more productive to approach the problem on the more basic level of what it must do. The formulation of the model will be guided by a set of assumptions about the evolutionary pressures which shaped the personality structure. These "evolutionary imperatives" are as follows:






4


1. The physiological substrate of personality evolved to

assure the emission of adaptive behavior by the organism. Survival pressures shaped an organization that was able to respond effectively to significant stimuli and produce behavior that enhanced the organism's well-being.

2. The evolution of this substrate proceeded from an organization based on "hard-wired" stimulus-response instincts to an organization which allowed increased latitude in behavior in order to take advantage of the organism's developing problem-solving abilities.

3. In order to permit self-determination of behavior and,

at the same time, to insure survival, new mechanisms were required to assure that the organism would (a) attend to significant stimuli; (b) be motivated to respond effectively to those stimuli;

(c) accomplish the mobilization of the necessary psychological resources to determine the form of that response (referred to hereafter as the Monitoring, Motivating, and Mobilization systems).

4. Although these systems interact, they must be functionally separated to the extent that they do not interfere with each other's normal operation. Similar functions might be carried out in other brain areas, but the automatic activation of these systems will assure that they dominate responding to stimuli which relate to the survival and well-being of the organism.

5. As these mechanisms are essential for the survival of the individual and species they will form the central organizing processes-of personality; personality dynamics will center on their operation.





5


Neurological theories of mental function center on clinical

observations of patients with localized brain lesions. The history of neurological thought has revolved around the question of how these data are to be interpreted. For many years higher mental functions were treated as discrete "faculties." The results were inconsistent and of little value. The failures of these "narrow localizers" led to theories which attempted to account for mental functions on the basis of the "mass action" of the brain. Where earlier models were too specific, these theories proved too general to be useful.

In the 1920s Goldstein broke with the tradition of attempting to infer functions directly from deficits and proposed instead an "analysis of basic disturbances." This approach led finally to Luria's conceptualization of mental activities as the product of the interaction of complex functional systems. In Luria's formulation a mental function is the result of contributions from a number of concertedly working zones. Therefore, that function may be destroyed, or disturbed differently, by lesions in different locations. Luria (1973a) described the characteristics of a functional system:

The presence of a constant (invariant) task, performed
by variable (variative) mechanisms, bringing the process
to a constant (invariant) result, is one of the basic features distinguishing the work of every "functional
system." The second distinguishing feature is the
complex composition of the "functional system," which always includes a series of afferent (adjusting) and efferent (effector) impulses. (Luria, 1973a, p. 28)

Luria outlined three principal functional units in the brain: the "units for regulating tone and waking and mental states," centered on the reticular activating system in the brainstem; the





6


"unit for receiving, analyzing and storing information," operating in the post-central (sensory) areas of the brain; and the "unit for programming, regulation and verification of activity," operating in the frontal lobes (Luria, 1973(i. ch. 2).

Luria's concepts represent a major advance in the understanding of the fundamental operating characteristics of brain systems. Although his formulations are too basic to be of much use to the applied psychologist, he has established a format and a methodology which will be followed here. The present investigation will focus on identifying and describing the interactions of the functional brain systems which satisfy the requirements of the evolutionary imperatives outlined above. Evidence suggests that the substrate for these systems will be found in those anatomical areas for which Luria acknowledged he had inadequate data for his own analyses: the medio-basal zones of the cortex and the right hemisphere of the brain.














CHAPTER II
THE PHYSIOLOGICAL SUBSTRATE OF PERSONALITY:
A SELECTIVE REVIEW OF THE LITERATURE

In the following sections the neurological and physiological psychology literature pertaining to the functional/anatomical organization of personality will be reviewed. The data will be related to psychological factors and any conclusions will be noted in discussions at the end of each section. The first section is a review of basic brain anatomy and organization. The second section will examine the physiological substrata of consciousness and conclude that human consciousness, characterized by self-awareness, is a manifestation of processes which occur in the left hemisphere of the brain. The third section will trace the systems involved in cortical activation and note the existence of two mechanisms within each hemisphere which have opposite effects on the form of cognitive processes. The fourth section will examine the role of the amygdala and frontal lobes in the subjective experience of emotion and of the right hemisphere in the expression of affect. The fifth section will develop the basis for a theory of memory function and the role of memory systems in organizing affective, arousal, and cognitive processes. The sixth section will focus on the interactions of the emotion, arousal, and memory systems and examine the role of biochemically mediated systems in coordinating these processes. In the seventh section a model of four functional systems which



7






8


constitute the basic elements of the personality structure will be proposed. The eighth section will describe the way in which the basic elements are organized into a functional meta-system which

forms the infrastructure of personality and functions to assure the emission of adaptive behavior. This model will be supported with

evidence concerning the psychological correlates of neurological syndromes. In the final section the psychological phenomena associated with various forms of psychopathology will be related to neurological indices which reflect the operation of the lateralized subsystems and their interaction within the functional meta-system. It Oill be concluded that many psychodiagnostic entities may be classified as hypo- or hyper-dominance spectrum disorders which are functionally related to chronic, and maladaptive, under- or over-utilization of the mechanisms in the left hemisphere of the brain.

Review of Basic Brain Anatomy and Organization

Cortical Mechanisms

Although certain phylogenetically new areas of the cerebral cortex are of special interest when discussing "higher mental functions," these areas must be considered in their physiological and evolutionary context. A brief review of basic brain systems and anatomy will establish this perspective.

In man, as in all mammals, large portions of the cerebral

cortex are devoted to the more elementary functions of processing sensory stimuli and the initiation and control of movements. The brain structures subserving these basic functions are divided at the






9


central fissure (Rolando). Incoming somato-sensory, visual and auditory nerve impulses are relayed, via the thalamus, from contralateral receptor surfaces to primary projection areas located in the parietal, occipital and temporal lobes (see Fig. 1, areas 1, 2, 3; 41; 17), respectively (Noback & Demerest, 1972). In each case the modally specific, somatotopic organization of nerve impulses in the projection areas is transformed into functional information (i.e., acquires meaning) in an adjacent secondary association area (areas 5, 7, 18, 19; 42). With each new level of processing there is increasingly complex synthesis of information and decreased modal specificity (Luria, 1973a). Damage to a primary projection area results in a loss of sensation (e.g., blindness) while lesions of a secondary association area are likely to produce the inability to recognize a stimulus in that modality (agnosia). Conversely,

artificial stimulation of a projection area produces a discrete sensory experience while stimulation of a secondary association area elicits a more elaborate sensory hallucination whose complexity is related to the level within the hierarchy that is activated (see Mullan & Penfield, 1959).

Progression within the hierarchy is reversed in the motor

systems. Specificity of control increases as the secondary (premotor) areas (areas 6 and 8) coordinate and fine tune their influence on the pyramidal cells of the primary motor cortex (area 4) with the assistance of continuous feedback from the sensory modalities. Lesions of the primary motor cortex produce contralateral paresis while stimulation elicits flexion of individual muscle groups.





10







/,- 64 7
8 65









187
9-219 19






818











Figure 1. Cytoarchitectural map of the lateral and medial surfaces of the human cerebral cortex, with numbers representing the areas of Brodman. (Redrawn from Noback & Demerest,
1972).
10 23 7 9

12 1

19




Figure 1. Cytoarchitectural map of the lateral and medial surfaces of the human 'Cerebral cortex, with numbers representing the areas of Brodman.' (Redrawrn from Noback & Demerest, 1972).









Stimulation of secondary motor areas produces smooth, coordinated movements and ablation of these areas may result in the loss of ability to perform skilled motor acts (apraxia) (Noback & Demerest, 1972).

The systems described thus far are typical of all mammals and become progressively more elaborate and efficient in the higher primates. Evolution proceeds generally by modifying and elaborating existing hardware, allocating new functions to tissues which are in some way pre-adapted to assume the new tasks (Campbell, 1974). The development of higher mental functions in man is correlated with bilateral anatomical expansion of two cortical areas which are adjacent to the secondary association areas described above. These regions subserve the highest level of organization in the hierarchies and are called teritary association areas (Luria, 1973a). Both areas are involved in what Penfield (1975) referred to as "transactions of the mind."

The inferior parietal lobule (IPL) (areas 23; 39, 40) lies at the anatomical confluence of the secondary association areas in the post-central cortex. This area is the "association cortex of association cortexes" (Geshwind, 1979). Here processed information from the surrounding sensory association areas is further integrated and synthesized. The area is called "supramodal" because its

individual units can only be excited by the simultaneous stimulation of two or more sensory modes (Luria, 1973a). Information processing at this level is "abstract" in that it is independent of a particular sensory modality (cf. Osgood, 1953). This ability allows the






12



simultaneous synthesis of information which permits the mental manipulation of the relationship between information units and as such is a prerequisite for the high level mental functions that are characteristic of human beings (Luria, 1973a).

At the anterior pole of the brain, contiguous with the secondary motor association areas, the prefrontal lobes (areas 9-12) are dramatically enlarged and now represent up to one-fourth of the entire cortical mass. The prefrontal lobes have extensive two-way connections with all other parts of the cerebral cortex (Luria, 1973a). The

coordinating and control operations carried out by the lower level (motor) systems in this hierarchy are evident in the functions of the tertiary integration areas. The frontal lobes have been called the "executive of the brain" (Pribram, 1973). They are the seat of Luria's "unit for programming, regulation and verification of activity" (Luria, 1.973a). Lesions of the frontal lobes lead to a defect in the patient's "capacity for planned initiative" (Penfield & Evans, 1940), and to disturbances on impulse control (Pribram, 1973).

Subcortical Systems

The limbic system consists of a group of interconnected structures, situated between the midbrain and the neocortical mantle, including the hypothalamus, amygdala, hippocanpus, septal area, and cingulate cortex (see Fin. 2). Because of their relationship to the olfactory bulbs, these structures were originally thought to be concerned with that function and so this area of the brain was designated the rhinecephalon ("nose-brain"). Although








13





























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ro J W LLU- M: C to L.) =.
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14



lower animals depend on the sense of smell for such survivalrelated activites as food-getting, the detection of enemies, and mating (MacLean, 1949), the role of these subcortical structures in motivational and emotional processes was not appreciated until the 1930s. Since Kluver and Bucy (1938) published their description of altered emotional behavior following the bilateral removal of the temporal lobes, including the amygdalae and hippocampi, limbic system components have been the subject of intense scrutiny. The massive literature which has accumulated is filled with confusing inconsistencies and conflicting findings. This may be due in part to the fact that the function of these structures appears to vary according to enviromental circumstances (Olds, 1958). Although a number of tentative models of limbic system function have been offered (e.g., Papez, 1937; Gloor, 1956; Olds, 1958) none has found general acceptance.

MacLean (1970) distinguishes three major types of systems in the mammalian brain which correspond to stages in its evolutionary development. He describes a protoreptilian core-brain, a paleomammalian brain (the limbic system) and a neomanmnalian brain (the neocortical areas). This triune brain conceptualization provides a useful perspective on the hierarchical arrangement of anatomical and functional systems and their relationship to behavior (Isaacson, 1974),
The protoreptilian brain represents the fundamental core of the nervous system consisting of areas homologous to parts of the upper brainstem and midbrain, hypothalamus, and basal ganglia.





15



In primitive organisms this brain produces a repertoire of instinctive,

stereotyped behaviors which are sufficient to insure the survival of the individual and species. These unlearned, species-specific

behavior patterns relate to the elementary functions of obtaining food and shelter, establishing and defending a home territory, breeding, maternal behavior, etc. (Isaacson, 1974). Programs for these behavior patterns are apparently stored in the brainstem and triggered by the hypothalamus: complex behavior sequences such as eating, drinking, sexual acts, aggression and many other types of unlearned behavior can be elicited by electrical stimulation of the hypothalamus at levels below those needed to activate motivational systems (Isaacson, 1974). In protoreptilian animals the performance of these acts would be coordinated by the basal ganglia. In these

forms the striatum is the highest center for sensory and motor processing, these functions being subserved by structures corresponding to the :audate/putamin and globus palladus, respectively (Schade' & Ford, 1973).

Stimulus and response are yoked in the protoreptilian brain. Since its repertoire of behaviors is unlearned (i.e., instinctive) this neural system "reacts to changes in the environment by increasing or decreasing the intensity of the predominant response sequence. . Suppression of response on the basis of non-reward or punishment is difficult" (Isaacson, 1974, p. 273).

The development of the paleomammalian brain allows the suppression of stereotyped ways of responding. The addition of the limbic

system circuitry permits the organism to adjust its behavior based






16



on new sources of information about the internal and external environments and to utilize new and more efficient forms of learning in organizing its responding. It is important to appreciate that the final product of limbic system operations in the paleomammalian brain is the inhibition of lower centers. However, these same mechanisms later came to exert important influences on the neocortical systems.

The cerebral cortex of the neomammalian brain developed in close association with the limbic system and basal ganglia. New nuclei were added to existing sensory and motor systems, culminating finally in the arrangement now found in primates. More efficient information processing in the neocortical additions permits more precise sensory discriminations and rapid, fine-grain movements of the extremities (Isaacson, 1974) but these new systems still operate in close conjunction with the older subcortical mechanisms (Schade' & Ford,

-1973).

The importance of the limbic system in human experience can hardly be overstated. These structures are implicated in most, if not all, forms of psychopathology. They are the probable substrate for the therapeutic effect of most psychotropic medications (Broekkamp & Lloyd, 1981) and the targets for all forms of "psychiatric surgery." Unfortunately, existing theory regarding the functional significance of the limbic system is primarily descriptive in nature or consists of generalizations so broad as to be of little use to the applied psychologist.





17



The Interpretation of Neurology Literature

The neurological data which form the basis of this review were gathered over many decades, from a variety of populations, by scientists with widely divergent theoretical beliefs. The psychologist who is unfamiliar with this literature must become aware of its inherent problems before venturing interpretations based upon it.

Those who come under the neurologist's care have, almost invariably, suffered a catastrophic change in their lives and personalities. Much of the data is drawn from individuals who have incurred brain damage. as a result of a head wound, cerebrovascular accident (stroke), or cerebral tumor (neoplasm). While offering an otherwise unavailable opportunity to study higher brain functions, these devastating "natural experiments" invariably preclude the rigorous application of scientific method to that study.

Although techniques continue to improve, it is unusually

impossible to define the parameters of a lesion accurately. Thus, crucial independent variables cannot be precisely isolated and controlled. Cause and effect interpretations are further hampered by the fact that a patient's pre-morbid level of functioning may be impossible to ascertain. Generalizability is also compromised when the subject has a chronic brain disease, such as epilepsy, or has undergone a surgical intervention to control that condition, such as temporal lobectomy or cerebral commissurotomy. In these cases, as with patients who have been subjected to "psychiatric surgery," it is reasonable to suggest that the "pre-treatment" psychological organization may have been grossly abnormal.





18



The human brain has remarkable ability to compensate for

injury by utilizing undamaged tissue. This recovery of function in the time elapsed between injury and testing is a further

confounding factor.

Much of this literature is in the form of case reports and the problem of individual variability is especially difficult to manage. Research designs utilizing groups and statistical analyses are becoming more frequent, but these studies invariably suffer from the difficulties noted above.

The most difficult problems to be encountered here have to do

with the specification of dependent variables: "higher brain functions" tend to resist definition and to defy quantification. Generally, only gross operational definitions have been available. The physician/experimentor was often forced to resort to a judgement as to whether a given (and hypothetical) function was "intact' or "lost," with the attendant risks of experimentor bias and error. Related to this is the understanding, which emerged slowly in the development of this literature, that the function of a circumscribed anatomical area cannot be inferred directly from a deficit which follows the ablation of that area: one can only specify how the brain functions in the absence of that tissue.

Human Consciousness
Right versus Left

There have been three basic approaches to the study of lateralized cortical function: the comparison of patients with unilateral brain damage; the lateralized presentation of stimuli or task to the






19


sensory receptor fields of normal subjects; and the study of patients whose cerebral hemispheres have been surgically separated (Nebes, 1977). The early studies of split-brain subjects prompted a great deal of speculation about the abilities of the right hemisphere and a listing of the published works concerned with asymmetrical hemispheric functioning is beyond the scope of this paper. Evidence pertaining to the differences between the cognitive operations performed by the left and right hemispheres has been ably reviewed by a number of writers (e.g., Bogan 1969a, 1969b; Galin, 1974, 1977; Gazzaniga, 1970; Gazzaniga & Ledoux, 1977; Nebes, 1974, 1977; Sperry, 1968). Some of the more important differences will be noted very briefly here.

The orimary deficits seen after right hemisphere lesions involve difficulties in perceiving, manipulating, and remembering spatial relationships and in perceiving and remembering sensory stimuli which resist verbal description (Nebes, 1977). The right hemipshere is superior to the left at generating a percept of the whole from fragmentary information and seems predisposed to notice complex gestalts or patterns rather than the parts (Nebes, 1971, 1974, 1977). The right hemisphere tends to process input simultaneously (in parallel) as opposed to the left hemisphere's preference for sequential (serial) processing, Cohen, 1973). In spite of many claims to the contrary, the evidence indicates that the right

hemisphere considers only the immediately perceived context and performs its tasks in a reflexive, automatic fashion.





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Language and the Left Hemisphe

The prominent Soviet geneticist, Theodosius Dobzhansky, observed that ". . while all other organisms become masters of their environments by changing their genes, man does so mostly by changing his culture, which he acquires by learning and transmits by teaching" (Dobzhansky, 1964, p. 145). Cultural evolution was made possible by the development of language, but language is more than a special form of social communication by which culture is transmitted: it is the mechanism which produces the adaptive behavior patterns that released the species from the constraints of biological evolution. Speech is the fundamental tool of intellectual activity. Language processes are essential in the operations of abstraction and generalization, the basis of categorical thinking, and the vehicle for organizing and regulating mental processes and behavior (Luria, 1973).

The clearest example of functional brain asymmetry is the lateralization of language to the left hemisphere, a fact long established by two observations: deficits in language functions (aphasia) following damage to the left hemisphere and the retention of language following right hemisphere injury (right handedness is assumed throughout the text unless otherwise noted), The patterns of language deficit following circumscribed lesions provide the most concrete evidence that is available about the mechanics of consciousness and its product, cognition. The focus of the following review is not on language itself, but on the evidence which indicates that the active operations of consciousness which produce






21


adaptive behavior are part of a functional system that is lateralized in the left hemisphere of the brain. Aphasia: Anatomy and Syndromes

Three general types of aphasia have been defined and traced

to different anatomical structures in the left hemisphere. Through an analysis of these syndromes it is possible to deduce the outline of a functional system in which these separate areas work together to produce the complex function of language. The following summary is based on reviews by Gardner (1975) and Zaidel (1978).

Broca's aphasia, resulting from damage to the inferior, postfrontal zones of the left hemisphere, is an expressive disorder. Utterances are difficult to initiate and speech is painfully slow and labored. Patients are usually able to name objects and to repeat, having less trouble with words that are familiar and concrete. Although they manage to convey their meanings in a peculiar, telegraphicc" style, they are unable to produce a fully formed sentence. Their productions consist almost entirely of substantives; grammatical parts and forms are absent or impoverished, Nouns are usually delivered in the singular and verbs appear in their simplest,

noninflected form. Parts of speech that are purely gramatical in function (conjunctions, prepositions, articles, adverbs) are exceedingly uncommon. The same deficit pattern is evident in the patient's reading and writing.

Although patients with Broca's aphasia may have difficulty in unravelling complex grammatical relationships (e.g., "The lion was eaten by the tiger: which animal is dead?"), their comorehen-






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of language is generally intact. The ability to understand and utilize nonverbal symbolism (e.g., gesture, pantomime) is also spared. Intellectual functioning is relatively unimpaired. Patients retain the ability to reason logically, to abstract and to generalize, and to respond to context appropriately. Their associational processes are not loosened, tangential or pressured. They do not produce paraphasias or confabulate. Finally, these patients are acutely aware of their deficits; they may be appropriately depressed and are prone to sudden, transient emotional outbursts.

Exactly the opposite clinical picture is evident when Wernicke's area, on the lateral, convex surfaces of the left temporal lobe, is damaged. Wernicke's aphasia is characterized by impaired language comprehension with fluently articulated but nonsensical speech. Unlike the Broca's aphasic, words are spoken clearly with normal sounding cadence, intonation and melbdy (prosody). However, the speech of the Wernicke's aphasic is lacking in content and may consist almost entirely of semantic jargon which has little conmunication value. These patients appear to have lost control of the language mechanisms at all levels. It is as if the selection thresholds for phonemes, words, and ideas were all lowered. Repetition is poor and marked by paraphasic errors in which the correct sounds may be present but emerge in the wrong order. Patients are able to name only the most familiar objects accurately, although the word produced may come from the same category as the target. Prompting with the initial wordsounds seldom helps.





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In spontaneous speech the key substantives are often missing and the remaining parts of speech lack their organizing influence. Grammatical parts and forms are used abundantly but incorrectly. Adjectives,, adverbs, conjunctions and prepositional phrases are strung together haphazardly and the result may resemble a schizophrenic's "word salad" (Luria, 1973). Nonsense syllables and neologisms are frequent. The guiding thought behind a verbal production may be evident, but it becomes obscured by tangential associations and incomprehensible babbling.

The Wernicke's aphasic understands little of what is said and seems to rely on nonverbal cues in order to respond to a situation. They are also unable to comprehend nonverbal symbolism and so their communication is vague and concrete at all levels.

Perhaps the most striking feature of Wernicke''s syndrome is

the patients.' complete lack of awareness of and indifference to their deficits. They appear unconcerned and will vehemently deny any problems.

The same deficits are evident in reading and writing tasks.

The patient can read single words but does not seem to grasp their meaning or relate them to him or herself. They may correctly repeat a simple command written on a card but will make no attempt to comply. Unable to formulate an acceptable sentence on their own, they are able to arrange cards with words printed on them to form a syntactically correct sentence. However, the key substantives are likely to be misplaced (e.g., "The man bit the dog.") confirming again that the patient's facility is with grammar as Opposed to meaning.





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There is less agreement about the third major language

disorder, known as anomic or amnesic aphasia, which results from more posterior lesions located in the angular gyrus in the parietooccipital area. Part of the confusion may stem from the fact that its main symptom, the loss of the ability to name objects, is common to all aphasic disorders. However, the anomic aphasia syndrome is distinguished by the fact that the naming disorder is accompanied by relatively intact comprehension of written and spoken language and normal spontaneous speech. The ability to read and to repeat are also spared in anomic aphasia.

The anomic aphasic has no difficulty using words in their appropriate context but cannot find the word in isolation of context; he is unable to divorce himself from the immediate situation. The anomic aphasic cannot produce the name of objects on demand even though he knows what they are. When an object is designated the patient is unable to produce its name, and conversely, given a name, the patient is not certain what it refers to.

Other language difficulties are evident. The patient's

spontaneous speech seems to be either too detailed or too general. Thinking is very concrete; the patient will interpret proverbs literally. The patient is aware of the diabilities and will often develop strategies to compensate for them. Lanugage, Symbolism, and Meanini

The theory of the functional organization of language developed by Wernicke in 1885 is still generally accepted today. According to this model the underlying structure of an utterance arises in





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Wernicke's area. It is then transferred via large fibre bundles (the arcuate fasciculus) to Broca's area where it evokes a detailed program for vocalization which is governed by the rules of grammar and syntax. The program developed in Broca's area, the linear scheme of the sentence, is supplied to the adjacent face area of the motor cortex which in turn drives the muscles which produce the vocalization. Thus, the content of speech originates in Wernicke's area and finds its form in Broca's area.

Wernicke's area is also essential for the comprehension of language. Auditory stimuli are relayed from the Organ of Corti to the primary auditory projection areas in Heschell's gyrus in the left temporal lobe. At this point, as with the other sensory projection areas, the information is somatotopically organized and retains its modal specificity; to be understood it must be transferred to the secondary auditory association area (Wernicke's area) where the somatotopical organization is converted into a functional organization (Luria, 1973a). Here, the fundamental phonemic characteristics of language are isolated and identified. Processing by Wernicke's area is essential for both the encoding and decoding of meaning. An intact Wernicke's area is also essential for the expression and comprehension of meaning through symbolic gesture and pantomime (Goodglass & Kaplan, 1973; Gionotti & Lemmo, 1976). It is interesting to note in this regard that "illusions of interpretation" emerge in consciousness after electrical stimulation of the temporal lobe in the right hemisphere but not after stimulation of any other brain area (Mullan & Penfield, 1959).





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Human Consciousness, Self-Awareness, and Thought

In the 1960s the general belief in cerebral dominance gave

way to the idea of cerebral specialization due, in large part, to the "split-brain" studies conducted on the cerebral commissurotomy patients of Drs. Vogel and Bogan. In 1969 Bogan took the progression a step further when he revived Wigan's (1844) notion of "the duality of mind." Wigan, (noting the anatomical duality of the brain, autopsy findings of hemispheric atrophy in patients whose personality was apparently intact, and introspective evidence of concurrent, opposing trains of thought) argued that if one hemisphere can sustain a mind, "it necessarily follows" that a man with two hemispheres must have two minds (p. 271). Bogan endorsed this concept in the third of his influential "Other side of the Brain" papers (Bogan. 1969a) and concluded that

Pending further evidence, I believe (with Wigan) that each of us has two minds in one person. . Various
kinds of evidence, especially from hemispherectomy,
have made it clear that one hemisphere is sufficient
to sustain a personality or mind. We may then
conclude that the individual with two intact hemispheres has the capacity for two distinct minds.
This conclusion finds its experimental proof in
the split-brain animals who can be trained to perceive, consider, and act independently. (1969,
pp. 156-157)
The "dual mind" concept implies two relatively equal but

functionally independent entities which act as opposed forces in the process of determining behavior. Bogan contributed the hypothesis that the two hemispheres utilize different "modes of thought" in this process: "propositional" on the left and "appositional" on the right (1969, p. 160). These appealing ideas were enthusiastically





27


embraced by laymen and professionals alike. They have been cited to support all manner of theories concerning psychological, philosophical, and spiritual dualities (see the critique by Kinsbourne, 1982). However, a closer analysis of the data cited by Bogan suggests that ushc conclusions are misleading.

Michael Gazzaniga, an author of the pioneering animal studies referred to by Bogan as perhaps the most dedicated and prolific of the "split-brain" researchers, complained about the "overpopularization" of basic data produced by himself and his colleagues:

These popular psychological interpretations of "mind left: and "mind right" are not only erronious: they
are inhibitory and blinding to the new students of
behavior who believe classic styles of mental activity
break down along simple hemispheric lines. (1977, p. 416)

There is no doubt that one cerebral hemisphere can, in the absence of its counterpart, support high level intellectual activity if the loss of the other hemisphere occurs early in development. Griffith and Davidson (1966), for example, report that children show relatively good recovery from hemispherectomy for infantile hemiplegia. Smith and Sugar (1975) reported on a 26 year old man who showed superior intelligence (WAIS VIQ:126, PIQ:102, FSIQ:116) 21 years after undergoing left hemispherectomy at age five and one-half. However, it is improper to infer normal functioning directly from grossly abnormal cases such as this, or from animal studies. While it is clear that the right hemisphere may have the capacity to develop higher mental processes, there is no evidence that it does so normally, and considerable evidence to the contrary. Research






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efforts in this area have been hampered by the lack of adequate operational definitions (for such phenomena as consciousness, cognition, and thought) which distinguish the hither mental processes of humans from the brain functions of lower forms.

Cerebral dominance for consciousness has been investigated

using the Wada carotid amobarbital test, a procedure developed to localize language functions prior to neurosurgery. The Wada test involves injecting sodium amobarbital into one common carotid artery and results in the anesthetization of only the cerebral hemisphere on the side of the injection. Terzian (1964) reported an absolute and immediate arrest of any communication, both verbal and nonverbal, in the first thirty to sixty seconds after the injection of the drug into the carotid artery of the dominant side which he interpreted as a transient loss of consciousness. Serafetinides and his co-workers reported similar results and noted that the phenomenon rarely occurred following barbiturization of the non-dominant hemisphere (Serafetinides, Driver & Hoare, 1964, 1965a, 1965b). They concluded that unconsciousness, and by

implication consciousness, is in general linked with the function of the hemisphere dominant for speech (Serafetinides et al., 1965a).

Rosadini and Rossi ('1967) attemDted to replicate these findings using more strictly operationalized definitions of consciousness. In one group (48 cases) the criteria consisted of an "analysis of the capacity of the patient to keep in contact with the examiner through verbalizations or movements, to react to noxious stimuli





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and to describe at the end of the examination what happened during the examination itself" (p. 103). In a second group the criteria

for consciousness consisted of "a simple stimulus-response test" in which "the patients were instructed to work a switch held in the hand ipsilateral to the intracarotid injection [i.e., the hand controlled by the unanesthetized hemisphere] any time they heard a given sound or saw a flash of light" (p. 103). Behavioral and clinical events indicating unconsciousness were required to last more than one minute in order to "Permit their safe detection." They found that 47 of their 69 cases did not meet their criteria for unconsciousness and the 22 cases which did occurred in roughly the same percentage following left and right injections; aphasia occurred in 16 cases (15 left, one right); only five of the 21 cases (three left, two right) evaluated with the stimulus-response test failed to operate the switch held in the hand controlled by the unanesthetized hemisphere in response to the signals used.

Although their results were complicated by existing neuropathological and cerebrovascular abnormalities, these authors concluded that "the existence of a cerebral dominance for consciousness is not supported" (,p. 111). The usefulness of this study appears to be severely limited by the criteria used to evaluate consciousness in the unanesthetized hemisphere: a large number of animals of various species have demonstrated the ability to "work a switch" in response to visual and auditory stimuli and to respond to noxious stimuli, but these lower forms are not considered conscious in the same sense that humans are. The authors were looking for "the






30



occurrence of signs revealing the capacity of the subject to keep in contact with the external world" (p. 103). They acknowledged that "the Suppression of expressive and receptive speech functions make such a task quite difficult with the patients receiving barbiturate in the dominant hemisphere" (p. 109). It seems that it is only the appearance of aphasiz (after dominant hemisphere anesthetization) that is specific to human consciousness in this study, and these

findings are consistent with those of Terzian and Serafetinides et al.

Rosadini and Rossi did not report the results of their test of subjects' ability to recall what had occurred during the Wada procedure but this question was addressed directly in an experiment reported by Gazzaniga (1977). This author found that "information encoded while the left hemisphere was anesthetized was uninterpretable by the verbal system when the left hemisphere returned to normal functioning .. when information is encoded by other than the verbal system the person is not consciously aware of the information" (p. 150).

Another approach to the localization of conscious awareness

involved an analysis of the temporal discrimination for simultaneity when two visual stimuli were presented separately to the left and

right visual half-fields, separated by a very brief interval. Efron (1963a; 1963b) found that normal right-handed subjects reported that the two flashes occurred simultaneously only when the light flashed in the left visual half-field was presented several milliseconds earlier than the light flashed in the right visual field.






31



Efron argued that this was because the "conscious comparison" of the two flashes takes place only in the hemisphere dominant for language, the time lag representing the extra neural steps involved

in relaying sensory information from the right hemisphere over the corpus callosum:

It is only after sensory data have reached the left hemisphere that one is "conscious" of the occurrence of an
event. . To be conscious of something is to be conscious
of something now. It is the thesis of this paper that
the "now" is the moment of arrival of sensory data in
the dominant temporal lobe. (1963b, p. 421)

The most convincing evidence of a correlation between human consciousness and language ability emerged from studies of a unique cerebral commissurotomy patient known as "case P.S." P.S., a right handed boy, developed epilepsy following an injury to his left hemisphere incurred at age two. He subsequently developed language skills in both his right and left hemispheres.

At age 14 he underwent complete surgical section of his corpus callosum to relieve his epilepsy. Following surgery it was found that P.S.'s right hemisphere could spell, comprehend verbal commands, process parts of speech and make conceptual judgements involving verbal information. It was also discovered that his right hemisphere, although unable to speak, could generate answers to printed questions presented tachistocopically to his left visual half-fields. He accomplished this by arranging Scrabble letters with his left

hand. These answers were often different from those given verbally by his isolated left hemisphere. Gazzaniga and Ledoux (1977) argued that P.S.'s right hemisphere possessed qualities deserving of conscious status because.






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His right hemisphere has a sense of self, for it knows the name it collectively shares with the left. It has
feelings, for it can describe its mood. It has a sense
of who it likes and what it likes, for it can name its
favorite people and its favorite hobby. The right
hemisphere in P.S. also has a sense of the future, for
it knows what day tomorrow is. Furthermore, it has
goals and aspirations for the future, for it can name
its occupational choice. . The fact that this mute half-brain could generate personal answers to ambiguous and subjective questions demonstrates that in P.S. the
right hemisphere has its own inde *pendent responsepriority determining mechanisms, which is to say, its
own volitional control system. (pp. 143-145)

P.S. is the only split-brain patient with advanced language skills in his right hemisphere and the only patient to demonstrate double consciousness. Ledoux, Wilson and Gazzaniga (1977) stressed the fact that "in all other patients, where linguistic sophistication is lacking in the right hemisphere, so too is the evidence for cons,,_-Iousness" (p. 420).

The capacity for speech and conceptual thought is clearly innate in homo sapiens; only the symbols themselves must be learned (Campbell, 1974). Recent evidence has clarified the anatomical substrate of this genetically transmitted specialization. The development of language capabilities in the human species is correlated with the anatomical expansion and interconnection of the association areas in the left hemisphere (Campbell, 1974). The posterior area of the planum temporale, .%hich forms a part of the secondary auditory cortex (Wernicke's area) is significantly larger on the left side (Geshwind & Levitsky, 1963. The enlargement of this area can be explained in terms of its distinctive cellular organization (Galaburda, LeMay, Kemper & Geshwind, 1978) and the





33


incomplete development of this cellular architecture has been related to language dysfunction (see Geshwind, 1979).

In her exhaustive study of hundreds of brain injured war veterans, Semmes (1967) discovered that elementary sensory and motor capacities were focally represented in the left hemisphere and diffusely represented in the right. She proposed that this difference indicates the mechanism of hemispheric specialization: focal organization favoring fine control and the integration of similar units (e.g., manual skills and speech) and diffuse organization favoring multimodal coordination (e.g., the various spatial abilities).

Gazzaniqa and Ledoux (1977) observed that nearly every demonstration of a right hemisphere advantage in split-brain patients has involved mani pul 0-spatial acti vi ties and concluded that

[iThis advantage] exists so long as manipulative activities
are involved in either the stimulus perceptions or the
response production . . The probable neural substrate of
these manipulo-spatial acts involves the inferior parietal
lobule of the right hemisphere in humans. In the left
hemisphere, however, linguistic functions occupy the
inferior parietal lobule .. . The superior performance
of the right hemisphere of split-brain patients on such
tasks does not reflect the evolutionary specialization
of the right hemisphere, but instead represents the
price paid by the left hemisphere in acquiring language,
Our view is not that the right hemisphere is
specialized in some unique way in man. Rather, it
continues to do what it does elsewhere in the phyla.
(pp. 420-421, emphasis added)

Campbell (1974) noted that "spatial relationships involving

depth and distance may appear to be predominately spatial concepts, but they are not of space but about space; of themselves they are spaceless and concerned with pattern rather than place" (p. 337).






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It appears, then, that the left hemisphere extracts meaning from the relationship of individual parts to each other, while the right gathers meaning from the pattern of the whole.

Bogan (1969b) proposed that the right "mind" utilized a

different mode of thought which he characterized as aDpositional to denote the ability to appose, or compare, information. He contrasted this with the propositional mode utilized by the left hemisphere. A number of investigators have distinguished similar dichotomies of information processing style. Luria (1973a) spoke of narrative versus relational processes. Galin (1974) suggested that the right hemisphere solved problems through a process Of Multiple converging determinants as opposed to a left hemispheric style which utilized a single causal chain. Sechenov (quoted by Luria, 1973a) postulated that the human brain utilizes two forms of integrative activity: organization into simultaneous and primarily Spatial groups, and into temporally organized successive series. This is consistent with Spearman's conclusion that intelligence comprises two components: the eduction of correlates used in analogical reasoning and the eduction of relations, the basis of abstract reasoning (see McFie & Piercy, 1952). Campbell (1974) noted that "abstraction means escape from the present . what distinguishes man from animals is the length of time through which his consciousness extends (p. 335). Finally, Bogan (1969b) observed that the most important distinction between the left and right hemispheric modes miqht be "the extent to which the linear concept of time participates in the ordering of thought" (p. 160).






35


There is a clear consensus recognizing two modes of information processing. However, the ability to process information is not necessarily a sufficient condition for consciousness. Although the notion of right brain "thought" has gained wide currency, to date, there has been no conclusive evidence that any cognitive operation occurring in the right hemisphere is directly experienced in consciousness. A possibility not considered by any of the above authors is that one mode might be an ancillary resource utilized by the other. The data reviewed thus far suggests that one must be

very careful to avoid anthropomorphizing when attempting to describe right brain processes. However, some inductive conclusions may be drawn,

Discussion

It is evident that human consciousness is inexorably linked

with the abstract symbolic processes associated with language. Perhaps the most universally accepted characteristic of human thought is self-awareness. Self-awareness (and its product, the self-concept) requires the abstraction and appreciation of defining features which are consistent over time and situation. Only the left hemisphere, with its temporal acuity, can consider and appreciate changed or conserved relationships in different conditions or contexts. Thus, only the left hemisphere can define itself. The resulting selfawareness provides a reference Point for all of the memories, feelings, intentions and thoughts that are collectively known as themind and which allow the individual, thus defined, to interact intelligently with the environment. Lacking the temporal organizing skills to






36


construct such a consistent frame of reference the right hemisphere is bound to the immediate context with only the influences of the physiological status of the organism (and the left hemisphere) to guide its processes. Complex motivations, therefore, cannot exist in the right hemisphere. Likewise, so-called "pictorial thinking," if temporally ordered and goal directed, must be organized by the left hemisphere. Galin (1974) suggested that the context bound, egocentric and impulsive nature of right hemisphere condition resembled Freud's notion of primary process thinking. Higher mental processes in the right hemisphere almost certainly qualify as cognitions (i.e., a way of knowing) and may account for the phenomenon of intuition (knowledge without awareness of the process by which it was gained). However, the term "thought" seems misleading and "information processing" might be preferable. As noted above, there is no direct evidence that mental events occurring in the right hemisphere are directly experienced in the conscious left hemisphere; one is left to ponder the question of whether a tree falling in the right brain would make a sound if the left wasn't listening.

The restrictions outlined above are in no way inconsistent with the demonstrated role of the human right hemisphere in the analysis of emotional communications and the modulation of affective expression (e.g., Ross & Mesulam, 1979). The brains of lower forms (and, apparently, the right brain in humans) are primarily concerned with neuronal signals which represent the survival needs of the organism within the immediate environment. Interaction with






37


the social environment is critically important to survival throughout the phylum. Campbell (1974) noted that animal vocalizations and signals are "emitted only in the presence of the appropriate stimulus" (p. 349) and warned against equating these vocalizations with human speech: "the signals . are generated or motivated by the phenomenon of emotion, and find their neurological origin not in the cortex but in the limbic system of the brain" (1974, p. 348). The cortical organization of these functions in the right brain of humans appears to mirror that of language in the left hemisphere with comprehension and expression utilizing anatomical areas homologous to Wernicke's area and Broca's area, respectively (Ross & Mesulam, 1979). Right hemisphere responses might achieve direct expression in circumstances where control by Broca's area in the left hemisphere is impaired or attenuated, a case in point being the clearly enunciated emotional exclamations of the frustrated Broca's aphasic. Similarly,

poorly defined and undifferentiated emotionally generated behavioral impulses (e.g., approach, avoidance) might also achieve motor expression in the absence of adequate left hemisphere control.

Hughlings Jackson (1864) suggested that if the "faculty of expression" was proven to be lateralized in the left cerebral hemisphere it would then be reasonable to expect that its corresponding opposite, perception, might be lateralized to the right. Although the concept of mental "faculties" has given way to an appreciation of complex functional systems, the role of the right hemisphere within those systems might, in a broad sense, be said to conform to Jackson's prediction. While the information processing





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style of the right hemisphere is not suited to solVing problems or making decisionsit is uniquely qualified to perceive the quality of significance (as defined by the individual's experience) in complex environmental stimuli. The evidence suggests that the human right hemisphere attends to the overall pattern of stimuli, searches out ("apposes") associations which are correlated with important stimulus configurations and collates them into percepts that have meaning for the organism. Its associational processes are unencumbered by rules of logic and its perceptions uninfluenced by expectations.

The thrust of Bogan's (1969b)"dual mind" thesis was a reaction against the traditional concept of hemispheric dominance which relegated the right hemisphere to the role of an "automaton" or reserve organ (e.g., Henschen, 1926; Strong & Elwyn, 1943). A basic assumption in the present work, however, is that evolutionary pressures required in the development of an automatic environment monitoring system in order to permit the transition from instinctual to self-determined behavior. It appears that evolution solved this problem by taking advantage of the fact that the human central nervous system contains two relatively autonomous brains which could be yoked together by the limbic system. Within this configuration the left hemisphere may be seen as a problem-solving and responsegenerating system and the right hemisphere might be said to

function as the repository and librarian of the individual's reinforcement history.





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Cortical Mobilization: Attention, ArousaT-, and Activation

Consciousness and cognition become possible only when minimum levels of cortical tone are attained. These tonic levels, reflected in the desynchronized EEG pattern, permit sensory discriminations, motor acts and other cognitive operations to take place. Once activated, the cortex has the ability to make phasic modifications of its level of activity and to voluntarily direct its attention. The fundamental systems which govern cortical tone and attention, however, are automatic and capable of overriding voluntary controls. It is clear that these systems, with their ability to control the level and content of consciousness, will exert a significant influence on the personality structure. The Reticular Activating System and Tonic Arousal

The mobilization of the cortex is accomplished by the brainstem reticular formation (RF). This structure is a network of highly interconnected neurons which adjusts its level of activation by integrating input from the sQnsory pathways, limbic system structures, and the neocortex. Impulses from the reticular formation lower the activation thresholds of the neurons it projects to. When the reticular formation is relaxed, cortical tone is lowered and the organism sleeps (Moruzzi & Magoun, 1949).

The reticular activating system (RAS) regulates the state of activation of the brain in two ways: the ascending reticular activating system (ARAS) affects the brain diffusely and sets the generalized (tonic) level of arousal; descending influences direct





40


RAS impulses to accomplish localized (phasic) arousal of specific areas of the brain. The ascending pathways of the RAS project rostrally from the brainstem reticular formation (via the central tegmental tract/medial forebrain bundle) to the hypothalamus, septal area, and nonspecific intralaminar thalamic nuclei (ILTN). The second path extends from the interpeduncular nucleus to the ILTN

via the habenula. The only direct ascending connections from the RAS to the neocortex are projections from the nonspecific thalamic nuclei (midline and ILTN) to the orbitofrontal cortex (via the ventral anterior nucleus of the thalamus). Descending influences are conveyed from the prefrontal neocortex to lower structures by way of the thalamocortical radiations, corticoreticular fibers,

medial forebrain bundle, and thalamotegmental fibers. Hippocampal output reaches the reticular formation via the fornix, mammillary bodies, and mammillotegmental tract. The septal area, has an additional connection with the reticular formation by way of a stria terminalis--habenula--habenulointerpeduncular tract--interpeduncular nucleus pathway (Noback & Demerest, 1972).

Based on their analysis of some 200 experiments, Pribram

and McGuinness (1975) outlined two major subsystems in the brainstem which control cortical mobilization and identified separate forebrain mechanisms which modulate their functioning. These systems initiate two different types of cortical activity. Diffuse

cortical "arousal," which is associated with the orienting response, is ased on the serotonergic brainstem median raphe" nuclei located in the core of the reticular formation. Arousal is modulated by






41


a lateral-frontal--amygdala--lateral-hypothalamus facilitory circuit and an inhibitory orbitofrontal--amygdala--medial-hypothalamus circuit. "Activation"is an attention focusing process involved in perceptual expectancies and motor readiness to respond. This system is based on the locus ceruleus, in the periaqueductal gray, which supplies norepinephrine to the forebrain. Activation was thought to be modulated by the ancient motor control system in the basal ganglia. Together, these systems provide for appropriate attending to novel or significant stimuli and prepare the organism to respond cognitively and behaviorally. The hippocampus was seen to integrate the functioning of these systems and to exert ultimate control over cortical mobilization through a mutually inhibitory relationship with the reticular formation.
Phasic Control Systems: The Frontal Lobes and Thalamus

A novel (possibly significant) stimulus elicits an orienting response (OR) from the organism. The psychological phenomena associated with the OR are familiar to all who have had experience with "things that go bump in the night." The complete orienting reaction includes
The suppression of ongoing behavior, the orienting of
the body and receptor towards the new stimulus, changes in the peripheral autonomic nervous system, and, perhaps
less obvious, preparations for associating the new stimulus
with memories from the past and expectancies of the
future. (Issacson, 1974, p. 110)
Pribram (1973) noted that the stimulus sampling aspects of the

orienting response differed from the processes necessary to register a stimulus in awareness and memory (which must be accomplished





42


before the organism can habituate to a stimulus). In contrast to the indiscriminate arousal associated with orienting, these latter processes require the focusing of attention.

The mobilization of selective attention ("activation") appears

to be reflected in the contingent negative variation (CNV) or expectancyy wave" (Tecce, 1970). The CNV is a special form of

cortical evoked response which consists of a spreading wave of negative potential that appears whenever there is a contingent relationship between two stimuli. Negativity develops when brain tissue is maintaining a readiness for processing (Pribram & McGuinness, 1975). Thus, the CNV appears whenever the organism is expecting to perform a perceptual or motor act. The negativity becomes abruptly positive when that act is executed (Walter, Cooper, Aldridge, McCallum, & Winter, 1964). High amplitude CNVs are related

to greater efficiency of perceptual and motor responses; concentration facilitates the CNV while inattention, boredom, or fatigue decrease it (Cohen, 1974). Elithorn et al. (1958) postulated that frontal lobe injuries somehow damaged the mechanism underlying anticipatory sets. It is interesting, in this light, that the CNV generally appears in the prefrontal lobes and sweeps posteriorally over the post-central cortex.

Patients with frontal lesions are unable to sustain their

attention. While intelligence, as measured by standardized tests, may be unimpaired, these individuals are highly distractable and cannot carry out purposeful activity which is normally directed by intentions (Luria, 1973 i). Luria pointed out that patients with





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lesions of the temporal, parietal, or occipital lobes may have

sensory, orientation, or intellectual deficits, but their attention and concentration remain sustained and directed by intentions. Luria and his co-workers suggested that the frontal syndrome reflected the loss of this selectivity (Luria, Homskaya, Blinkov, & Critchley, 1967). An understanding of the functioning of thalamic systems suggests a mechanism by which the frontal lobes might select or
"recruit" psychological operations in the post-central cortex.

The thalami are a pair of egg-shaped masses located beneath the cortex in the center of the cerebral hemispheres, The thalamus

is the final processing point for cortical input and the central integration station of the nervous system. The brief review of thalamic anatomy and functioning presented below is based on reviews by Noback and Demerest (1972) and Chusid (1976).

The ventral half of the thalamus contains the specific relay

of nuclei of the sensory-motor systems. The nuclei of the dorsal tier are association nuclei which have reciprocal connections with the association areas of the post-central cortex and no subcortical connections; the dorsolateral and posterolateral nuclei are interconneected with the parietal lobe, and the pulvinar with the temporal and Parietal lobe.

The dorsomedial and anterior thalamic nuclei are association nuclei involved in emotion and memory, respectively. The dorsomedial nucleus receives input from the amygdala and lateral hypothalamus and has reciprocal connections with the association areas of the prefrontal lobe. The anterior nuclei of the thalamus receive






44


the output from the hippocampus and have reciprocal connections with the cingulate cortex.

Lying between and separating the major thalamic association nuclei, the nonspecific (intralaminar, midline and reticulate) nuclei have only intrathalamic and subcortical connections. They receive their main input from the brainstem reticular formation and from the rostral end of the ascending reticular activating system (ARAS). The intralaminar nuclei have the "remarkable property of being able to exert a controlling influence upon the rhythmic electrical activity of the entire cortex" (Jasper, 1949, p. 406). Jasper noted that this system is in a position to provide a central

coordinating mechanism for cerebral activities:

A central integrative mechanism with ready access to all
afferent and elaborative systems of both hemispheres,
and closely related to autonomic spring of action; is necessary to explain consciously directed thought and
behavior. It seems that the thalamic reticular system,
with its diffuse cortical projections, relations to
afferent and efferent systems, relations to mesencephalic
hypothalamic and striatal systems, is a good candidate
for this office. (Jasper, 1949, p. 419) Discussion

The brain's mobilization systems with their brainstem, thalamic, and forebrain components, regul ate consciousness, unconsciousness, and the differential consciousness of attention. Sensory signals representing possibly significant stimuli cause the reticular formation to initiate diffuse cortical arousal. When a stimulus has been identified, cortical activation systems facilitate the organization of cerebral activity to deal with the situation appropriately. It appears that the frontal lobes direct this process





45



by recruiting psychological operations in the post-central cortex. The frontal lobe may accomplish this through its influence on the nonspecific thalamic nuclei which, in turn, control the phasic activation of specific cortical systems.

It is important to note that the two biochemically mediated subsystems which control the mobilization processes are duplicated in both halves of the brain. Unilateral prefrontal lobe lesions have been found to produce deficits which resemble those seen after damage to post-central lesions on the same side: right frontal lesions have been associated with disturbances of emotion and spatial abilities while left-sided injuries lead to disorders of speech and thought (Zangwill, 1966; Benton, 1968; Luria, 1973a;l. Processes which disrupt the biochemical balance between the two control mechanisms have far-reaching psychological consequences which will be reviewed in a later section.

i motivation: Emotion and Affect

The greatest risk involved in giving up instinct-based responding in favor of self-determined behavior is the possibility that the individual might fail to respond appropriately in survival-related situations. The evolution of the species could not have occurred if this problem had not been solved. The forces which motivate adaptive behavior must, by definition, be the single most powerful influence on the personality structure. The evidence indicates that the source of these forces lies in the limbic system. It appears that separate cortical mechanisms mediate their internal experience and external expression.






46


Amygdala Circuits and the Prefrontal Lobes

The role of the amygdala in emotional processes established by Kluver and Bucy in 1938, has been assumed to be affected through this structure's close relationship with the hypothalamus. The amygdala seems to direct behavior toward biological goals (Halgren, 1981) and is implicated in the control of species-sDecific behaviors related to survival needs, including defensive and aggressive behaviors, sexual activity, and feeding (Isaacson, 1974). In lower forms these processes might depend on a simplified (instinctive) form of memory in which stimulus and response are yoked (Pribram & McGuiness, 1975). In addition to mediating emotional states the amygdala is involved in the analysis of reinforcement contingencies. Amygdala lesions have been shown to produce impaired recognition of stimuli associated with rewards (Weiskrantz, 1956; Schwartzbaum, Thompson & Kellicut, 1965; Jones & Mishkin, 1972) and inability to respond appropriately to changes in the magnitude of rewards (Schwartzbaum, 1960).
Strong interconnections with the hypothalamus (via the stria terminalis and ventral amygdalofugal fibers) give the amygdala immediate access to information concerning the internal status of the organism (Price, 1981). The amygdala also receives processed sensory information from all of the secondary sensory association areas (Van Hoesen, 1981). Mishkin and Aggleton (1981) noted that this arrangement places the amygdalae in a position to integrate external events with their internal consequences, which would permit the attachment of emotional and motivational significance





47


to sensory stimuli. Kessner (1981) reported experimental evidence that demonstrated the essential role of the amygdala in encoding and retrieving the positive and negative attributes of a specific memory. In lower forms the identification of a motivationally significant stimulus might result in the release of species-specific behaviors, but in humans behavior is self-determined. Halgren (.1981) concluded from his amygdala stimultion studies with humans that "the amygdala helps organize the discharge of emotional tension into consciousness" (p. 404) and noted that this would allow the directing of consciousness toward biological goals. The amygdala's input to the neocortex is directed to the entire prefrontal lobe both directly, via the uncinate fasciculus, and indirectly, by way of the dorosomedial thalamus (Noback & Demerest, 1972; Price, 1981).
While damage to the dorsolateral area of the prefrontal lobes has been associated with intellectual disturbances, lesions of the orbito-frontal cortex (and orbital undercutting, which disconnects this area from the amygdala) result in emotional changes (Lewin, 1961). Elithorn, et al. (1958) concluded that this type of damage produced a "generalized impairment of the ability to form appropriate emotional responses" (p. 250),,including the ability to elaborate on the affect appropriate to the concepts present in consciousness.

In contrast to the planning deficits, loss of energy and interest, and affective dullness seen after dorsolateral frontal damage,.orbitofrontal injuries often lead to euphoria, impulsive (disinhibited) behavior, and the appearance of "greediness, selfishness, and






48


tactlessness" (see Lewin, 1961). Faust (1966) noted that such patients resemble psychopaths in that they are unable to profit from experience and are in constant conflict with their environment and the law. Zangwill (1966) pointed out that the tactlessness

common in frontal lobe patients does not result from a loss of knowledge of social conventions, but from the failure to regulate behavior in accordance with those standards.

Disconnecting the orbito-frontal cortex from the am~ygdala

(orbital undercutting) has been reported to be the most effective psychosurgical operation for relieving the symptoms of anxiety and depression (Elithorn et al., 1958; Lewin, 1961; Levinson & Meyer, 1965). Elithorn et al. (1958) noted that this procedure

increased reactions of "a hysterical type" and is contraindicated for those conditions. It is interesting to note that drugs that reduce anxiety and produce euphoria (e.g., the barbiturates) have been found to exert an uncoupling effect between the frontal lobes and the limbic system (Heath & Galbraith, 1965). Affective Express ion

The right hemisphere plays an essential role in both the

comprehension of emotional communications and the expression of affect. Patients with right hemisphere lesions showed impaired recall of stories with emotional content versus neutral stories (Wechsler, 1973). Hielman, Scholes and Watson (1975) demonstrated that judgements of the emotional mood of a speaker (sad, happy, angry, indifferent) made by patients with right temporoparietal

lesions were significantly impaired relative to patients with leftsided lesions and controls. This finding was replicated by Tucker,






49


Watson and Hielman (1976) who showed also that right hemisphere patients were impaired in the vocal expression of emotion. The efforts of patients with right temporoparietal damage to impart a sad, happy or angry tone to their voices were rated as incorrect significantly more often than controls. Ross and Mesulam (1979) presented case studies of two well-educated patients who had comnparable damage in the right supra-sylvian area, which is homologous to Broca's area on the left side. Both patients showed flattened affect and had completely lost the ability to laugh, cry, or otherwise express any emotion in their speech. Their ability to experience and comprehend emotions was unchanged. The authors noted that the organization of emotion in the right hemisphere seems to mirror that of language in the left: the area homologous to Wernicke's area being essential for comprehension, and to Broca's

area, for expression.

Gazzaniga and Ledoux (1977) suggested that right hemisphere functioning in humans is distinguished only by contrast to the left; it continues to perform its functions in the same manner as elsewhere in the phylum. Although it should be noted that the human right hemisphere is in possession of tertiary association areas and so would perform those tasks more efficiently, the data reviewed above are not inconsistent with Gazzaniga's interpretation. The brains of lower forms (and, apparently, the right brain in humans) are primarily concerned with neuronal signal !s which represent the survival needs of the organism within the immediate environment. Interaction with the social environment





50


is critically important to survival throughout the phylum. Campbell (1974) noted that animal vocalizations and signals are "emitted only in the presence of the appropriate stimulus" (p. 349) and warned against equating these vocalizations with human speech: "the signals . are generated or motivated by the phenomenon of emotion, and find their neurological origin not in the cortex but in the limbic system of the brain" (1974, p. 348). Discussion

It appears that basic human emotional experience is an

emergent property of the functioning of mechanisms that originally served to regulate the emission of soecies-specific behaviors which were elicited directly by releasing stimuli in survivalrelated situations. The functional systei which evolved in humans decouples stimulus and response. T1us, in humans, it is t~le emtional experience evoked by a stimulus--rather than the stimulus itself-that is the primary motivating factor (reinforcer) which ultimately determines behavior. Further, this emotional experience might be most properly considered to be a part of the experiencing person's environment, since that experience is involuntary and has the power to condition the person's response. These processes appear to have their functional impact in the left hemisphere, where the formulation of behavioral responses occurs.

The physiological mechanisms which motivate adaptive behavior in humans are centered on the amygdala which integrates information from the internal and external enviroments in order to attach emotional significance to stimuli. This structure is involved in the





51



encoding and retrieval of this information in memory and forwards its signals to the prefrontal lobes where they are experienced as subjective emotions. The prefrontal lobes appear to utilize these signals in the process of forming intentions to direct adaptive behavior. This amygdala-prefrontal pathway appears to be the substrate of anxiety and depression. When the amygdala-frontal connection is severed surgically, or uncoupled pharmacologically, the neocortex experiences euphoria, but fails to behave in an adaptive manner.

Memory Functions
It is evident that the functional brain systems which form the infrastructure of personality include separate cognitive, affective and arousal components. It appears that these subsystems evolved to take full advantage of a form of learning which utilizes reinforcement and emotional experience in determining behavior. The product of any learning experience is memory. The importance of memories (or "associations") in the organization of cognitive operations is obvious and most, if not all, arousal and affective processes must depend on the ability to discriminate personally relevant stimuli. Clearly, memory is fundamental to all aspects of personality function, but the material substrate of memory remains a complete mystery (e.g., Lashley, 1950; Luria, 1975a);-our concepts regarding it are, of necessity, only abstract descriptions, Before reviewing the physiological organization of memory systems it will be necessary to define and delimit, as far as possible, those abstract





52



concepts of memory phenomena that are pertinent to the interests of the applied psychologist.

Experimental psychologists have traditionally approached,

the study of memory by subdividing it into registration, retention, and recall, attempting to isolate and measure these aspects and the variables which affect them. Rapaport (1961) criticized this methodology as artificial, insisting that these functions are inextricably related and that such experiments merely demonstrate how memory can function under given laboratory conditions, Working from a psychoanalytic perspective, Rapaport preferred to treat memory as an aspect of cognition. He argued that "actual memory phenomena are encountered only in the context of thought processes; at best the classical memory experiments could ignore this fact and make us ignore it, but they could not produce memory phenomena outside this context" (Rapaport, 1961, p. 6). He acknowledged the difficulties in determining the relation of indistinct entities such as emotion and memory and attempted to clarify the psychoanalytic viewpoint by suggesting that "memory is a motivated behavior phenomenon and emotions are motivating factors" (Rapaport,

1961, p. 8). This statement, however, appears to beg the question; if memories are activated by emotions, then what initiates arousal and affective processes?

The interaction of cognition, affect and memory in the etiology and cure of psychopathology were central themes in the work, published in 1893 by Josef Breuer and Sigmund Freud, which gave Psychoanalysis its start. The emphasis on the significance of






53


memory phenomena in psychoanalytic literature (e.g., slips of the

tongue, forgetting, false remembering, repression) can be traced

to this seminal paper in which the authors concluded that "hysterics

suffer mainly from reminiscences." In this work, Breuer and Freud

made a crucial distinction regarding the memory processes operating

in psychoneurosis which may have sowed the seed from which the

notion of unconscious causation of psychological phenomena germinated:

...the causal relation between the determining psychical
trauma tan experience which calls up distressing affects
such as those of fright, anxiety, shame or physical pain]
and the hysterical phenomenon is not of a kind implying
that the trauma merely acts like an agent provocateur
in releasing the symptom, which thereafte leads an
independent existence. We must presume rather that the
psychical trauma--or more precisely the memory of the trauma--acts like a foreign body which long after its
entry must continue to be regarded as an agent that
is still at work.' (Breuer & Freud, 1974, p. 355)

The authors became aware of this "highly remarkable phenomenon"

and its relation to affective processes in the course of their

experimental treatment of hysterical conversion symptoms:

[we found] that each individual hysterical symptom immediately and permanently disappeared when we had
succeeded in bringing clearly to light the memory of
the event by which it was provoked and in arousing its accompanying affect, and when the patient had
described that event in the greatest possible detail
and had put the affect into words. Recollection
without affect almost invariably produces no result.
(Breuer & Freud, 1974, p. 355)

Hillix and Marx (1974) have suggested that "it was necessary for

Freud to invent the psychic apparatus and much of his psychoanalytic theory just to account for what he and Breuer had already

observed" (p. 352). It is to be hoped that recent evidence will

make a more parsimonious accounting possible.





54



The implicitly verbal form of memory referred to by Rapaport seems to be qualitatively different from the "foreign body" which Breuer and Freud assumed to be the culprit in hysterical neurosis. They referred to the latter type of memory by the less formal term "idea" and indicated that symptom removal depends.on the transformation of this "idea" into a more formal thought process so that its associated affect can be abreacted:

[the therapeutic procedure] brings to an end the operative force of the idea which was not abreacted in the
first instance, by allowing its strangulated affect to
find a way out through speech; and it subjects it to associative correction by introducing it into normal
consciousness. (p. 356).

This special type of memory would seem to merit a more detailed description. It is evident that we are concerned here with a subset of memories which have significance for the individual. By definition, these are memories that are associated with reinforcement and/or emotional experience. They are experiential (nonverbal) and may be isolated from consciousness. It may be noted that this subset of memories will define the relationship between the individual and his or her environment and might be the organism's most important survival resource. A concept from social learning theory seems to encompass this type of memory comfortably and provides a more

operational definition.

Julian Rotter (1966) theorized that "a reinforcement acts to

strengthen an expectancy that a particular behavior will be followed by that reinforcement in the future" (p. 2). Further, "when an organism perceives two situations as similar, then his expectations





55

for a particular kind of reinforcement, or class of reinforcements, will generalize from one situation to another" (Rotter, 1975, p. 57). Rotter distinguished two types of "generalized expectancies" (GE). The first has to do with the nature of the reinforcement: expectations for a particular kind of reinforcement in a given situation. The second type deals with other properties of situational stimuli and has to do with the perception of control that one can exercise to change or maintain the situation: the kind of behavior that is likely to produce or terminate reinforcement. The first type is designated with a subscript r for reinforcement (GE r ). The second type is designated a problem-solving generalized expectancy (GEPS). Striking insights into the nature and mechanics of this sort of experiential memory were afforded by Penfield's observations of certain psychical phenomena elicited by direct electrical stimulation of the conscious brain (see Mullan & Penfield, 1959).

Wilder Penfield's data were collected from patients undergoing

radical brain surgery, with local anesthesia, for the relief of intrac-, table epilepsy. His observations consist of spontaneous reports from these patients following applications of a mild electric current to the exposed cortex from the tip of a unipolar electrode. The responses to such stimulation which are of interest here fall into three categories:

1. The emergence in consciousness of vivid and coherent

experiential hallucinations which appeared to be recollections of (or abstractions from) the subject's past experiences.

2. Changes in a patient's subjective experience of his or her relationship with the immediate environment.






56


3. "Illusory" emotional experiences.

Based on his analysis of the data, Penfield (1975) postulated the existence of two related brain systems: a "mechanism of recall," and a "mechanism of interpretation." The latter involved the temporal cortex (exclusive of the speech areas) and was referred to by Penfield as the "nonverbal concept mechanism." Penfield comDared its function with nonverbal concepts to the operation of the speech cortex with verbal concepts.

Somehow [this mechanism] seems to analyze the components
of sensation, compares them with previous experience,
and by that analysis and comparison, transmits into
consciousness their present and immediate significance
: I [an emotional response] is a signal that rises
into consciousness as a result of an interpretation of
what the present situation may bring the subject in
the immediate future.. (Mullan & Penfield, 1959, p. 283) It appears that the cognitive products of these mechanisms fit the criteria for generalized expectations. Penfield's evidence assists in understanding the different types of memory referred to by Rapaport and by Breuer and Freud and indicates the neural substrate of these processes. (Penfield's data and conclusions will be reviewed and evaluated in detail presently.)

It has been assumed here that at the base of personality there are mechanisms whereby cognitive operations and affective processes are appropriately activated by significant memories. The present task is to describe neurological evidence which accounts for the memory phenomena reviewed above in terms that fit the criteria for "functional systems" as defined by Luria, and satisfy the evolutionary imperatives outlined at the beginning of this chapter.





57


The neuropathological correlates of human amnesia syndromes and data from related animal studies will be reviewed. It will be hypothesized that human beings possess two autonomous, lateralized memory systems, centered on the hippocampi, each of which is intimately associated with its own emotional and cortical activating system. Human Amnesia Syndromes

A number of terms are used to describe different aspects of

memory function and dysfunction. Immediate memory, usually measured

by digit span, probably reflects the ability to hold information in the primary or secondary sensory cortex as long as voluntary attention (directed by the nonspecific thalamic nuclei) is focused upon it (Smithies, 1966). The terms short-term ("recent") and long-term ("remote") memory indicate recollection over increasingly greater periods of time, but both are almost certainly subserved by the same physiological processes (Brierley, 1977). The term retrograde amnesia refers to a period of time before an accident or illness for which the patient's ability to recall is diminished or lott Ant rde amnesia is an inability to retain in memory events that occur after such an injury or illness.

The combination of a severe retrograde amnesia and a debilitating anterograde amnesia is the hallmark of the Wernicke-Kors~koff syndrome, the most common form of memory disease. This illness is most fr ,quently seen as a result of brain lesions brought on by dietary (thiamine) deficiencies in chronic alcoholics, although the lesions and illness may be produced by a number of toxic or disease processes (see the excellent review by Brierley, 1977). This illness






58


was described independently in the 1880s by Karl Wernicke (whose work with aphasia was reviewed earlier) and S. S. Korsakoff, a Russian psychiatrist. Wernicke focused on the acute stage of the illness during which the patient is usually depressed, fearful and anxious; often paranoid; and always severely confused and disoriented (Wernicke's encephalopathy). Patients who survive this acute stage become stabilized in the phase known as Korsakoff's psychosis. This chronic state is characterized by severe memory disorders and profound changes in the patients personality, motivation, and affect.

Against a background of retained intellectual skills and intact remote memory, the victim of Korsakoff's psychosis suffers a retrograde amnesia for periods of up to several years before the onset of the illness, and an almost total inability to recall any new information once his or her attention is distracted from it. Consequently, these patients live virtually in the immediate present and are always disoriented as to time, place and situation. These patients are, at best, only vaguely aware of their inability to learn new material. They often produce confabulations to cover gaps in

their recollection and fuse or combine ("reduplicate") experiences from different periods in their lives (Gardner, 1975). In both cases they believe that their statements reflect reality. In most cases the patient's affect is blunted, although some patients have exhibited chronic euphoria (e.g., Remy, 1942). They show reduced spontaneity and initiative and a "lack of desire for alcohol, sex, and other traditional reinforcers"'(Gardner, 1975, p.'188).






59


(Such indifference to alcohol is especially interesting in light of the fact that the brain damage in most of these patients was caused by years of sustained, heavy drinking.)

Post-mortem examination of the brains of persons who suffered

from Wernicke-Korsakoff disease reveals lesions of certain anatomical structures in the limbic system associated with the well-known circuit of Papez. Such damage almost invariably includes, and may be confined to, injury to the mammillary bodies, the relay for hippocampal output on its way to the anterior thalamic nuclei

(Brierly, 1977).

A pure form of this memory disorder was the unfortunate consequence of bilateral removal of the hippocampi in humans. In the 1950s Scoville performed a series of experimental operations designed to relieve the symptoms of chronic schizophrenia without the undesirable side-effects of a complete frontal lobotomy. The surgical procedure involved the resection of the medial surface of the temporal lobes from 5.0 to 8.0 cm posterior to the tip of the lobe combined in some cases with orbital undercutting. Thirty severely deteriorated schizophrenics had undergone the operation, with slight improvement in their conditions, when a purely temporal resection was performed on a nonpsychotic epileptic patient whose seizures were unresponsive to medication. When this patient, "case H. M.," recovered from the operation it became apparent that he had developed a severe amnestic disorder which resembled Korsakoff's psychosis and which persisted at 14 years (Milner, Corkin & Teuber, 1969). Scoville

and Milner subsequently examined eight of the psychotic patients






60


who had undergone the operation and who were able to participate in formal testing. They discovered "some generalized memory disturbance in all patients with removals extending far enough posteriorly to damage portions of the hippocampus and hippocampal gyrus (Milner, 1958, p. 112). The degree of memory impairment was more or less proportional to the amount of these structures removed. Bilateral resection of the uncus and amygdaloid nucleus alone did not result in amnesia (Brierly, 1977), nor did removal of the gyri of the outer aspects of the temporal lobes (Bailey, 1946). It has been concluded, therefore, that "the structures necessary for normal memorizing are the hippocampal formations within the temporal lobes, the mammillary bodies and, possibly, certain thalamic nuclei within the diencephalon" (Brierley, 1977, p. 221); that is, the hippocampi,. their output pathways and related projection sites.
There has been, as yet, no definitive explanation of either the nature of the amnesic deficit described above or of its underlying mechanisms. The fact that remote memory seems to be intact in these patients has led many to believe the the hippocampaldiencephalic structures are not involved in the process of recall, although Brierley (1977) pointed out that such a conclusion is unjustified in the absence of adequate pre-post evaluation of this function. Milner (1966, ch. 5) attempted to account for the pairing of a period of retrograde amnesia with an inability to learn new material by hypothesizing that the establishment of a permanent memory trace requires an extended period of "consolidation," which is somehow disrupted in this syndrome. In Milner's view the deficit






61


represents a failure to transfer sensory impressions into long-term store. The adequacy of the consolidation hypothesis is called into question, however, by demonstrations that amnesic patients are in fact able to recall new information under certain conditions. The most convincing evidence comes from experiments using the technique of "cued recall" in which a subject is given partial information about a stimulus (e.g., a previously presented word or picture) and asked to identify the whole item. Under these circumstances the performance of amnesic subjects was not significantly different from that of controls (Weiskrantz & Warrington, 1970). This suggested to the authors that the amnesic deficit involved problems with mechanisms of retrieval rather than those of acquisition or retention.

Weiskrantz (1979) reviewed a number of experimental paradigms in which normal learning has been demonstrated in amnesic subjects and underscored the fact that, in each case, the patients themselves persistently failed to acknowledge the fact that their performance was based on specific past experience or that they had been confronted with the task before). Thus, amnesia vi,.ctims do not have access to their memories on a conscious level, nor is such awareness necessary for that memory to be demonstrated objectively. Weiskrantz pointed out that this- "striking dissociation between the subjects' commentaries and their objective performance . suggests a dissociation between levels of processing rather than a failure on any particu-lar level" (p. 385).





62


Levels of processing in memory are the subject of a theory

(summarized by Gaffen, 1972) which is based on arguments by Talland (1965) and supported by experimental evidence (Peterson, 1967; Kintsch, 1970). Briefly, the theory postulates that the process of recall consists of two separate and autonomous stages: retrieval (or search) and recognition. The retrieval process "proceeds at several levels . each being terminated by an implicit act of recognition" (Talland, 1965, p. 304). The recognition stage is based on a record from which the past cannot be read directly, but which can assign a particular response in a particular context a value of "familiarity--unfamiliarity" (the correct response being the most familiar in that context). Thus "[in the retrieval stage] various responses are generated (but not emitted); when finally the correct response is generated, it is recognized as such by the recognition stage, and is then emitted" (Gaffen, 1972, p. 328). The theory postulates that amnesic subjects (animal and human) lack the faculty of discriminating familiarity. This basic deficit is manifested in the premature termination of search cycles, resulting in an incorrect match. These formulations are not inconsistent with those of Butters and Cermack (1974), who concluded from their experiments that increased sensitivity to proactive interference, subsequent to inappropriate encoding of information, was the critical factor- underlying the amnesic disorder. Finally, it is interesting to note that modern theories regarding amnesia seem to have arrived at the point at which they began: Korsakoff (1889), in keeping with the associationist doctrine of his time, believed





63


that his patients were deficient in making associations among new ideas and in-connecting past and present experience.

The suggestion of bilevel processes in memory noted above are particularly interesting given that language and conscious awareness have been associated with temporal lobe structures of the left hemisphere, and Penfield's (1975) report that electrically induced "illusions of recognition" were elicited only by stimulation of temporal structures of the right hemisphere. Memory and the Neocortex

As noted earlier, Penfield's experiments with electrical stimulation of the cortex in conscious patients led him to postulate the existence of two separate memory systems: a "mechanism of recall" and a "mechanism of interpretation." The existence of the former was suggested by the fact that, following stimulations of the exposed cortex, some of his patients reported vivid auditory and/or visual experiences ("flashbacks"); it seemed to the patients as if they were reliving prior experiences in their lives, although they retained their awareness of the operating room environment. Since many of these sensory sequences were trivial, yet perceived as familiar, Penfield concluded that his electrode was tapping a "continuous record of conscious experience." Although widespread areas of the cortex were exposed and explored, these "experiential responses come only from the temporal lobe, never from any other part of the brain" (Penfield, 1975, p. 31).

Niesser (1967) presented persuasive arguments refuting Penfield's claim that "nothing is lost . the brain of every man contains an





64


unchanging ganglionic record of subjective experience" (Penfield, 1954, p. 67). Niesser suggested that "most of the cases described by Penfield seem more like generic and repeated categories of events rather than specific instances" (p. 168). (It will be noted that Niesser's formulation is congruent with the notion of a "generalized expectation," as defined earlier.)

Penfield's second mechanism was suggested by another category of electrically induced phenomena which consisted of the "misrepresentation or altered interpretation of present experience" (M611an & Penfield, 1959, p. 269). Prominent among these were "illusions of recognition" during which "present experience seemed familiar, strange, altered, or unreal" (p. 270).' These "illusions of comparative interpretation" were associated with stimulation of the temporal cortex in the hemisphere that was minor for handedness and speech. The authors believed that "in normal life, these are signals that rise into consciousness, signals tnat depend on subconscious comparison of past experience with the present" (p. 283).

In 1951 Penfield proposed that portions of the temporal lobes be called "memory cortex" in the belief that his electrode had activated a neuronal record which was stored there. He was obliged to revise this theory in 1958 because of a new understanding of the physiology of electrical brain stimulation. When an electrode passes a current into the cerebral cortex, the current completely disrupts the patient's normal use of that gray matter (e.g., stimulation of the speech areas produces momentary aphasia). Therefore, any positive responses are produced by axon-conduction and the functional





65


activation of a distant, secondary ganglionic station (Penfield, 1975, ch. 7). In his later formulations then, Penfield referred to those temporal structures as the "interpretive cortex" and postulated that his electrode had activated a final common pathway to a secondary center which in turn produced the illusions of comparative interpretation. Since the temporal lobe forms the principal source of input into the hippocampus (which was known to be related to memory), Penfield assumed that this was the secondary center in question. He suggested that

The hippocampi seem to store keys-of-access to the
record of the stream of consciousness. With the
interpretive cortex, they make possible the scanning
and the recall of experiential memory.* (Penfield,
1975, p. 36)

Penfield's finding that illusions of familiarity were associated with activity of the temporal lobe in the right hemisphere is complemented by Kimura's (1963) evidence that the right temporal lobe appears to be more involved in the analysis of unfamiliar stimuli. Kimura presented familiar and unfamiliar visual stimuli to the right and left visual fields of patients with lesions of the right or the left temporal lobe. The right temporal group was impaired in the perception of the unfamiliar stimuli but not the familiar. Kimura interpreted her results in terms of the verbal identifiability of a stimulus:

It seems clear that a frequent (though not a necessary) concomitant of familiarity in a perceptual sense is the
possibility of verbal identification. Where increased
familiarity with a stimulus object, or class of objects,
is associated with the repeated naming of the object,
the ability instantly to attach a name to it represents an important step in the development of a concept. It






66



seems probable that in such cases a large part of the increase in permanent neural -. epresentation which is assumed to correlate with familiarity will take place
in the language centers, that is, in the dominant
hemisphere. (Kimura, 1963, p. 269)

Thus, unfamiliar stimuli, which are not represented in verbal memory by a permanent neural model (e.g., a name or concept) are more likely to be processed by the right hemisphere, which Kimura suggests is more important, than the left in the establishment of such "cell assemblies." Since all of the material in the memory store are initially unfamiliar it follows that many, if not all, verbal concepts (in the left hemisphere) might be based on neural models, or gestalto7n, which were assembled (and are stored) in the right hemisphere. Such an hypothesis is supported by evidence from split-brain studies that the right hemisphere is far superior to the left in the discrimination of part-whole relationships (Nebes, 1974).

The appearance of material-specific amnesia syndromes following unilateral temporal lobectomies suggests that the isolation of language in the left hemisphere extends to verbal learning also, and is thus virtually complete. Milner (1971) reports that,

A comparison of left and right anterior temporal lobectomy in epileptic patients has revealed certain specific
memory defects that vary with the side of the lesion.
These material-specific disorders are to be distinguished
from the global amnesia that follows bilateral damage
in the hippocampal zone (Milner, 1958). Thus, left
temporal lobectomy, in the dominant hemisphere for
speech, selectively impairs the learning and retention of verbal material (Meyer & Yates, 1955; Milner, 1958).
...Conversely, removal of the right, nondominant
temporal lobe leaves verbal memory intact but impairs
the recognition and recall of visual and auditory patterns






67


that do not lend themselves easily to verbal encoding.
...Thus within the sphere of learning and memory
there is a double dissociation between the effects of
these two lesions.* (Milner, 1971, p. 274)

Butters and Cermack (1974) focused on the specifically verbal aspects of the amnesic disorder in Korsakoff patients and noted that, during learning tasks, these patients did not react to changes in semantic categories on successive lists. They concluded that the amnesic deficit was due to the patient's

inability to encode verbal information along semantic
or meaning dimensions. . Korsakoff patients do not
spontaneously employ semantic encoding strategies,
but rely on basic acoustic and associative categorizations.
If the Korsakoff patient is instructed to encode semantically. he will do so, but in an impaired manner.
(pp. 74-75)

Butters and Cermack assumed that the Korsakoff patients'

deficient utilization of "meaning" in learning tasks was a specifically verbal (i.e., left hemisphere) phenomenon. However, Gazzaniga and his colleagues produced evidence which suggests that the right hemisphere may play an important role in imparting "meaning" to verbal memory processes. These authors tested patients with partial or complete section of the cerebral commissures for recall of two lists of paired-associate nouns. On presentation of the second list each patient was instructed to "form a 'picture in his mind' of the two items interacting in some unusual or amusing way" (Gazzaniga, Risse, Springer, Clark & Wilson, 1975, p. 12). Patients with partial sectioning of the cerebral commissures showed marked improvement with the imagery instructions but none was seen where there was complete section of the hemispheric interconnections.





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Discussion

The evidence reviewed thus far suggests that the central problem in the amnesia syndromes involves mechanisms which normally facilitate access to stored memory traces. The problem might occur at the point of encoding and deposition, or retrieval, or both. The hippocampus appears to be critical to these coding and decoding operations; this structure may normally provide the memory "cues" which must be externally administered in its absence.

The evidence is supportive of Niesser's (1967) proposal that

(rather than tapping a "continuous memory strip") "Penfield's electrode may have touched on the mechanisms of perceptual synthesis" (p. 169). There can be little doubt that Penfield's "final common pathways" in the temporal lobes are related to hippocampal afferents. However, it is now clear that the hippocampus cannot be the only secondary ganglionic station which must be activated before a signal indicating familiarity, originating in the right hemisphere, can "rise into consciousness." Studies of split-brain patients have shown conclusively that the ability to give a verbal account of events occurring in the right hemisphere is dependent on the integrity of the cerebral commissures (e.g., Sperry, 1968). The general layout of the commissures is such that a specific area in one hemisphere is connected via commissural fibers to the homologous area in the contralateral hemisphere. The temporal lobes have their own private interconnection in the anterior commissure, wiiijch also connects the two amygdalae (Gray, 1977). Studies of patients with only partial sectioning of these commissures have indicated






69


that highly processed information might be even more "transferrable" than elementary sensory information and may be able to utilize any commissural pathway that is available (Gazzaniga- et al., 1975).

It appears that there are two separate and autonomous memory systems in the brain which are specialized as to their function: a verbal system, lateralized in the left hemisphere, and a nonverbal (experiential) system lateralized in the right. The evidence suggests that memory functions might be conceptualized as having both vertical and lateral dimensions: the scanning (or search) within each system may involve a temporal-hippocampal interaction and the phenomenon of recognition may be a function of right-left temporal lobe interaction.

A voluntarily initiated search of memory would, most certainly, begin in the left-hemisphere system, but an environmental stimulus might activate an initial nonverbal (right-hemisphere) scan and analysis. In either case, the result of these processes would seem to be the emergence in conscious awareness of a signal indicating the "familiarity" (or "strangeness") of the stimulus and, finally, the facilitation of generalized expectations and verbal associations related to that stimulus. The implications of such a formulation

for the understanding of psychopathological processes and the practice of psychotherapy will be discussed in later sections, following an evaluation of the limbic system's role in experiential and emotional memory.






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The Limbic System, RAS, and Memory

Little distinction is made in the human amnesia literature

between the verbal, experiential, and emotional aspects of memory. The verbal manifestations of the disorder are the most obvious, the most amenable to description and testing, and so have become the primary focus of scientific attention. Although it is seldom emphasized, most case studies of amnesia victims relate anecdotal evidence of emotional dysfunction (e.g., blunted or flattened affect; euphoria). There are also more or less vague, but consistent, references to what might be termed disturbances of tonic arousal ("decreased spotaniety," "lack of initiative," "indifference," "passivity"). These emotional and arousal difficulties appear to be associated with systems centered on the amygdala and RAS, respectively. Both of these will be described below following a consideration of the nonverbal, predominantly unconscious mechanisms of experiential memory.

Before an organism can respond to a stimulus on any level

(emotional, verbal, or behavioral) its meaning must be ascertained. At the most elementary level the organism's survival depends on its ability to make appropriate decisions about whether to invest neural energy in attending to and further analyzing a particular stimulus (orienting) or to ignore it (habituation). Such a decision demands a judgement as to the apparent novelty, possible significance, or lack of these qualities in the stimulus configuration, a process which requires access to a patterned memory trace or "neural model" (Sokolov, 1963). Such a process must begin with sensory





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input and end with afferents which are capable of modulating the

activity of the brainstem RAS.

A determination of novelty or significance might be made at the level of the secondary sensory association cortices where raw receptor impulses are converted into functional (i.e., "meaningful") information, although such duplication of effort in all the modalities would be cumbersome and inefficient. Further, the relative significance of a stimulus may depend on the internal state of the organism (e.g., satiation, the presence or absence of certain hormones, etc.). The fact that information related to this added consideration is most readily available in subcortical structures is another argument favoring a centralized location for the mechanisms involved with the decision to orient or habituate. The hippocampus meets all of the criteria specified above.

Luria concluded that "many nuerons in the hippocampus and connected nuclei do not respond to modality-specific stimuli of any sort, but serve to compare present stimuli with traces of past experience; they react to every change in the stimulus and thus play to some extent the role both of 'attention neurons' and of 'memory neurons"' (1973a, p. 289). According to Luria the hippocampus provides for the "elimination of responses to irrelevant

stimuli and enables the organism to behave in a strictly selective manner" (1973a, pp. 271-272). It appears that the hippocampus accomplishes this complex task by coordinating the activities of the cortical and subcortical mechanisms which are directly involved in the processes of attention, memory and learning.





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Early investigators were puzzled by the fact that cortical

activation (EEG desynchronization) was accompanied by synchronous slow-wave activity (4-8 Hz theta rhythms) in the hippocampus (e.g., Green & Arduini, 1954). The most constant behavioral correlation of hippocampal theta activity in animals of different species is orienting towards, and attending to, stimuli in the environment (Isaacson, 1974). Cortical activation, such as that seen in the orienting response, is accomplished by the brainstem RAS in conjunction with the nonspecific thalamic nuclei. The hippocampus appears to regulate the process of involuntary attention by performing switching functions through a mutually inhibitory relationship with the reticular formation (Smithies, 1966). The Soviet neuropsycholgist Vinogradova provided important insights into the mechanics of this process.

By observing unit activity with microelectrodes Vinogradova

(1970) determined that all of the neurons in the hippocampus monitor incoming stimuli, habituating to repetition and dishabituating to any change in the stimulus configuration. Such responsiveness requires constant matching of the stimulus with a related neuronal model (Sokolov, 1963). That these models exist in the cortex, and not in the hippocampus itself, is established by evidence that the quality of sensory information is almost completely erased in hippocampal neurons (Gloor, 1961).

Vinogradova distinguished two types of neurons in the hippocampus: A-neurons (30-40%) which are activated by a stimulus and I-neurons (60%) which are inhibited. She went on to propose a mechanism whereby the hippocampus is able to modulate the processes of attention and learning:





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The hippocampus exerts a tonic inhibitory influence upon
the reticular formation, blocking activatory processes
through the tonic discharge of its I-neurons when
novelty is absent and registration [a change in the
neuronal model] is not needed. But when a stimulus which
is not registered in the memory system appears, this
inhibitory control is blocked (I-neurons become silent),
arousal occurs, and the process of registration starts.
(1970, p. 114)

Vinogradova's hypothesis is supported by observations of electrical activity in the brains of animals in classical conditioning paradigms. Theta rhythms (associated with activity of Vinogradova's A-neurons) are found in the early stages of learning but disappear when the response has become well established (Isaacson, 1974). During conditioning the time course of the theta rhythm and the orienting response are matched. As the latter is replaced by the stabilized conditioned response, theta dies out (I-neurons become active again) and the hippocampus resumes its inhibitory control-over the RAS, thus ending the orienting response and thereby allowing the fully developed conditioned response to materialize (Smithies, 1966, p. 90).

The hippocampal-cortical interaction was apparent in an

analysis of the characteristics of hippocampal theta rhythms in situations requiring different types of cortical information processing. Bremner (1970) investigated the effects of orienting, simple conditioning, discrimination, and discrimination reversal tasks on various parameters of the theta rhythm using the habituated organism (rat and man) as a baseline. He found that theta*.power (amount of energy) increased in the presence of stimuli which elicited orienting and arousal and decreased in the interval preceding a response in the conditioning situation; the range of energy distribution





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around the peak frequency narrowed during discrimination; and the location of the peak shifted in discrimination reversal procedures.

In summary, the hippocampus appears to be able to monitor

incoming stimuli and match them against (cortical) neuronal models which represent the past experience of the organism with related stimuli. In addition to facilitating appropriate access to these memory traces hippocampal activities have been directly associated

with the triggering of cortical processes which permit further analysis of a stimulus, emotional and behavioral responses as warranted, and/or alterations in the neuronal model itself (i.e., learning).

Papez Circuit and Memory

The hippocampus forms part of a continuous pathway within the limbic system which Papez (1937) believed to be the substrate of emotion. Papez was aware that destruction or stimulation of limbic structures produced major alterations in emotional behavior and believed that emotional expression depended on the integrative action of the hypothalamus. He was also convinced that subjective emotional

experience required the participation of the cerebral cortex. Papez outlined an anatomical circuit through which he thought

emotion might arise in either of those two centers. Thus

Incitations of cortical origin would first pass to the hippocampal formation and then down by way of the fornix to the mammillary body. From this they
would pass upward through the mammillothalamic tract
t'eto the anterior nuclei of the thalamus and thence
by the medial thalamocortical radiation (in the cingulum) to the cortex of the gyrus cinguli .
Radiations of the emotive process from the gyrus cinguli to other regions in the cerebral cortex






75


would add emotional coloring to psychic processes
occurring elsewhere. (Papez, 1937, pp. 304-306)

Papez believed that sensory input to the system originated in the thalamus and was communicated via the subthalamus to the hypothalamus. The circuit is completed with the connection of the cingulate gyrus to the hippocampus by way of the cingulum bundle.

It is now known that other limbic structures are more actively involved in the specifically emotional processes. However, Papez's anatomical concepts might be rehabilitated if their context were changed from emotion to memory. Input to the hippocampal circuits would be seen as processed sensory information and its output as

memory indexing information capable of "cueing" associations and generalized expectations related to the input stimulus. It remains to place the role of the hippocampus in the context of the functional system it subserves and to examine the other components of that system.
Fornix. Hippocampal output makes its way via direct and

indirect pathways to the thalamus (anterior, dorsomedial, and intralaminar nuclei), the hypothalamus, and the midbrain reticular formation. Its main efferent fiber system, the fornix, is composed of axons from pyramidal cells in the body of the hippocampus. These fibers converge in the fimbria, traveling backwards within the temiporal lobe, and then arch forward under the corpus callosum as the crura (posterior pillars) of the fornix. Here a number of fibers cross to the other side, forming the hippocampal commissure. The two crura then join to form the body of the fornix which






76


continues to arch forward, following the course of the lateral ventricle to the rostral edge of the thalamus. Here the bundles separate again to form the anterior columns of the fornix which curve downward in front of the intraventricular foramen and above the anterior commissure (AC). Approximately half of the fibers descend behind the AC as the postcommissural fornix; the other half, in a less compact bundle, pass in front of the AC as the precommissural fornix. Postcommissural fibers pass through the hypothalamus to

the mammillary bodies, giving off fibers to the thalamus on their way. Some of the precommissural fibers distribute to the septal area and others join with septal fibers and continue into the same areas as the postcommissural fornix. Approximately one-third of the fornix fibers reach the mammillary bodies (Daitz, 1953); the anterior nuclei of the thalamus receive as many direct fibers from the fornix as they do from the mammillothalamic tract (Truax & Carpenter, 1969, ch. 21).

Liss (1968), working with the rat, found that hippocampal

and fornix lesions had analogous effects on learning and behavior in passive and active-avoidance tasks. In the monkey, fornix lesions led to impaired learning of a spatial reversal task that was "functionally similar" to the deficit seen after hippocampal removal (Mahut, 1972; Mahut & Zola, 1973). Gaffen (1972) reported a series of six experiments in which rats with fornix lesions were shown to have a defect in "recognition memory" that the author argued was equivalent to anterograde amnesia in humans. In Russel's (1971) survey of brain wound cases, the eight patients who showed






77


a Korsakoff-type of memory disorder were thought to have had their fornices damaged by metal fragments. Four of the five cases with typical amnesic syndromes in Jarho's (1973) study of brain-injured war veterans were considered to have bilateral interruption of the fornix or mammillothalamic tract.

Sweet, Talland, and Ervin (1959) reported the case of a woman

in whom the anterior columns of the fornix were sectioned to facilitate the removal of a colloid cyst of the third ventricle. This patient showed a rapid recovery of old skills, but developed a "severe loss of memory for recent events which persisted at two years] . a retrograde amnesia of at least several weeks and a subjective

complaint of amnesia for specific events of four or five years past" (p. 76), coupled with intact remote memory. In the discussion following the presentation of this patient, Brenda Milner reported on a similar case operated on by Welsh in 1954. This patient showed a gross initial memory disorder but, at one year, was able (with effort) to effect some compensation for his defect: "Whenever he deliberately sought associative links he was able to improve his performance considerable" (Sweet. et al., 1959, p. 79). Milner concluded that.

I think that one can only account for the paradoxical
diversity of data from the fornix cases, as contrasted
with the consistent and severe memory loss in the hippocampal cases (and maybe in the mammillary body cases
also), by supposing that you are only interfering
with a part of the system by fornix section. Thus,
you are most apt to see a temporary disruption of disturbance of memory with minimal residual loss. '(p. 79)






78


Data on the fornix is relatively scarce, and there have been reports of negative findings. In his influential review, Brierly (1977) took a very conservative stance on this subject:

The most discrete link between the hippocampal and
diencepahlic regions is the fornix. It is surprising,
therefore, that with the exception of the case reported
by Sweet, Talland and Ervin (1959), bilateral division
of the fornix (usually in the region of the intraventricular foramen) has not resulted in a disorder of memorizing (Dott, 1938; Cairnes & Mosberg, 1951;
Garcia-Bengochea and his colleagues, 1954). This finding suggests that the two groups of structures
linked by the fornix cannot be regarded as a unitary
system subserving the process of memorizing, at least
until major interconnections other than the fornix
have been identified. (pp. 221-222)

Other interconnections are available. Smithies (1966) describes a "massive direct hippocampal-hypothalamic pathway" that runs
diffusely through the subthalamus and which he suggests miaht be "quite able to carry on hippocamoal and limbic circuit function in the absence of [the fornix]" (p. 122). A seond look at the reports

cited by Brierley, however, allows the possibility that his conclusions are premature.

Sweet et al. (1959) emphasized that their patient's "conversations, social amenities, and general demeanor gave little evidence of [her] severe deficits unless they were specifically looked for" (p. 76). They also noted that she lacked spontaniety, made little

effort to converse, and was apparently indifferent to her deficits (cf. Korsakoff's syndrome). The very brief report of Bengochea, De la Torre, Esquival, Vieta and Fernandez (1954), after a "short follow-up" of their patients (whose fornices were severed in an experimental operation to relieve intractable epilepsy) did not






79



mention any attempt at quantification of behavior. They simply stated that "so far, in none of the 12 surviving cases there has been [sic] any unfavorable neurological or psychiatric sequela" (p. 177). It seems possible that these authors may have missed subtle symptoms in their apparently superficial evaluation.

Wilder Penfield (quoted in Sweet et al, 1959) underscored the fact that "patients who have [bilateral hippocampal lesions] do not forget their skills. Two of them were able to carry out most complicated skills learner previous--glove cutting and engineering drawing" (p. 81). Unfortunately, the only behavioral measure reported by Cairnes and Mosberg (1951) involved a return to work. These authors noted that some of their pateints (who had incurred fornix damage in the course of surgery to remove colloid cysts of the third ventricle) showed initial confusion, loss of memorizing, and amnesia for the period surrounding the operation, but: "after operation all [but one of their nine surving cases] returned to work, and . .showed no disturbance of emotion or intelligence" (p. 564). Thus

Four of the five young women ...are doing normal
housework; three have borne childrn. The other young woman is in regular work as a clerk, and is
free from complaints . . Two older women ...
are also doing their housework [although one has a
'slight impairment of memory'] . . Of the two men,
one is working regularly as a policemen. (p. 568)

The extent of the lesions in these patients is unclear. The authors report only that "each had . partial or complete division of the anterior columns of the fornix" (p. 564). (It seems possible that these surgical fornicotomies spared the precommissural fornix.)





80


It should be noted that a colloid cyst of the third ventricle tends to produce confusion, dulling of attention and memory, and sometimes a progressive dementia prior to its surgical removal. These factors would make a pre-post evaluation of memory function very difficult. Still, the scantiness of the reported date in the studies reviewed above is unfortunate. It is evident that injuries to different parts of this system result in different expressions of the disorder. It is reasonable to conclude, however, that damage to the fornix has adverse effects on memory function which vary as to the quality and degree, and may leave a greater possibility

of recovery of function.

Mammillary bodies and mammillothalamic tract. The mammillary bodies are a collection of nuclei at the posterior boundary of the hypothalamus. They form a major relay station for hippocampal output on its way to the thalamus (via the mammillothalamic tract) and to the midbrain reticular formation (by way of the mammillotegmental tract).

In humans, damage which is apparently limited to the mammillary bodies has resulted in the full Korsakoff amnesic syndrome (Remy; 1942; Delay & Brion, 1951; Gruner, 1956; Symonds, 1966), although Victor (1964) suggested that additional damage to the thalamus was necessary to produce the disorder. In the rat, lesions of the mammillary bodies or of the mammillothalamic tract impaired the ability to perform a spatial discrimination in a T-maze in order to avoid footshocks (Thompson, Langer & Rich, 1964). Krieckhaus (1962, 1964) found that complete or partial destruction of the






81


mammillothalamic tract in the cat reduced the retention of a two-way active avoidance task and produced a less striking deficit in the retention of a one-way active avoidance task. These findings were later replicated in the rat (Krieckhaus, 1965). Thomas, Frey, Slotnick and Krieckhaus (1963) studied the post-operative acquisition of the two-way active avoidance learning task and reported mixed results: four of their eleven cats were completely unable to learn the task, while seven of them mastered the problem within the number of trials required by normal animals. (The significance of the distinction between acquisition and retention will be discussed in the following section.)

Cingulate cortex. The cingulate cortex lies above the

corpus callosum on the medial side of the hemisphere, separated from the neocortex above by the cingulate sulcus. It merges with the hippocampal gyrus posteriorly and with the neocortex of the frontal lobe anteriorly. As noted by Papez, the cingulate gyrus receives its main afferent supply from the anterior nuclei of the thalamus and projects to the hippocampus via the cingulum bundle. Stimulation of the cingulate cortex also produces activity in the prefrontal and orbitofrontal regions of the neocortex (Dunsmore & Lennox, 1950). There are reciprocal connections with the anterior and other thalamic nuclei (including the dorsomedial). A strong projection to the interior parietal lobule (IPL) in the post-central neocortex has been demonstrated in the monkey (Mesulam, Van Hoesen, Pandya & Geshwind, 1977). These authors, using the horseradish peroxidase technique, found that "the cingulate gyrus contained one





82


of the heaviest concentrations of labeled neurons in most cases" following injection of that substance into the IPL (p. 324).

The literature documenting the efforts of physiological

psychologists to define the role of the cingulate cortex in learning and memory is confused somewhat by varying interpretations of the data by those authors (an objective review is available in Isaacson, 1974). Several experimenters have ascribed the learning deficit which follows cingulate lesions to an enhanced fear response. However, Kimble and Gostnell (1968), using two different behavioral measures, failed to find any support for this hypothesis. Lubar, Perachio, and Kavanagh (1966) suggested that incidental damage to the visual cortex could account for the deficits following cingulate lesions, but adequate control lesions (e.g., Kimble, 1968) and lesions which avoid damage to the visual cortex (Trafton, Fibley & Johnson, 1969) are adequate proof of the cingulate's role in the observed impairments (Isaacson, 1974). The essential findings on the effect of cingulate lesions on memory in the rat and cat are straightforward: such lesions impair the ability of these animals to acquire active avoidance conditioning (Peretz, 1960; McCleary, 1961; Thomas & Slotnick, 1962; Lubar & Perachio, 1965; Kimble & Gostnell, 1968; Trafton et al., 1969).

A number of relatively comnlex deficits have been noted after cingulate damage which resemble symptoms seen in human amnesia syndromes. These include disruption of the orderly-sequencing of behaviors (in nest-building and maternal behavior, Stamm, 1955; Slotnick, 1967), deficits in the temporal ordering of responding






83


(in runway problems and bar-press alternation: Barker & Thomas, 1965, 1966; Barker, 1967); and the failure to exhibit behaviors which were indicative of opium addiction (Marques, 1971). The last, especially, is supportive of the hypothesis voiced by some authors that cingulatelesioned animals are unable to anticipate the emotional consequences of their behavior for both rewards and punishments (Glass, Ison, & Thomas, 1969; Isaacson, 1974).

Anterior cingulectomy has been termed the psychosurgical
"operation of choice" for the treatment of severe obsessional and anxiety disorders (Lewin, 1961; Whitty, 1966). Whitty noted that one of the long-term effects of cingulectomy was "relative neglect of the impact of external events." Finally, Pechtel, McAvoy, Levitt, Kling & Massermann, (1958) concluded that lesions of the cingulate gyrus in humans resulted in, among other things, "amnesia for previous learning" and "impairment of new learning skills." Discussion

The failure to transfer learning between hemispheres seen in

cerebral commissurotomy preparations (animal and human) indicates that the storage of memories is a neocortical function (Geshwind, 1965). Generalized memory disorders, on the other hand, are only produced by bilateral damage to the diencephalic structures of the hippocampal system. It is evident that these structures in the circuit of Papez form part of a functional system which monitors the environment, matches incoming stimuli with representations of the organisms previous experience with similar stimulus configurations, activates the organism in the presence of potentially significant stimuli,






84


and finally, causes pertinent information concerning that stimulus to be presented to conscious awareness. It appears that there are two such systems in the human brain, each specialized to deal with a different type of information. The first, lateralized to the right hemisphere, performs the functions listed above with experiential data. The second, operating in the 16ft hemisphere, deals with language and verbal concepts which may be based on gestalten that are assembled and stored in the contralateral system. All of the functions noted above are impaired, to a greater or lesser extent, in various manifestations of the amnesic syndrome.

The evidence suggests that some learning does not take place in amnesia victims, that is, memory traces are stored. However, the amnesic subject has difficulty gaining access to those memory traces when they are needed. More specifically, they are unable to recognize and select the appropriate memory trace in a given situation from the set of available traces. Access to the proper trace is facilitated with adequate cueing. It appears that the function of the hippocampal system is to assure the activation of appropriate memory traces based on the requirements of the situation. In the present formulation, significant (experientially based) memory traces have been designated by the superordinate term "generalized expectations." It appears that the hippocampal system is responsible for the generalizing of these expectations from one situation to another, similar, situation.

In lower form, where survival is dependent on instincts, the

identification of a significant stimulus may culminate in the release





85



of species-specific behaviors that are organized at a subcortical level. The elicitation of unlearned behavior sequences (related to feeding, fighting, fleeing and reproduction) by hypothalamic stimulation is well established and seems to be related to programs: stored in the midbrain or brainstem (Isaacson, 1974). In humans, however, cortical functions are more important in determining the behaviors that insure survival.

Papez's circuit is an "anatomical identity which connects the temporal and cingulo-frontal cortex of both hemispheres" (Barbizet, 1963). Given the recently identified massive interconnection of the cingulate gyrus with the inferior parietal lobule (IPL), Papez's circle may be seen as forming a part of a larger cortical circuit (see Fig. .). As such, it is in a position to directly modulate the activity of the two areas of the brain that are identified with the highest levels of information processing: the specifically human tertiary association areas of the prefrontal lobe and the IPL.

Information flow within this cortical-hippocampal circuit

might be seen as follows: raw sensory data enter the system in the primary projection areas and are processed in the secondary association areas of the post-central cortex; as this information acquires meaning it is passed on to the temporal cortex and into the hippocampal circuit; here the meaningful bits are translated into a code that emerges in the cingulate cortex as memory indexing information; this, in turn, is referred to the IPL where it facilitates associations that fine-tune the continuing input into the system. This process continues until an adequate match is made and recognized at the








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87



level of the temporal cortex. This match would take the form of a

finished "gestalt" in the experiential system and a formal "concept" in the verbal system. With the appearance of such a match the temporal component would terminate the search process and relay the product to the c~ntralateral system. Thus, the recognition of a gestalt might initiate the search for a related verbal concept or, conversely, a verbal formulation may trigger a scan for pertinent experiential associations. The description of these two complementary systems seems to provide adequate explanation for the qualitatively different types of memory which were described by Breuer~and Freud and by Rapaport, respectively as noted at the beginning of this section.

Substantial support for the model described above may be found in a situation where hippocampal activity is induced from within the limbic system (rather than from the neocortex, as was the case in Penfield's experiments). Such is the case when amygdala stimulation produces after-discharges in the hippocampus. Halgren (1981) reviewed the effects of amygdala stimulation in conscious humans and noted that such stimulations sometimes produces "complex formed hallucinations, sometimes complete scenes as in a dream or vivid recollection and sometimes more vague, apparently similar to an intruding thought .. and illusions of familiarity (deja vu)" (p. 345). Halgren suggested that it is the activation of distant normal tissue subsequent to amygdala stimulation which produces the resulting mental phenomena. He cites evidence that





88


Amygdala stimulation seldom evokes a mental phenomenon
unless it also evokes an after-discharge . most amygdala stimulations, even if they evoke an afterdischarge (AD), do not evoke any reported mental phenomenon . thus, there is no direct connection between amygdala activity and hallucinatory experiences ....
Simultaneous recordings from multiple brain areas
indicate that amygdala ADs seldom remain localized.
Initial spread is to the ipsilaterla hippocampus and
hippocampal gyrus. AD may remain confined to these
structures, in which case no mental phenomenon is
necessarily evoked. Further spread is to the ipsi
lateral limbic cortex (orbital, insular and cingular) and diencephalon (especially the anterior nucleus of
the thalamus, but also to the centre median, pulvinar
and dorsomedial nuclei). ADs seldom spread to the
neocortex, which may however desynchronize." (pp. 396397, emphasis added)

These findings are congruent with the present formulation, which would interpret the appearance of "complex formed sensory hallucinations" following the stimulation described above as the result of hippocampal output evoking experiential memory traces in the posterior association cortex by way of the anterior thalamus and cingulate cortex. The appearance of identical "experiential hallucinations" following temporal lobe stimulation and limbic system ADs supports Penfield's contention that final common pathways in the temporal lobe may produce activity in the hippocampal system which utlimately results in the appearance of mental phenomena.

The hippocampal system is directly involved in the mechanics

of learning. Hippocampal activity seems to assure that the organism attends to novel stimuli and sets the conditions which permit changes to be encoded into the neural models which form the centeral representations of those stimuli. Pribram and McGuinness (1975) suggested that such changes in neural represenations "may be conceived as






89


changes of state, set, or 'attitude'" (p. 132). Such experientially based alterations in the disposition of the organism relative to a stimulus object, situation, or event provide an operational definition for the generic term "generalized expectation" as used in the present formulations. This type of memory appears to be the "idea" which Breuer and Freud believed to be of central importance in the etiology of psychoneurosis. (It will be suggested that it is precisely these generalized expectations, in this memory system, which will provide the answer to the question posed in the introduction: "What is to be changed in the process of psychotherapy?")

In addition to their mnemonic defects, amnesia victims with damage to the hippocampal systems also present anomalies in their affect and arousal. The latter may be traced to the failure of the hippocampus to trigger cortical activation (via the RAS) as it normally does in conditions of uncertainty or significance. The former seems to reflect the disruption of hippocampal modulation

of the emotion/motivation processes by way of its interactions with other limbic structures and with the prefrontal areas (via the dorsomedial thalamus).

Interfaces and Interactions of the Monitoring
Motivating, and Mobilization System
Pharmacological research with animals and humans has implicated the two classes of biogenic amines in the modulation of fundamental brain processes. The catecriolamines dopamine (DA) and its metabolite norepinephrine (NE) have been associated With cognitive function and dysfunction (e.g., schizophrenia) while affective disorders seem to





90


involve an interaction between NE and the indolamine serotonin (5-HT). Histological researchers have identified six major mono amine systems in the rat brain (Fuxe, 1965; Fuxe, Hamberger &

Hokfelt, 1968).

The nigro-striatal DA system originates in cell bodies in the

substancia nigra whose axons extend through the lateral hypothalamus to terminate on cells in the caudate nucleus. This tract is known to regulate the activity of the ancient extrapyramidal motor

control system.

The meso-limbic DA system originates in the ventral tegmental area of the pons and projects mainly to the septal area and related nuclei in the limbic forebrain. Dysfunction in this system is

assumed by many to be the source of schizophrenic disorders.

The meso-cortical DA system is less well defined but appears to include projections from the ventrotegmental to the frontal cortex and from the substancia nigra to the anterior cingulate cortex (see Meltzer, 1979).
The ventral NE system originates in the reticular formation (medulla and pons) and ascends through the median forebrain bundle (MFB) to terminate on cells throughout the hypothalamus and amygdala. Stein, Wise and Berger (1972) suggested that this system primarily regulates motivational activities.
The dorsal NE system arises in the locus ceruleus in the pons and may supply up to 70% of the NE in primate brains (Redmond, 1979). The axons in this system ascend through the MFB and give off branches to the hypothalamus, the hippocampus and amygdala,






91



the septal area (which receives the bulk of these terminals), the anterior-ventral thalamus, the cingulate gyrus and the neocortex. Stein and Wise (1971) suggest that this dorsal NE system is involved in regulating cognitive activities.

The ascending serotonin (5-HT) system originates in the median raphe' nucleus which is situated in the core of the reticular formation and receives collateral branches from the sensory nerves. 5-HT cells in the raphd; have terminations throughout the central gray of the midbrain. Axons from the raphe' also rise in the MEB and distribute to the same areas as the dorsal NE system, with the addition of terminals in the basal ganglia and more widespread distribution in the neocortex. This system seems to be most directly involved in regulating tonic arousal.

Brodie and Shore (1957) suggested that NE and 5-HT exert

opposing effects that modulate a variety of CNS functions (e.g., sleep, appetite, sexual drive). It appears that these parallel systems control behavior by reciprocal action in balanced systems.

While functional specificity is determined by the neural circuitry involved at a given anatomical level, the cumulative effects of specific functional outcomes produced a preponderance of one or

the other transmitter seem to be consistent and to result in coordination across levels. It appears that a preponderance of the indolamine 5-HT, for example, results in the release of tonic mobilizing energy at the brainstem level, the suppression of ongoing behavior at the hypothalamic level, the identification of negativee reinforcement" at the limbic system level, and




Full Text
CHAPTER III
METHOD
This study was designed to affirm or disaffirm the consequents of
the proposed theoretical model of personality function and psychopathol
ogy and to demonstrate a confluence of psychological and neurological
observations. Specifically, the investigation attempted to ascertain
whether groups of adult psychiatric patients classified according to
the constructs of the proposed model as suffering from hypo- or hyper
dominance spectrum disorders differed significantly from each other on
a ratio of test scores that have been demonstrated to be sensitive to
damage to the left and right cerebral hemispheres, respectively,
without differing significantly in overall performance on the
instruments.
Both spectrum disorders are personality disturbances defined by
chronically diminished social performance and/or dysfunctional behavior
patterns without a clear precipitating factor. Both exclude gross
impairments of perception, orientation or memory. Hyper-dominance
spectrum disorders are defined as disorders in which the primary
symptoms are manifested intrapsychically and/or are characterized by a
central tendency to maladaptive overutilization of formal thought
processes. Hypo-dominance spectrum disorders are defined as disorders
in which the primary symptoms are manifested extrapsychically and by
a central tendency to maladaptive underutilization of formal thought
processes. DSM III diagnostic categories which meet these criteria
are listed in Appendix A.
151


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89
changes of state, set, or 'attitude'" (p. 132). Such experiential ly
based alterations in the disposition of the organism relative to a
stimulus object, situation, or event provide an operational definition
for the generic term "generalized expectation" as used in the present
formulations. This type of memory appears to be the "idea" which
Breuer and Freud believed to be of central importance in the etiology
of psychoneurosis. (It will be suggested that it is precisely these
generalized expectations, in this memory system, which will provide
the answer to the question posed in the introduction: "What is to be
changed in the process of psychotherapy?")
In addition to their mnemonic defects, amnesia victims with
damage to the hippocampal systems also present anomalies in their
affect and arousal. The latter may be traced to the failure of the
hippocampus to trigger cortical activation (via the RAS) as it
normally does in conditions of uncertainty or significance. The
former seems to reflect the disruption of hippocampal modulation
of the emotion/motivation processes by way of its interactions
with other limbic structures and with the prefrontal areas (via the
dorsomedial thalamus).
Interfaces and Interactions of the Monitoring
Motivating, and Mobilization System
Pharmacological research with animals and humans has implicated
the two classes of biogenic amines in the modulation of fundamental
brain processes. The catecholamines dopamine (DA) and its metabolite
norepinephrine (NE) have been associated with cognitive function and
dysfunction (e.g., schizophrenia) while affective disorders seem to


18
The human brain has remarkable ability to compensate for
injury by utilizing undamaged tissue. This recovery of function
in the time elapsed between injury and testing is a further
confounding factor.
Much of this literature is in the form of case reports and the
problem of individual variability is especially difficult to manage.
Research designs utilizing groups and statistical analyses are
becoming more frequent, but these studies invariably suffer from
the difficulties noted above.
The most difficult problems to be encountered here have to do
with the specification of dependent variables: "higher brain func
tions" tend to resist definition and to defy quantification.
Generally, only gross operational definitions have been available.
The physician/experimentor was often forced to resort to a judgement
as to whether a given (and hypothetical) function was "intact"
or "lost," with the attendant risks of experimentar bias and error.
Related to this is the understanding, which emerged slowly in the
development of this literature, that the function of a circumscribed
anatomical area cannot be inferred directly from a deficit which
follows the ablation of that area: one can only specify how the
brain functions in the absence of that tissue.
Human Consciousness
Right versus Left
There have been three basic approaches to the study of lateralized
cortical function: the comparison of patients with unilateral brain
damage; the lateralized presentation of stimuli or task to the


35
There is a clear consensus recognizing two modes of informa
tion processing. However, the ability to process information is
not necessarily a sufficient condition for consciousness. Although
the notion of right brain "thought" has gained wide currency, to
date, there has been no conclusive evidence that any cognitive opera
tion occurring in the right hemisphere is directly experienced in
consciousness. A possiblity not considered by any of the above
authors is that one mode might be an ancillary resource utilized
by the other. The data reviewed thus far suggests that one must be
very careful to avoid anthropomorphizing when attempting to describe
right brain processes. However, some inductive conclusions may be
drawn.
Discussion
It is evident that human consciousness is inexorably linked
with the abstract symbolic processes associated with language. Perhaps
the most universally accepted characteristic of human thought is
self-awareness. Self-awareness (and its product, the self-concept)
requires the abstraction and appreciation of defining features which
are consistent over time and situation. Only the left hemisphere,
with its temporal acuity, can consider and aopreciate changed or
conserved relationships in different conditions or contexts. Thus,
only the left hemisphere can define itself. The resulting self-
awareness provides a reference point for all of the memories, feelings,
intentions and thoughts that are collectively known as the- mind
and which allow the individual, thus defined, to interact intelligently
with the environment. Lacking the temporal organizing skills to


106
By altering the cortical evoked response paradigm so that the
second of a pair of responses could be expressed as a function of
the first Pribram (1967) was able to make imoortant inferences
about the organization that is imposed on incoming information
When a double click or a double flash is used to evoke
a neural response, the amplitude of the second of the
pair of responses serves as an indicator of the duration
over which a part of the system is occupied in processing
the first of the pair of inputs. A depression in the
amplitude of the second of the responses thus indicates
a longer recover--a longer processing time for a signal
within the channel. Such an increase in processing
time effectively desynchronizes the channel to repetitive
inputs: fewer fibers are available for processing
any given signal in the series. Prolongation of re
covery thus reduces redundancy in the channel. At any
moment more information can be processed. . Thus
the rate of information processing is enhanced. (Pribram,
1967, p. 460)
Augmented redundancy in information processing channels would mean
that more neurons were available to fire in a stimulus-linked
fashion. Thus, the sensitivity of the organism would be increased
(e.g., decreased pain tolerance, increased subjective experience
of pain). Since information tends to be "chunked" in this mode
there would be a decrease in the focusing of attention on details
and decreased articulation of experience. Conversely, reduced
redundancy in information processing channels would leave fewer
neurons available to fire in a stimulus-linked fashion, thus
reducing the sensitivity of the organism; bits of information would
tend to be separated in this mode permitting increased focusing
of attention on details and increased articulation of experience.
It will be noted that the augmentation mode would favor the performance
of cognitive tasks associated with the orienting response (i.e.,


19
sensory receptor fields of normal subjects; and the study of patients
whose cerebral hemispheres have been surgically separated (Nebes,
1977). The early studies of split-brain subjects prompted a great
deal of speculation about the abilities of the right hemisphere
and a listing of the published works concerned with asymmetrical
hemispheric functioning is beyond the scope of this paper. Evidence
pertaining to the differences between the cognitive operations
performed by the left and right hemispheres has been ably reviewed
by a number of writers (e.g., Bogan 1969a, 1969b; Galin, 1974, 1977;
Gazzaniga, 1970; Gazzaniga & Ledoux, 1977; Nebes, 1974, 1977;
Sperry, 1968). Some of the more important differences will be noted
very briefly here.
The primary deficits seen after right hemisphere lesions involve
difficulties in perceiving, manipulating, and remembering spatial
relationships and in perceiving and remembering sensory stimuli
which resist verbal description (Nebes, 1977). The right hemipshere
is superior to the left at generating a percept of the whole from
fragmentary information and seems predisposed to notice complex
gestalts or patterns rather than the parts (Nebes, 1971, 1974,
1977). The right hemisphere tends to process input simultaneously
(in parallel) as opposed to the left hemisphere's preference for
sequential (serial) processing, Cohen, 1973). In spite of many
claims to the contrary, the evidence indicates that the right
hemisphere considers only the immediately perceived context and
performs its tasks in a reflexive, automatic fashion.


163
behavior. The functional meta-system model accounts for this,
and the fact that split-brain subjects do not tend to develop
pathological symptoms, by hypothesizing that the right hemisphere
normally monitors the environment for significant (e.g., social)
stimuli, relates them to personal experience, and signals the left
hemisphere appropriately via the (subcortical) limbic system.
Perhaps the most important asset of the proposed model is the
fact that a rigorous and comprehensive test of the full range of
its predictions can be accomplished with existing technology. The
essential cortical-limbic system hyoerconnection postulate could
be evaluated by observing galvanic skin response (GSR) on the left
side in the hyper-dominant group. Augmentation of cortical averaged
evoked potentials (AEP) to increasing stimulus intensity would be
predicted in the hypo-dominant group and reduction in the other,
thus verifying the predicted cognitive mode. Differences in the
contingent negative variation (CNV) between obsessives, anxiety
neurotics, hysterics, and psychopaths were noted earlier and are
congruent with the tenets of the model.
An ideal test of the theory would include replication of the
cognitive and personality results of the present study, peripheral
physiological observations of indices of limbic system function (bi
lateral GSR), perceptual reactance (augmenting/reducing), and
information processing efficiency (CNV). This could be done with
cell sizes large enough to permit meaningful comparisons of
diagnostic entities to be made within and between grouDS. Such a
study might result in complete operational definitions of


51
encoding and retrieval of this information in memory and forwards
its signals to the prefrontal lobes where they are experienced as
subjective emotions. The prefrontal lobes appear to utilize these
signals in the process of forming intentions to direct adaptive
behavior. This amygdala-prefrontal pathway appears to be the sub
strate of anxiety and depression. When the amygdala-frontal connec
tion is severed surgically, or uncoupled pharmacologically, the
neocortex experiences euphoria, but fails to behave in an adaptive
manner.
Memory Functions
It is evident that the functional brain systems which form the
infrastructure of personality include separate cognitive, affective
and arousal components. It appears that these subsystems evolved
to take full advantage of a form of learning which utilizes reinforce
ment and emotional experience in determining behavior. The product
of any learning experience is memory. The importance of memories
(or "associations") in the organization of cognitive operations is
obvious and most, if not all, arousal and affective processes must
depend on the ability to discriminate personally relevant stimuli.
Clearly, memory is fundamental to all aspects of personality function,
but the material substrate of memory remains a complete mystery
(e.g., Lashley, 1950; Luria, 1973a); our concepts regarding it are,
of necessity, only abstract descriptions. Before reviewing the
physiological organization of memory systems it will be necessary
to define and delimit, as far as possible, those abstract


123
meta-system the right hemisphere performs the functions of an
environment "monitoring" system and the left hemisphere constitutes
a problem-solving and response-generating system; the amygdalae
and the left prefrontal lobe form an emotional "motivation" system;
and the reticular activating system; (with their brainstem, thalamic
and prefrontal lobe components) compose a cortical "mobilization"
system.
While the differences between the cognitive operations performed
by the left and right cerebral hemispheres may have been over
popularized little, if any, attention has been paid to the fact
that the older medio-basal and subcortical structures (which sub
serve the processes of emotion and attention) are also bilaterally
represented. It is common practice to refer to "the" limbic system
when in fact the components of this system are duplicated within
each half brain. It is reasonable to expect that these lateralized
structures are also specialized as to function, or reflect the
specialization of the neocortical mechanisms which they subserve.
In the context of the functional meta-system, motivation and arousal
must normally be activated by the right (monitoring) system and
responded to by the left (problem-solving and response-generating)
system. If this were not the case then the automatic nature of the
overall system (which is guaranteed by the relative isolation of
its components) might be defeated with the result that adaptive
behavior would not be assured in survival situations. A basic
assumption here is that mother nature would not relinquish instinct-
based responding and permit self-determined behavior without firm


BIBLIOGRAPHY
Aggleton, J. P., Burton, M. H., & Passingham, R. E. Cortical and
Subcortical Afferents to the Amygdala of the Rhesus Monkey
(Macaca Mulatta). Brain Research, 1980, 190, 347.
Arbuthnott, G., Fuxe, K., & Ungerstedt, U. Central Catecholamine
Turnover and Self-stimulation Behavior. Brain Research, 1971,
27, 406-413.
Austin, G. M., & Grant, F. C. Physiologic Observations Following
Total Hemispherectomy in Man. Surgery, 1962, j!8, 239-258.
Bailey, F. W. Histopathology of Polioencephalitis Hemorrahagica
Superior (Wernicke's Disease). Archives of Neurology and
Psychiatry, Chicago, 1946, 56_, 609-630.
Baker, L. J., Kesner, R. P., & Mi chal, R. E. Journal of Comparative
and Physiological Psychology, 1981, 95, 312.
Ball's, G. U. (Ed.). Clinical Psychooatholoqy. Boston: Butterworth,
1978.
Barbizet, J. Defect of Memorizing of Hippocampal-Mammillary Origin:
A Review. Journal of Neurology, Neurosurgery, and Psychiatry,
1963, 26, 127-135.
Barker, D. J. Alterations in Sequential Behavior in Rats Following
Ablation of Midline Limbic Cortex. Journal of Comparative and
Physiological Psychology, 1967, 64, 453-460.
Barker, D. J., & Thomas, G. J. Ablation of Cingulate Cortex in
Rats Impairs Alternation Learning and Retention. Journal of
Comparative Physiological Psychology, 1965, 60, 353-359.
Barker, D. J., & Thomas, G. J. Effects of Regional Ablation of
Midline Cortex in Alternation Learning in Rats. Physioloay and
Behavior, 1966, 1, 313-317.
Bear, D. M. Temporal Lobe Epilepsy, a Syndrome of Sensory-Limbic
Hyperconnection. Cortex, 1979, 15, 357-384.
Bear, D. M., & Fedio, P. Quantitative Analysis of Interictal
Behavior in Temporal Lobe Epilepsy. Archives of Neuroloqy,
1977, 34, 454-467.
168


69
that highly processed information might be even more "transferable"
than elementary sensory information and may be able to utilize any
commissural pathway that is available (Gazzaniga. et al., 1975).
It appears that there are two separate and autonomous memory
systems in the brain which are specialized as to their function:
a verbal system, lateralized in the left hemisphere, and a nonverbal
(experiential) system lateralized in the right. The evidence
suggests that memory functions might be conceptualized as having
both vertical and lateral dimensions: the scanning (or search)
within each system may involve a temporal-hippocampal interaction
and the phenomenon of recognition may be a function of right-left
temporal lobe interaction.
A voluntarily initiated search of memory would, most certainly,
begin in the left-hemisphere system, but an environmental stimulus
might activate an initial nonverbal (right-hemisphere) scan and
analysis. In either case, the result of these processes would seem
to be the emergence in conscious awareness of a signal indicating
the "familiarity" (or "strangeness") of the stimulus and, finally,
the facilitation of generalized expectations and verbal associations
related to that stimulus. The implications of such a formulation
for the understanding of psychopathological processes and the practice
of psychotherapy will be discussed in later sections, following
an evaluation of the limbic system's role in experiential and emo
tional memory.


70
The Limbic System, RAS, and Memory
Little distinction is made in the human amnesia literature
between the verbal, experiential, and emotional aspects of memory.
The verbal manifestations of the disorder are the most obvious, the
most amenable to description and testing, and so have become the
primary focus of scientific attention. Although it is seldom
emphasized, most case studies of amnesia victims relate anecdotal
evidence of emotional dysfunction (e.g., blunted or flattened
affect; euphoria). There are also more or less vague, but consistent,
references to what might be termed disturbances of tonic arousal
("decreased spotaniety," "lack of initiative," "indifference,"
"passivity"). These emotional and arousal difficulties appear to
be associated with systems centered on the amygdala and RAS,
respectively. Both of these will be described below following a
consideration of the nonverbal, predominantly unconscious mechanisms
of experiential memory.
Before an organism can respond to a stimulus on any level
(emotional, verbal, or behavioral) its meaning must be ascertained.
At the most elementary level the organism's survival depends on
its ability to make appropriate decisions about whether to invest
neural energy in attending to and further analyzing a particular
stimulus (orienting) or to ignore it (habituation). Such a decision
demands a judgement as to the apparent novelty, possible significance,
or lack of these qualities in the stimulus configuration, a process
which requires access to a patterned memory trace or "neural
model" (Sokolov, 1963). Such a process must begin with sensory


85
of species-specific behaviors that are organized at a subcortical
level. The elicitation of unlearned behavior sequences (related to
feeding, fighting, fleeing and reproduction) by hypothalamic
stimulation is well established and seems to be related to programs
stored in the midbrain or brainstem (Isaacson, 1974). In humans,
however, cortical functions are more important in determining the
behaviors that insure survival.
Papez's circuit is an "anatomical identity which connects the
temporal and cingulo-frontal cortex of both hemispheres" (Barbizet,
1963). Given the recently identified massive interconnection of
the cingulate gyrus with the inferior parietal lobule (IPL), Papez's
circle may be seen as forming a part of a larger cortical circuit
(see Fig. $). As such, it is in a position to directly modulate
the activity of the two areas of the brain that are identified with
the highest levels of information processing: the specifically
human tertiary association areas of the prefrontal lobe and the IPL.
Information flow within this cortical-hippocampal circuit
might be seen as follows: raw sensory data enter the system in the
primary projection areas and are processed in the secondary association
areas of the post-central cortex; as this information acquires meaning
it is passed on to the temporal cortex and into the hippocampal
circuit; here the meaningful bits are translated into a code that
emerges in the cingulate cortex as memory indexing information; this,
in turn, is referred to the IPL where it facilitates associations
that fine-tune the continuing input into the system. This process
continues until an adequate match is made and recognized at the


Figure 2. Partially schematized representation of the limbic system. FR CTX frontal cortex;
CING cingulate cortex; TEMP CTX temporal cortex; CC corpus callosum; SEPT septal area;
AMYG amygdala; HPC hippocampus; HT hypothalamus; MB mammillary bodies; ANT anterior
thalamic nuclei; HAB habenula; ILTN intralaminar thalamic nuclei; IP interpenduncular
nucleus; RF reticular formation; DBB diagonal band of Broca; MFB median forebrain bundle;
SM stria medularis; ST stria terminal is; FNX fornix. (Redrawn from Isaacson et al, 1973)


142
Cognitive activity in the right hemisphere is limited by
its inability to manipulate time. The right half of the brain
is context-bound, both internally and externally; its products are
reflexive, not considered. While the right hemisphere is responsible
for mediating the external expression of immediate affect the
appreciation of subjective emotional experience is a left-brain
function. Right brain activities are seen here as ancillary processes
which are utilized by the left. Contrary to recent theories
which tend to conceptualize psychopathology as emerging from the
disruption of a balance between right and left brain functions
(e.g., Bogan, 1969b; Flor-Henry, 1979; Tucker, 1981) the present
model emphasizes the need for integration of the influences which
converge on the left hemisphere. Any process which interfered with
the integrated functioning of the meta-system would be, by
definition, pathological. A process which culminated in the relative
overactivation of the left hemisphere would diminish the right
hemisphere's modulation of behavior and, conversely the right
hemisphere's contribution would be exaggerated when activation of the
left was reduced or disrunted. Such processes would be exacerbated
by the effects of reciprocal transcallosal inhibition (Flor-Henry,
1979). Since considered, "self-determined" responding is coordinated
by the left hemisphere such processes would lead to pathological
conditions of hyper- and hypo-dominance, respectively.
In the following paragraphs standard psychodiagnostic entities
will be related to neurological indices which reflect the operation
and interaction of elements within the proposed functional meta-system.


82
of the heaviest concentrations of labeled neurons in most cases"
following injection of that substance into the IPL (p. 324).
The literature documenting the efforts of physiological
psychologists to define the role of the cingulate cortex in learning
and memory is confused somewhat by varying interpretations of the
data by those authors (an objective review is available in Isaacson,
1974). Several experimenters have ascribed the learning deficit
which follows cingulate lesions to an enhanced fear response. However,
Kimble and Gostnell (1968), using two different behavioral measures,
failed to find any support for this hypothesis. Lubar, Perachio,
and Kavanagh (1966) suggested that incidental damage to the visual
cortex could account for the deficits following cingulate lesions,
but adequate control lesions (e.g., Kimble, 1968) and lesions which
avoid damage to the visual cortex (Trafton, Fibley & Johnson, 1969)
are adequate proof of the cingulate's role in the observed impairments
(Isaacson, 1974). The essential findings on the effect of cingulate
lesions on memory in the rat and cat are straightforward: such
lesions impair the ability of these animals to acquire active
avoidance conditioning (Peretz, 1960; McCleary, 1961; Thomas &
Slotnick, 1962; Lubar & Perachio, 1965; Kimble & Gostnell, 1968;
Trafton et al., 1969).
A number of relatively complex deficits have been noted after
cingulate damage which resemble symptoms seen in human amnesia
syndromes. These include disruption of the orderlysequencing of
behaviors (in nest building and maternal behavior, Stamm, 1955;
Slotnick, 1967), deficits in the temporal ordering of responding


40
RAS impulses to accomplish localized (phasic) arousal of specific
areas of the brain. The ascending pathways of the RAS project
rostrally from the brainstem reticular formation (via the central
tegmental tract/medial forebrain bundle) to the hypothalamus, septal
area, and nonspecific intralaminar thalamic nuclei (ILTN). The
second path extends from the interpeduncular nucleus to the ILTN
via the habenula. The only direct ascending connections from the
RAS to the neocortex are projections from the nonspecific thalamic
nuclei (midline and ILTN) to the orbitofrontal cortex (via the
ventral anterior nucleus of the thalamus). Descending influences
are conveyed from the prefrontal neocortex to lower structures by
way of the thalamocortical radiations, corticoreticular fibers,
medial forebrain bundle, and thalamotegmental fibers. Hippocampal
output reaches the reticular formation via the fornix, mammillary
bodies, and mammillotegmental tract. The septal area has an addi
tional connection with the reticular formation by way of a stria
terminalis--habenul a--habenul ointerpeduncular tract--interpeduncular
nucleus pathway (Noback & Demerest, 1972).
Based on their analysis of some 200 experiments, Pribram
and McGuinness (1975) outlined two major subsystems in the brain
stem which control cortical mobilization and identified separate
forebrain mechanisms which modulate their functioning. These
systems initiate two different types of cortical activity. Diffuse
cortical "arousal," which is associated with the orienting response,
is based on the serotonergic brainstem median raphe' nuclei located
in the core of the reticular formation. Arousal is modulated by


12
simultaneous synthesis of information which permits the mental manipu
lation of the relationship between information units and as such is
a prerequisite for the high level mental functions that are charac
teristic of human beings (Luria, 1973a).
At the anterior pole of the brain, contiguous with the secondary
motor association areas, the prefrontal lobes (areas 9-12) are
dramatically enlarged and now represent up to one-fourth of the entire
cortical mass. The prefrontal lobes have extensive two-way connections
with all other parts of the cerebral cortex (Luria, 1973a). The
coordinating and control operations carried out by the lower level
(motor) systems in this hierarchy are evident in the functions of
the tertiary integration areas. The frontal lobes have been called
the "executive of the brain" (Pribram, 1973). They are the seat of
Luria1 s "unit for programming, regulation and verification of
activity" (Luria, 1973a). Lesions of the frontal lobes lead to
a defect in the patient's "capacity for planned initiative"
(Penfield & Evans, 1940), and to disturbances on impulse control
(Pribram, 1973).
Subcortical Systems
The limbic system consists of a group of interconnected
structures, situated between the midbrain and the neocortical
mantle, including the hypothalamus, amygdala, hippocampus, septal
area, and cingulate cortex (see Fig. 2). Because of their relation
ship to the olfactory bulbs, these structures were originally
thought to be concerned with that function and so this area of the
brain was designated the rhinecephalon ("nose-brain"). Although


103
7. In obsessive-compulsive illness certain thoughts and/or
actions acquire an inappropriate level of significance for the
individual and social functioning is impaired. Cingulectomy is
the psychosurgical "operation of choice for the classical syndrome
of obsessive neurosis;" after surgery "the obsessive thoughts
retreat gradually into the background and the drive behind them
lessens" (Lewin, 1961). Humans who have undergone cingulectomy
show some disinhibition of behavior (but decreased aggressiveness)
and seem to neglect the emotional impact of external events
(Whitty, 1966).
Discussion
It is evident that the cingulate cortex is involved in the
coordination of emotional, attentional and cognitive processes.
It appears that this structure modulates the infusion of emotional
significance into ongoing perceptual and/or associational operations
and influences the allocation of neural mobilizing energy, thus
permitting normal emotional reactivity and appropriate inhibition
of behavioral and cognitive activity. While it is not possible at
present to sort out the complexities of 5-HT/NE interactions
across brainstem, cortical, and right/left dimensions, the available
evidence suggests that the cingulate gyrus evolved as a cortical
extension of those brainstem control mechanisms. In the context
of functional systems, the cingulate cortex is in a position to
encode emotional data from the amygdala-orbitofrontal circuit into
neural models being "indexed" by the hippocampal memory system.


95
normal responses to actual pain. Arbuthnott, Fuxe, and Ungerstedt
(1971) presented evidence that all of the "positive reinforcing"
(self-stimulation) points are within the ventral NE system. Stein
and Wise (1969) demonstrated (with permanently implanted push-pull
cannulae) that "rewarding" self-stimulation of the MFB caused
increased synthesis and turnover of NE and a marked increase in the
release of radioactively labeled ME into perfusates of the amygdala
and hypothalamus.
Isaacson (1974) suggested that the almost total'lack of fear
and aggression resDonses in animals after bilateral amygdalectomy
reflects a decrease in the ability of enviromental stimuli to
elicit reactions from the organism. Increased 5-HT activity has
been associated with increased fear and decreased aggression while
decreased NE activity has been associated with decreased fear and
increased aggressiveness. It appears that in lower forms these
systems mediate the release or suppression of instinctive behavior
at the level of the hypothalamus. It follows from all of the data
presented above that the ability to learn from experience, a
prerequisite for self-determined behavior, involves an integration
of emotional experience and memory which is based on an interaction
of NE and 5-HT in the amygdala.
Emotion, Amygdala Circuits, and Memory
The role of the amygdala in emotional processes, established
by Kluver and Buey in 1933, has been assumed to be affected through
this structure's close relationship with the hypothalamus. The
amygdala seems to direct behavior toward biological goals (Halgren,


31
Efron argued that this was because the "conscious comparison"
of the two flashes takes place only in the hemisphere dominant for
language, the time lag representing the extra neural steps involved
in relaying sensory information from the right hemisphere over the
corpus callosum:
It is only after sensory data have reached the left hemi
sphere that one is "conscious" of the occurrence of an
event. ... To be conscious of something is to be conscious
of something now. It is the thesis of this paper that
the "now" is the moment of arrival of sensory data in
the dominant temporal lobe. (1963b, p. 421)
The most convincing evidence of a correlation between human
consciousness and language ability emerged from studies of a
unique cerebral commissurotomy patient known as "case P.S."
P.S., a right handed boy, developed epilepsy following an injury
to his left hemisphere incurred at age two. He subsequently
developed language skills in both his right and left hemispheres.
At age 14 he underwent complete surgical section of his corpus
callosum to relieve his epilepsy. Following surgery it was found
that P.S.'s right hemisphere could spell, comprehend verbal commands,
process parts of speech and make conceptual judgements involving
verbal information. It was also discovered that his right hemisphere,
although unable to speak, could generate answers to printed ques
tions presented tachistocopically to his left visual half-fields.
He accomplished this by arranging Scrabble letters with his left
hand. These answers were often different from those given
verbally by his isolated left hemisphere. Gazzaniga and Ledoux
(1977) argued that P.S.'s right hemisphere possessed qualities
deserving of conscious status because


55
for a particular kind of reinforcement, or class of reinforcements,
will generalize from one situation to another" (Rotter, 1975, p. 57).
Rotter distinguished two types of "generalized expectancies" (GE).
The first has to do with the nature of the reinforcement: expectations
for a particular kind of reinforcement in a given situation. The
second type deals with other properties of situational stimuli and
has to do with the perception of control that one can exercise to
change or maintain the situation: the kind of behavior that is
likely to produce or terminate reinforcement. The first type is
designated with a subscript _r for reinforcement (GEr). The second
type is designated a problem-solving generalized expectancy (GEps).
Striking insights into the nature and mechanics of this sort of
experiential memory were afforded by Penfield's observations of
certain psychical phenomena elicited by direct electrical stimulation
of the conscious brain (see Mullan & Penfield, 1959).
Wilder Penfield1s data were collected from patients undergoing
radical brain surgery, with local anesthesia, for the relief of intrac
table epilepsy. His observations consist of spontaneous reports from
these patients following applications of a mild electric current to
the exposed cortex from the tip of a unipolar electrode. The responses
to such stimulation which are of interest here fall into three
categories:
1. The emergence in consciousness of vivid and coherent
experiential hallucinations which appeared to be recollections of
(or abstractions from) the subject's past experiences.
2. Changes in a patient's subjective experience of his or her
relationship with the immediate environment.


66
seems probable that in such cases a large part of the
increase in permanent neural representation which is
assumed to correlate with familiarity will take place
in the language centers, that is, in the dominant
hemisphere. (Kimura, 1963, p. 269)
Thus, unfamiliar stimuli, which are not represented in verbal
memory by a permanent neural model (e.g., a name or concept) are
more likely to be processed by the right hemisphere, which Kimura
suggests is more important than the left in the establishment of
such "cell assemblies." Since all of the material in the memory store
are initially unfamiliar it follows that many, if not all, verbal
concepts (in the left hemisphere) might be based on neural models,
or gestaltan, which were assembled (and are stored) in the right
hemisphere. Such an hypothesis is supported by evidence from
split-brain studies that the right hemisphere is far superior to
the left in the discrimination of part-whole relationships (Nebes,
1974).
The appearance of material-specific amnesia syndromes following
unilateral temporal lobectomies suggests that the isolation of
language in the left hemisphere extends to verbal learning also, and
is thus virtually complete. Milner (1971) reports that
A comparison of left and right anterior temporal lobec
tomy in epileptic patients has revealed certain specific
memory defects that vary with the side of the lesion.
These material-specific disorders are to be distinguished
from the global amnesia that follows bilateral damage
in the hippocampal zone (Milner, 1958). Thus, left
temporal lobectomy, in the dominant hemisphere for
speech, selectively impairs the learning and retention
of verbal material (Meyer & Yates, 1955; Milner, 1958).
. . Conversely, removal of the right, nondominant
temporal lobe leaves verbal memory intact but impairs
the recognition and recall of visual and auditory patterns


Differences in MMPI Scores 157
Age 159
V DISCUSSION 160
APPENDICES
A HYPO- AND HYPER-DOMINANCE SPECTRUM DISORDERS BY
DSM III DIAGNOSTIC CLASSIFICATION 165
B INFORMED CONSENT STATEMENT 166
C REVISED SCORING CRITERIA FOR THE STREET (1931)
GESTALT COMPLETION TEST 167
BIBLIOGRAPHY 168
BIOGRAPHICAL SKETCH 188

vi


53
memory phenomena in psychoanalytic literature (e.g., slips of the
tongue, forgetting, false remembering, repression) can be traced
to this seminal paper in which the authors concluded that "hysterics
suffer mainly from reminiscences." In this work, Breuer and Freud
made a crucial distinction regarding the memory processes operating
in psychoneurosis which may have sowed the seed from which the
notion of unconscious causation of psychological phenomena germinated:
. . the causal relation between the determining psychical
trauma Ian experience which calls up distressing affects
such as those of fright, anxiety, shame or physical pain]
and the hysterical phenomenon is not of a kind implying
that the trauma merely acts like an agent provocateur
in releasing the symptom, which thereafter leads an
independent existence. We must presume rather that the
psychical traumaor more precisely the memory of the
traumaacts like a foreign body which long after its
entry must continue to be regarded as an agent that
is still at work. (Breuer & Freud, 1974, p. 355)
The authors became aware of this "highly remarkable phenomenon"
and its relation to affective processes in the course of their
experimental treatment of hysterical conversion symptoms:
[We found] that each individual hysterical symptom
immediately and permanently disappeared when we had
succeeded in bringing clearly to light the memory of
the event by which it was provoked and in arousing
its accompanying affect, and when the patient had
described that event in the greatest possible detail
and had put the affect into words. Recollection
without affect almost invariably produces no result.
(Breuer & Freud, 1974, p. 355)
Hillix and Marx (1974) have suggested that "it was necessary for
Freud to invent the psychic apparatus and much of his psycho
analytic theory just to account for what he and Breuer had already
observed" (p. 352). It is to be hoped that recent evidence will
make a more parsimonious accounting possible.


62
Levels of processing in memory are the subject of a theory
(summarized by Gaffen, 1972) which is based on arguments by Tall and
(1965) and supported by experimental evidence (Peterson, 1967;
Kintsch, 1970). Briefly, the theory postulates that the process
of recall consists of two separate and autonomous stages: retrieval
(or search) and recognition. The retrieval process "proceeds at
several levels . each being terminated by an implicit act of
recognition" (Talland, 1965, p. 304). The recognition stage is
based on a record from which the past cannot be read directly,
but which can assign a particular response in a particular context
a value of "familiarity--unfamiliarity" (the correct response being
the most familiar in that context). Thus "[in the retrieval stage]
various responses are generated (but not emitted); when finally the
correct response is generated, it is recognized as such by the
recognition stage, and is then emitted" (Gaffen, 1972, p. 328).
The theory postulates that amnesic subjects (animal and human)
lack the faculty of discriminating familiarity. This basic deficit
is manifested in the premature termination of search cycles, resulting
in an incorrect match. These formulations are not inconsistent
with those of Butters and Cermack (1974), who concluded from their
experiments that increased sensitivity to proactive interference,
subsequent to inappropriate encoding of information, was the
critical factor underlying the amnesic disorder. Finally, it is
interesting to note that modern theories regarding amnesia seem
to have arrived at the point at which they began: Korsakoff (1889),
in keeping with the associationist doctrine of his time, believed


93
inputs either genetically or experientially endowed with significance
to the organism. The emotional state would be accompanied by the
"intercerbral release of a trophic neurogenic substance . .
[which would permit] transcription into more permanent form . .
to present and immediately preceding states of neuronal activation
where the outcome had been significant for the organism" (Kety,
1972, p. 71).
Simply put, the results of the organism's adaptive efforts (i.e.,
positive or negative reinforcement) might produce a characteristic
biochemical signature within these circuits which reflected the
consequent affective status of the organism in that situation. These
data could then be encoded along with the memory of the situational
context. If that same stimulus configuration was confronted again
at a later time the encoded emotional information would cause the
biochemical pattern to be replicated in these circuits, the effect
of which would be to recreate the original emotional state to
motivate the organism appropriately based on the original experience.
Extensive work by Stein and his colleagues indicates that the
circuits involved include a noradrenergic median forebrain bundle
reward/behavior-release system which is antagonized by the seroto
nergic periventricular system (PVS) which functions as a punishment/
behavior-suppression mechanism. Stein, Wise and Berger (1972)
summarize experimental evidence which implicates these systems in
the release and suppression of unlearned behavior:
Briefly, electrical stimulation of the median fore
brain bundle . elicits species-typical consummatory
responses, such as feeding and copulation, which


77
a Korsakoff-type of memory disorder were thought to have had their
fornices damaged by metal fragments. Four of the five cases with
typical amnesic syndromes in Jarho's (1973) study of brain-injured war
veterans were considered to have bilateral interruption of the fornix
or mammillothalamic tract.
Sweet, Talland, and Ervin (1959) reported the case of a woman
in whom the anterior columns of the fornix were sectioned to facilitate
the removal of a colloid cyst of the third ventricle. This patient
showed a rapid recovery of old skills, but developed a "severe loss
of memory for recent events [which persisted at two years] ... a
retrograde amnesia of at least several weeks and a subjective
complaint of amnesia for specific events of four or five years past"
(p. 76), coupled with intact remote memory. In the discussion
following the presentation of this patient, Brenda Milner reported
on a similar case operated on by Welsh in 1954. This patient showed
a gross initial memory disorder but, at one year, was able (with
effort) to effect some compensation for his defect: "Whenever he
deliberately sought associative links he was able to improve his
performance considerable" (Sweet et al., 1959, p. 79). Milner
concluded that
I think that one can only account for the paradoxical
diversity of data from the fornix cases, as contrasted
with the consistent and severe memory loss in the hippo
campal cases (and maybe in the mammillary body cases
also), by supposing that you are only interfering
with a part of the system by fornix section. Thus,
you are most apt to see a temporary disruption of dis
turbance of memory with minimal residual loss. (p. 79)


54
The implicitly verbal form of memory referred to by Rapaport
seems to be qualitatively different from the "foreign body" which
Breuer and Freud assumed to be the culprit in hysterical neurosis.
They referred to the latter type of memory by the less formal term
"idea" and indicated that symptom removal depends on the transforma
tion of this "idea" into a more formal thought process so that its
associated affect can be abreacted:
[the therapeutic procedure] brings to an end the opera
tive force of the idea which was not abreacted in the
first instance, by allowing its strangulated affect to
find a way out through speech; and it subjects it to
associative correction by introducing it into normal
consciousness, (p. 356).
This special type of memory would seem to merit a more detailed
description. It is evident that we are concerned here with a
subset of memories which have significanee for the individual.
By definition, these are memories that are associated with reinforce
ment and/or emotional experience. They are experiential (nonverbal)
and may be isolated from consciousness. It may be noted that this
subset of memories will define the relationship between the individual
and his or her environment and might be the organism's most important
survival resource. A concept from social learning theory seems to
encompass this type of memory comfortably and provides a more
operational definition.
Julian Rotter (1966) theorized that "a reinforcement acts to
strengthen an expectancy that a particular behavior will be followed
by that reinforcement in the future" (p. 2). Further, "when an
organism perceives two situations as similar, then his expectations


27
embraced by laymen and professionals alike. They have been cited
to suDport all manner of theories concerning psychological, philo
sophical, and spiritual dualities (see the critique by Kinsbourne,
1982). However, a closer analysis of the data cited by Bogan
suggests that ushc conclusions are misleading.
Michael Gazzaniga, an author of the pioneering animal studies
referred to by Bogan as perhaps the most dedicated and prolific of
the "split-brain" researchers, complained about the "overpopulari
zation" of basic data produced by himself and his colleagues:
These popular psychological interpretations of "mind
left: and "mind right" are not only erronious: they
are inhibitory and blinding to the new students of
behavior who believe classic styles of mental activity
break down along simple hemispheric lines. (1977, p. 416)
There is no doubt that one cerebral hemisphere can, in the absence of
its counterpart, support high level intellectual activity if the
loss of the other hemisphere occurs early in development. Griffith
and Davidson (1966), for example, report that children show relatively
good recovery from hemispherectomy for infantile hemiplegia.
Smith and Sugar (1975) renorted on a 26 year old man who showed
superior intelligence (WAIS VIQ:126, PIQ:102, FSIQ:116) 21 years
after undergoing left hemispherectomy at age five and one-half.
However, it is improper to infer normal functioning directly from
grossly abnormal cases such as this, or from animal studies. While
it is clear that the right hemisphere may have the capacity to
develop higher mental processes, there is no evidence that it does
so normally, and considerable evidence to the contrary. Research


15
In primitive organisms this brain produces a repertoire of instinctive,
stereotyped behaviors which are sufficient to insure the survival of
the individual and species. These unlearned, species-specific
behavior patterns relate to the elementary functions of obtaining
food and shelter, establishing and defending a home territory,
breeding, maternal behavior, etc. (Isaacson, 1974). Programs for
these behavior patterns are apparently stored in the brainstem
and triggered by the hypothalamus: complex behavior sequences such
as eating, drinking, sexual acts, aggression and many other types of
unlearned behavior can be elicited by electrical stimulation of the
hypothalamus at levels below those needed to activate motivational
systems (Isaacson, 1974). In protoreptilian animals the performance
of these acts would be coordinated by the basal ganglia. In these
forms the striatum is the highest center for sensory and motor
processing, these functions being subserved by structures corresponding
to the caudate/putamin and globus palladus, respectively (Schade'
& Ford, 1973).
Stimulus and response are yoked in the protoreptilian brain.
Since its repertoire of behaviors is unlearned (i.e., instinctive)
this neural system "reacts to changes in the environment by increasing
or decreasing the intensity of the predominant response sequence.
. . Suppression of response on the basis of non-reward or punishment
is difficult" (Isaacson, 1974, p. 273).
The development of the paleomammalian brain allows the suppres
sion of stereotyped ways of responding. The addition of the limbic
system circuitry permits the organism to adjust its behavior based


CHAPTER I
INTRODUCTION
The various schools of psychotherapy agree that the practitioner's
task is to facilitate change in the client. They agree on little else.
There is, as yet, no consensus regarding the two major issues in
psychotherapy: what is to be changed, and how that change is to be
brought about (Strupp, 1978). Strong opinions about both of these
basic problems are available; data based conclusions are not. Pro
ponents of fundamentally different viewpoints debate acrimoniously
(Eysenck, 1974); yet verifiable differences in outcome between theory-
based forms of intervention are rare (Bergin & Lambert, 1978).
This unsatisfactory state of affairs in applied psychology has un
fortunate consequences for practitioners and their clients alike.
The problem has been ascribed by Watson to the "preparadigmatic"
state of the discipline of psychology. According to Watson "psychology
has not experienced anything comparable to what atomic theory has done
for biology, what laws of motion have done for physics. Either
psychology's first paradigm has not been discovered yet or it has
not been recognized for what it is" (Watson, 1967, p. 53). Hanson
stated the problem succinctly; "The issue is not theory using, but
theory finding" (1965, p. 3).
The two major forces in psychological thought, psychoanalysis
and behaviorism, have encountered significant problems. Theories
1


61
represents a failure to transfer sensory impressions into long-term
store. The adequacy of the consolidation hypothesis is called into
question, however, by demonstrations that amnesic patients are in
fact able to recall new information under certain conditions. The
most convincing evidence comes from experiments using the technique
of "cued recall" in which a subject is given partial information
about a stimulus (e.g., a previously presented word or picture)
and asked to identify the whole item. Under these circumstances the
performance of amnesic subjects was not significantly different
from that of controls (Weiskrantz & Warrington, 1970). This
suggested to the authors that the amnesic deficit involved problems
with mechanisms of retrieval rather than those of acquisition or
retention.
Weiskrantz (1979) reviewed a number of experimental paradigms
in which normal learning has been demonstrated in amnesic subjects
and underscored the fact that, in each case, the patients themselves
persistently failed to acknowledge the fact that their performance
was based on specific past experience (or that they had been
confronted with the task before). Thus, amnesia victims do not have
access to their memories on a conscious level, nor is such awareness
necessary for that memory to be demonstrated objectively. Weiskrantz
pointed out that this "striking dissociation between the subjects'
commentaries and their objective performance . suggests a
dissociation between levels of processing rather than a failure on
any particular level" (p. 385).


76
continues to arch forward, following the course of the lateral
ventricle to the rostral edge of the thalamus. Here the bundles
separate again to form the anterior columns of the fornix which
curve downward in front of the intraventricular foramen and above
the anterior commissure (AC). Approximately half of the fibers
descend behind the AC as the postcommissural fornix; the other half,
in a less compact bundle, pass in front of the AC as the precommissural
fornix. Postcommissural fibers pass through the hypothalamus to
the mammillary bodies, giving off fibers to the thalamus on their
way. Some of the precommissural fibers distribute to the septal
area and others join with septal fibers and continue into the same
areas as the postcommissural fornix. Approximately one-third of the
fornix fibers reach the mammillary bodies (Daitz, 1953); the anterior
nuclei of the thalamus receive as many direct fibers from the fornix
as they do from the mammillothalamic tract (Truax & Carpenter, 1969,
ch. 21).
Liss (1968), working with the rat, found that hippocampal
and fornix lesions had analogous effects on learning and behavior
in passive and active-avoidance tasks. In the monkey, fornix
lesions led to impaired learning of a spatial reversal task that
was "functionally similar" to the deficit seen after hippocampal
removal (Mahut, 1972; Mahut & Zola, 1973). Gaffen (1972) reported
a series of six experiments in which rats with fornix lesions were
shown to have a defect in "recognition memory" that the author
argued was eguivalent to anterograde amnesia in humans. In Russel's
(1971) survey of brain wound cases, the eight patients who showed


75
would add emotional coloring to psychic processes
occurring elsewhere. (Papez, 1937, pp. 304-306)
Papez believed that sensory input to the system originated in the
thalamus and was communicated via the subthalamus to the hypothalamus.
The circuit is completed with the connection of the cingulate
gyrus to the hippocampus by way of the cingulum bundle.
It is now known that other limbic structures are more actively
involved in the specifically emotional processes. However, Papez's
anatomical concepts might be rehabilitated if their context were
changed from emotion to memory. Input to the hippocampal circuits
would be seen as processed sensory information and its output as
memory indexing information capable of "cueing" associations and
generalized expectations related to the input stimulus. It remains
to place the role of the hippocampus in the context of the functional
system it subserves and to examine the other components of that
system.
Fornix. Hippocampal output makes its way via direct and
indirect pathways to the thalamus (anterior, dorsomedial, and intra
laminar nuclei), the hypothalamus, and the midbrain reticular
formation. Its main efferent fiber system, the fornix, is composed
of axons from pyramidal cells in the body of the hippocampus.
These fibers converge in the fimbria, traveling backwards within the
temporal lobe, and then arch forward under the corpus callosum as
the crura (posterior pillars) of the fornix. Here a number of
a
fibers cross to the other side, forming the hippocampal commissure.
The two crura then join to form the body of the fornix which


157
differed significantly (p < 0.001) from patients assigned to the
hypo-dominant group, who scored relatively better on tests sensitive
sensitive to right, relative to left, ehmisphere cognitive functioning,
as predicted by the proposed model.
Overall Performance on the Tests Sensitive to
Right versus Left Hemisphere Cognitive Functioning
Hypothesis 2 suggested that any differences discovered in the
test of Hypothesis 1 would be due only to differences in the ratios
between groups and not to any other pattern of overall performance
on the tests (e.g., intelligence). To test this hypothesis,
standardized scores (z-scores based on the total n) were derived for
each subject's individual test scores (following Robson, 1973) and
added togehter for each group (Street + Object Assembly +
Similarities + Information). The mean z-score was x = 0.09 for
the hypo-dominant group and x = -0.04 for the hyper-dominant group.
A student's t-test for independent samples (Robson, 1973) performing
on these means revealed no significant differences between groups
(t = 0.87^gg, p<0.4, two-tailed, N.S.). It was concluded that
there were no significant differences on overall test performance
between groups and that the differences in ratios found in the
test of Hypothesis 1 were not due to any factor (e.g., intelligence)
other than the one being tested.
Differences in MMPI Scores
Hypothesis 3 amounted to a verification of the subjects' diagnoses
and provided an additional test of the ability of the proposed
model to predict membership in diagnostic categories based on an
assessment of personality traits. Specifically, it was hypothesized


139
functioning the system which has greater access to the motivating and
mobilizing system would have the greatest impact on cognitive and
behavioral responding. This conclusion is perhaps best exemplified
in the psychological sequelae of unilateral temporal lobe epilepsy.
The Psychopathological Correlates of Unilateral Temporal Lobe
Epilepsy
In all members of the phylum, biologically important experiences
elicit an emotional resonse which facilitates the learning of
biologically important behavior (Campbell, 1974). Destruction of the
amygdala, or disconnection of the amygdala from the sensory cortices,
results in the dissociation of affective qualities from sensory
stimuli (e.g., Kluver & Buey, 1938). Sensory-limbic association
depends on a neural pathway which extends to the amygdala via the
ventral temporal cortex (Bear, 1979).
Patients with temporal lobe epilepsy often show progressive
personality changes which culminate in patterns which are indis
tinguishable from certain psychodiagnostic entities. Bear (1979)
reviewed evidence suggesting that these changes are produced by
chronic over-stimulation of the amygdala. In contrast to the
consequences of temporo-1imbic disconnection, he postulated that
the alternations in behavior, emotion, and thought observed in this
disease stem from a process of temporo-1imbic hyperconnection:
a "progressive overinvestment of perception and thought with affective
significance" (p. 359).
Bear and Fedio (1977) compared the self-reported and observer
ratings of personality traits in patients with left and right temporal
lobe epileptic foci. Factor analysis revealed two factors which


122
association areas, responsible for an increasingly complex synthesis
of incoming information, becomes specialized to perform different
tasks in different ways. The mode of information processing in the
two hemispheres begins to diverge as speech comes to dominate the
functional organization of association areas on the left, leaving
the high level integration of nonverbal sensory input to their
counterparts on the right. In the course of ontogenetic development
the tertiary zones within each hemisphere begin to control the work
of the secondary areas, which become subordinate to them (Luria,
1973a, ch. 2). In this manner the basic perceptual processes are
altered to conform to the needs of the higher centers. The secondary
association areas on the left become specialized to perceive semantic
data which can be coded into logical systems. The left tertiary area
analyzes this input sequentially, abstracting relevant details and
associating these with verbal symbols (see Nebes, 1977). The secon
dary areas on the right specialize in analyzing concrete structural
data and relationships which are organized into complex wholes by
the tertiary association area on that side.
Evidence (to be detailed below) indicates that during develop
ment these two increasingly specialized cortical information
processing systems (each with its own subjacent memory, emotion,
and arousal subsystems) become organized into a single functional
meta-system in which the cortical components are yoked together by
the limbic system. This meta-system optimizes the utilization of
its complementary components in the service of producing adaptive
behavior and assuring the survival of the species. Within this


161
and considered response generation), it is convenient to express
these two forms of pathology in terms of hyper- or hyDo-dominance.
Any given pathological syndrome will reflect both the point of
A
disturbance within the meta-system and the attempts of the rest of
the system to compensate, but most may be classified as hyper-
or hypo-dominance spectrum disorders with the primary symptoms
manifesting themselves either intra- or extrapsychically, resoectively.
A major consequence of hyper- or hypo-dominance is the avail
ability of right hemisphere cognitive products to the left for
utilization in response generation. The validity of the instruments
used as dependent measures in this study is well established and
their sensitivity to lateralized cognitive processes has been
adequately documented. The relationship between these indices of
lateralized brain functions and personality traits (as measured by
the relevant clinical scales of the MMPI) was clearly demonstrated
and congruent with the predictions of the theory. The agreement
between the MMPI data and the diagnoses used to assign subjects to
the experimental groups also enhances the value of the results.
A number of other factors contribute to the validity and general -
izability of the results in this study. Considerable design risks
were undertaken in that variables were introduced which would have
invalidated the results had they not conformed to predictions.
The assignment of mentally disordered sex offenders exclusively to
the hypo-dominant group and general psychiatric forensic inpatients
to only the hyper-dominant groups could not have been justified if
the two experimental groups had differed in overall performance on the


180
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Moruzzi, G., & Magoun, H.W. Brainstem Reticular Formation and
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Nebes, R. D. Superiority of the Minor Hemisphere in Commissurotomized
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152
Subjects
A total of 42 adult psychiatric patients served as subjects in
this study. Twenty outpatients were taken from an urban community
mental health center and three rural satellite clinics. Twenty-two
inpatients were taken from a state forensic hospital. Subjects were
assigned to the hyper- or hypo-dominant groups by DSM III Axis I and
II diagnoses (see Appendix A).
The hypo-dominant group (N=22) consisted of 13 males and nine
females. Eighteen were right-handed, one left-handed and three
ambidextrous by self-report. The mean age was 30.45 years with a
standard deviation of 12.14 years. The diagnoses within this
group included 14 antisocial personality disorder (eleven of whom
were classified as mentally disordered sex offenders undergoing
inpatient treatment at the state forensic hospital); three histrionic
personality disorder; two dependent personality disorder; one
avoidant personality disorder; one narcissistic personality disorder;
one psychogenic Dain disorder.
The hyper-dominant groun (N=20) consisted of 14 males and six
females. Ten were right-handed, one left-handed and three ambidextrous
by self-report. The mean age was 42.65 years with a standard
deviation of 13.73 years. The diagnoses within this grouo included
11 schizophrenia, paranoid types (all of whom were adjudicated
incompetent to stand trial and undergoing inpatient treatment at
the state forensic hospital); two generalized anxiety disorder;
two compulsive personality disorder; three dysthymic disorder;
one simple phobia; one agoraphobia with panic attacks.


155
scoring criteria for two items were altered for this study. (The
complete socring criteria used are presented in Appendix C.)
Hypotheses
The following hypotheses were generated:
1. The ratio of the sum of the raw scores on the Street Gestalt
Completion Test and the WAIS-R Object Assembly Subtest to the
QAIS-R Similarities and Information Subtests (Street + Object
Assembly/Similarities + Information ) will differ between groups
with the hyper-dominant grouD showing relatively better performance
on the Similarities and Information Tests and the hypo-dominant
group showing relatively better performance on the Street and Object
Assembly Tests.
2. The sum of the standardized scores (z-scores) on all the
tests (Street + Object Assembly + Similarities + Information) will
not differ between the groups.
3. The ratio of the sum of the MMPI T-scores for scales
three and four to the sum of the T-scores for scales six and
seven (3T + 4T/6T + 7T) will differ significantly between groups
with the hyper-dominant group showing relatively higher scores on
scales six and seven and the hypo-dominant groups showing relatively
higher scores on scales three and four.


145
Anxiety Disorders, Obsessive-Compulsive Illness, and Paranoia
On the WAIS, a verbal I.Q. that is significantly higher than
performance I.Q. is especially characteristic of patients with
anxiety neurosis (see evidence summaried by Ogden, 1967). This
relationship indicates a relative predominance of left hemispheric
processes in this disorder. Tucker and his colleagues (1978)
reported two experiments which linked anxiety with left hemisphere
overactivation and dysfunction. Subjects reporting high anxiety
showed performance decrements on tasks lateralized (via the visual
half-fields) to the left, but not the right, hemisphere. High
trait anxiety was also associated with a right-ear attentional bias
and a low incidence of left lateral eye movements (Tucker, Antes,
Stenslie & Barnhardt, 1978).
Anxiety patients show lower than normal CNV amplitude. The CNV
develops slowly and irregularly in acute anxiety states, extinguishes
rapidly during deconditioning, and reappears slowly, if at all,
on reconditioning trials (Cohen, 1974). Since the CNV has been
related to the dopamine-mediated activation system, the elevated
levels of psychological activity and reduced cognitive efficiency
seen in anxiety states appears to be related to excessive serotonin-
mediated arousal processes in the left hemisphere. By contrast,
obsessive-compulsive subjects have an exaggerated CNV amplitude which
shows slow resolution, less decrement with partial reinforcement, and
fails to habituate (Cohen, 1974). Thus, the excessive (verbal)
cognitive activity which is characteristic in these individuals
seems to be related to overactivation of the dooamine-mediated


98
cue showed a decrease in latency on the second retention test as compared
to the first, and thus resembled the groups that never received a cue.
By contrast, animals which received hippocampal stimulation following
the reminder cue exhibited an increase in latency, similar to the
control animals which benefited from the reminder FS. Kessner (1981)
concluded from these results that "amygdala stimulation disrupted the
encoding of negative affective attributes associated with the reminder
FS negating its efficacy in enhancing subsequent retention. In
contrast, hippocampal stimulation had only a small effect and did
not totally disrupt the efficacy of the FS reminder" (p. 336). It
seems, therefore, that the amygdala is selectively involved in the
memory of emotional experience, at least for the negative affect
associated with passive-avoidance learning situation.
Kessner (1981) reviewed several studies which suggested that
the amygdala was not involved in the encoding of positive affective
attributes. However, he cites evidence that amygdala stimulation
does appear to disrupt retention when there is a magnitude of reward
effect (i.e., fewer errors for the expectation of a greater magnitude
of reward). Kessner concluded from the studies reviewed above that
"the amygdala encodes, stores, and retrieves both positive and
negative emotional attributes of a specific memory, providing the
input is of sufficient intensity to elicit a relatively strong
emotional reaction. . Different neural systems (e.g., hippo
campus) would encode other attributes (e.g., enviromental context)
of the same specific memory" (1981, p. 340).


140
differentiated between the two groups. On an emotive-ideative
dimension, patients with right temporal foci were distinguished by
externally demonstrated affect while the left group was characterized
by ideational/ruminative traits. On a normal-severe dimension the
left group of patients reported more severity compared to the right
and endorsed more socially disapproved traits (e.g., paranoia,
aggression, dependence) but the rater evaluated the right group
as more severely disturbed. Bear (1979) observed that both groups
displayed a characteristic high intensity of affect but differed
in verbal awareness of the emotions. The major difference between
the groups was in the way they interpreted and reported their abnormal
affective experience. Bear suggested that "strong affect expressed
as mood excess differs from cognitive elaboration, often verbal or
logical, of specific relationships between stimuli and affect"
(1979, p. 370). This difference might account for the fact that
the right hemisphere epileptic foci have been associated with mood-
affective disorders and left foci with cognitive-paranoid psychosis
(Flor-Henry, 1974). Bear and Fedio (1977) speculated about the
ways in which a patient might interpret the enhanced affective
associations to previously neutral events or concepts which result
from chronic temporo-limbic hyperconnection:
Experiencing objects and events shot through with affec
tive coloration engenders a mystically religious world
view if a patient's immediate actions and thoughts are so
cathected, the result is an augmented sense of personal
destiny. A felt significance behind events that others
dismiss constitutes a seed bed for paranoia or may
confirm the feeling that the patient is a passive
pawn in the hands of powerful forces that structure
the world. Feeling fervently about rules and laws


68
Discussion
The evidence reviewed thus far suggests that the central problem
in the amnesia syndromes involves mechanisms which normally facili
tate access to stored memory traces. The problem might occur at
the point of encoding and deposition, or retrieval, or both.
The hippocampus appears to be critical to these coding and decoding
operations; this structure may normally provide the memory "cues"
which must be externally administered in its absence.
The evidence is supportive of Niesser's (1967) proposal that
(rather than tapping a "continuous memory strip'1) "Penfield's
electrode may have touched on the mechanisms of perceptual synthesis"
(p. 169). There can be little doubt that Penfield's "final common
pathways" in the temporal lobes are related to hippocampal afferents.
However, it is now clear that the hippocampus cannot be the only
secondary ganglionic station which must be activated before a signal
indicating familiarity, originating in the right hemisphere, can
"rise into consciousness." Studies of split-brain patients have
shown conclusively that the ability to give a verbal account of
events occurring in the right hemisphere is dependent on the
integrity of the cerebral commissures (e.g., Sperry, 1968).
The general layout of the commissures is such that a specific area
in one hemisphere is connected via commissural fibers to the homolo
gous area in the contralateral hemisphere. The temporal lobes have
their own private interconnection in the anterior commissure, which
also connects the two amygdalae (Gray, 1977). Studies of patients
with only partial sectioning of these commissures have indicated


FUNCTIONAL BRAIN SYSTEMS AND PERSONALITY DYNAMICS
By
DAVID LINDQUIST
A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN
PARTIAL FULFILLMENT OF THE REQUIREMENTS
FOR THE DEGREE OF DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA

Copyright 1985
by
David Lindquist

ACKNOWLEDGEMENTS
I am deeply grateful to Max and Ruth Lindquist for their patience
and the opportunities they gave me. My debt to my undergraduate
mentor, Dr. Carol Van Hartesveldt, who taught me about science and
let me make my own mistakes, is also gratefully acknowledged.
I want to thank my doctoral committee members, Doctors Grater,
Morgan, Schauble, Suchman and Ziller, who were also my most respected
graduate teachers. Special thanks are due to David Suchman, who
first encouraged me in this project, and to my chairman, Harry Grater,
who shepherded me through the crises with grace and allowed me to
keeD my dignity.
The help of Dr. Bill Froming in the design of this study, and of
Mssrs. Denny Gies, Bill Baxter and Ted Shaw (of the North Florida
Evaluation and Treatment Center) and of Ms. Janet Despard (of Mental
Health Services, Inc.) in facilitating its execution, is much
appreciated. I am especially grateful to Ms. Cheryl Shaw, who typed
the manuscript, and without whose friendship and organizational help
this project would not have been completed.
Warmest heartfelt gratitude goes to my dear friends Gabriel
Rodriguez, Marshall and Laura Knudson, and David Kurtzman, whose
love sustained me through these difficult years.
I can never properly express my thanks to Ruth Lindquist, to whom
this piece of work is lovingly dedicated.
iii

TABLE OF CONTENTS
ACKNOWLEDGEMENTS iii
LIST OF FIGURES vii
ABSTRACT viii
CHAPTER
I INTRODUCTION 1
II THE PHYSIOLOGICAL SUBSTRATE OF PERSONALITY:
A SELECTIVE REVIEW OF THE LITERATURE 7
Review of Basic Brain Anatoniy and Organization 8
Cortical Mechanisms 8
Subcortical Systems 12
The Interpretation of Neurology Literature 17
Human Consciousness 18
Right versus Left 18
Language and the Left Hemisphere 20
Aphasia: Anatoniy and Syndromes 21
Language, Symbolism, and Meaning 24
Human Consciousness, Self-Awareness,
and Thought 26
Discussion 35
Cortical Mobilization: Attention, Arousal,
and Activation 39
The Reticular Activating System and
Tonic Arousal 39
Phasic Control Systems: The Frontal Lobes
and Thalamus 41
Discussion 44
Motivation: Emotion and Affect 45
Amygdala Circuits and the Prefrontal Lobes 46
Affective Expression 48
Discussion 50
Memory Functions 51
Human Amnesia Syndromes 57
Memory and the Neocortex 63
Discussion 68
The Limbic System, RAS, and Memory 70
Papez Circuit and Memory 74
Fornix 75
iv

Mammillary bodies and mammillothalamic tract..80
Cingulate cortex 81
Discussion 83
Interfaces and Interactions of the Monitoring,
Motivating, and Mobilization Systems 89
The Biochemistry of Emotion, Motivation,
and Learning 92
Emotion, Amygdala Circuits and Memory 95
Discussion 103
Biochemical and Electrophysiological Aspects
of Cortical Mobilization Processes 104
The Functional Elements of the Personality
Structure 108
The Problem-Solving/Response-Generatinq
System 109
The Memory System 110
The Motivating System 112
The Mobilization System 114
Learning and Memory: Animal Studies 115
A Functional Meta-system 120
Lateralized Mobilization Processes 125
Asymmetrical reaction time to laterally
presented stimuli 125
Altered GSR following unilateral brain
injury 126
Asyrmietrical biochemical and electro-
physiological processes 127
Bilateral Interaction in Emotion and Cognition..128
Mental Health and Psychopatholgy 136
The Psychopathological Correlates of
Unilateral Temporal Lobe Epilepsy 139
Discussion: Hyper- and Hypo-dominance
Spectrum Disorders 141
Schizophrenia and the Affective Disorders 143
Anxiety Disorders, Obsessive-Compulsive
Illness, and Paranoia 145
Sociopathy and Hysteria 147
III METHOD 151
Subjects 152
Instruments 153
Procedure 154
Hypotheses 155
IV RESULTS 156
Left versus Right Hemisphere Cognitive
Functioning Between Groups 156
Overall Performance on the Tests Sensitive to
Right versus Left Hemisphere Cognitive
Functioning 157
v

Differences in MMPI Scores 157
Age 159
V DISCUSSION 160
APPENDICES
A HYPO- AND HYPER-DOMINANCE SPECTRUM DISORDERS BY
DSM III DIAGNOSTIC CLASSIFICATION 165
B INFORMED CONSENT STATEMENT 166
C REVISED SCORING CRITERIA FOR THE STREET (1931)
GESTALT COMPLETION TEST 167
BIBLIOGRAPHY 168
BIOGRAPHICAL SKETCH 188

vi

LIST OF FIGURES
FIGURES
1 Cytoarchitectural map of the lateral and medial
surfaces of the human cerebral cortex, with numbers
representing the areas of Brodman 10
2 Partially schematized representation of the
limbic system 13
3 The position of Papez's circuit within a
larger cortical circuit 86
4 Schematic representation of the proposed model of
the physiological substrate of personality Ill

Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy
FUNCTIONAL BRAIN SYSTEMS AND PERSONALITY DYNAMICS
By
David Lindquist
May 1985
Chairman: Dr. Harry Grater
Major Department: Psychology
A heuristic model of the physiological substrate of personality
structure was developed from a review of recent neurological and physio
logical psychology literature. The formulation of the model was based
on assumptions that the phylogenetic transition from instinctive
responding to self-determined behavior required the evolution of auto
matic neural mechanisms to insure that the organism would monitor the
environment for significant stimuli, be motivated to respond in
the presence of those stimuli, and mobilize the appropriate psycho
logical operations to determine the form of that response. Functional
elements of these basic mechanisms were identified and a functional
meta-system was outlined which would organize the elements and
optimize the utilization of lateralized cognitive processes in the
interest of assuring the emission of adaptive behavior.
The proposed model suggests that certain psychodiagnostic entities
might be classified as hyper- and hypo-dominance spectrum disorders
vi 1 i

based on the form of dysfunction within the meta-system. The ability
of the model to predict membership in diagnostic categories was
tested by assigning 42 adult psychiatric inpatient and outpatient
subjects to hyper-dominant and hypo-dominant groups by diagnosis,
according to the constructs of the model, and comparing performance
on instruments shown to be sensitive to right and left cerebral hemi
sphere dysfunction. The Street Gestalt Completion Test and the Object
Assembly, Similarities, and Information Subtests of the Wechsler Adult
Intelligence Scale-Revised were administered to each subject. An
abbreviated form of the Minnesota Multi phasic Personality Inventory
(MMPI) was used to compare symptomology between groups.
Significant between-group differences (p < 0.001) in the ratios
of test scores sensitive to right versus left cognitive functioning
were found in the predicted directions, while the groups did not differ
in overall performance on the instruments. Significant differences
(p < 0.005) in the ratios of selected MMPI clinical scales, in the
predicted direction, provided further support for the hypothesized
relationship between lateralized cognitive functioning, symptomology,
and diagnosis.
It was concluded that the proposed model orovides tenable and
potentially useful operational definitions of personality functions
and psychopathology. Results were discussed in terms of their impli
cations for psychotherapeutic interventions and additional methods
to test the validity of the model.
IX

CHAPTER I
INTRODUCTION
The various schools of psychotherapy agree that the practitioner's
task is to facilitate change in the client. They agree on little else.
There is, as yet, no consensus regarding the two major issues in
psychotherapy: what is to be changed, and how that change is to be
brought about (Strupp, 1978). Strong opinions about both of these
basic problems are available; data based conclusions are not. Pro
ponents of fundamentally different viewpoints debate acrimoniously
(Eysenck, 1974); yet verifiable differences in outcome between theory-
based forms of intervention are rare (Bergin & Lambert, 1978).
This unsatisfactory state of affairs in applied psychology has un
fortunate consequences for practitioners and their clients alike.
The problem has been ascribed by Watson to the "preparadigmatic"
state of the discipline of psychology. According to Watson "psychology
has not experienced anything comparable to what atomic theory has done
for biology, what laws of motion have done for physics. Either
psychology's first paradigm has not been discovered yet or it has
not been recognized for what it is" (Watson, 1967, p. 53). Hanson
stated the problem succinctly; "The issue is not theory using, but
theory finding" (1965, p. 3).
The two major forces in psychological thought, psychoanalysis
and behaviorism, have encountered significant problems. Theories
1

2
based on the former have been criticized as untestable and therefore
unscientific (Eysenck, 1970). The latter movement lost impetus with
the discovery by Olds and Milner (1954) of "pleasure" centers in the
brain. This revelation undermined the basic assumption of the learning
theorists that behavior could be explained simply by defining the
rules governing stimulus-response relationships.
Successful scientific theories are built on paradigms that
describe the fundamental properties and mechanics of their subject.
The fundamental units of personality are networks of neurons in the
brain. Sigmund Freud (1948) attempted to relate mental structures
to anatomical locations but was forced to abandon his effort because
the neurology of the time was not adequate. Instead he and subsequent
theorists were forced to base their models on suppositions about
the products of the personality processes. As noted above, the
results have been less than satisfactory. The science of neurology
has made significant progress in the interim and a large amount of
useful information has accumulated. These data have been virtually
ignored by the discipline of psychology. The integration of neuro
logical data and psychological theory may provide a basis for a useful
paradigm for the psychotherapist. The present work is intended as
a step toward such an integration.
The purpose of this study is to develop and test a heuristic
model of personality function, based on an understanding of its
physiological substrate, with the ultimate goal of imoroving the
effectiveness of psychotherapeutic interventions. Such a model
should identify the basic elements and processes of the personality

3
structure and describe the ways in which these interact to produce
psychological health and psychopathology. Such a model might lead to
new operational definitions of psychological phenomena which, in
turn, may suggest new intervention points and methods.
The purpose of a theory is to integrate known facts within a
single framework and account for them in terms of a small number of
interrelated concepts. Existing theories suffer from a lack of
integration. The discipline of psychology has hindered such integra
tion by institutionalizing a tradition of subspecialization: theories
of attention, perception, cognition, motivation, emotion, behavior,
etc. are developed in relative isolation. The failure of individual
theorists to understand and to appreciate the interrelatedness of
these various personality operations may account for the inadequacy
of psychological theory in general. It is assumed here that the
component parts of the personality can only be properly understood
in the context of their relationship to the whole; that more will
be gained from a gross (but comprehensive and testable) description
of the personality infrastructure than from a detailed examination
of a single personality operation.
Rather than subdivide the personality structure on the basis of
notions of what it should do; it might be more productive to approach
the problem on the more basic level of what it must do. The formula
tion of the model will be guided by a set of assumptions about the
evolutionary pressures which shaped the personality structure.
These "evolutionary imperatives" are as follows:

4
1. The physiological substrate of personality evolved to
assure the emission of adaptive behavior by the organism. Survival
pressures shaped an organization that was able to respond effectively
to significant stimuli and produce behavior that enhanced the
organism's well-being.
2. The evolution of this substrate proceeded from an organiza
tion based on "hard-wired" stimulus-response instincts to an organiza
tion which allowed increased latitude in behavior in order to take
advantage of the organism's developing problem-solving abilities.
3. In order to permit self-determination of behavior and,
at the same time, to insure survival, new mechanisms were required
to assure that the organism would (a) attend to significant
stimuli; (b) be motivated to respond effectively to those stimuli;
(c) accomplish the mobilization of the necessary psychological
resources to determine the form of that response (referred to
hereafter as the Monitoring, Motivating, and Mobilization systems).
4. Although these systems interact, they must be functionally
separated to the extent that they do not interfere with each other's
normal operation. Similar functions might be carried out in other
brain areas, but the automatic activation of these systems will
assure that they dominate responding to stimuli which relate to the
survival and well-being of the organism.
5. As these mechanisms are essential for the survival of
the individual and species they will form the central organizing
processes of personality; personality dynamics will center on their
operation.

5
Neurological theories of mental function center on clinical
observations of patients with localized brain lesions. The history
of neurological thought has revolved around the question of how these
data are to be interpreted. For many years higher mental functions
were treated as discrete "faculties." The results were inconsistent
and of little value. The failures of these "narrow localizers" led
to theories which attempted to account for mental functions on the
basis of the "mass action" of the brain. Where earlier models were
too specific, these theories proved too general to be useful.
In the 1920s Goldstein broke with the tradition of attempting to
infer functions directly from deficits and proposed instead an
"analysis of basic disturbances." This approach led finally to
Luria's conceptualization of mental activities as the product of the
interaction of complex functional systems. In Luria's formulation
a mental function is the result of contributions from a number of
concertedly working zones. Therefore, that function may be destroyed,
or disturbed differently, by lesions in different locations. Luria
(1973a) described the characteristics of a functional system:
The presence of a constant (invariant) task, performed
by variable (variative) mechanisms, bringing the process
to a constant (invariant) result, is one of the basic
features distinguishing the work of every "functional
system." The second distinguishing feature is the
complex composition of the "functional system," which
always includes a series of afferent (adjusting) and
efferent (effector) impulses. (Luria, 1973a, p. 28)
Luria outlined three principal functional units in the brain:
the "units for regulating tone and waking and mental states,"
v t
centered on the reticular activating system in the brainstem; the

6
"unit for receiving, analyzing and storing information," operating
in the post-central (sensory) areas of the brain; and the "unit
for programming, regulation and verification of activity," operating
in the frontal lobes (Luria, 1973a, ch. 2).
Luria's concepts represent a major advance in the understanding
of the fundamental operating characteristics of brain systems.
Although his formulations are too basic to be of much use to
the applied psychologist, he has established a format and a methodol
ogy which will be followed here. The present investigation will
focus on identifying and describing the interactions of the functional
brain systems which satisfy the requirements of the evolutionary
imperatives outlined above. Evidence suggests that the substrate
for these systems will be found in those anatomical areas for which
Luria acknowledged he had inadequate data for his own analyses: the
medio-basal zones of the cortex and the right hemisphere of the brain.

CHAPTER II
THE PHYSIOLOGICAL SUBSTRATE OF PERSONALITY:
A SELECTIVE REVIEW OF THE LITERATURE
In the following sections the neurological and physiological
psychology literature pertaining to the functional/anatomical
organization of personality will be reviewed. The data will be
related to psychological factors and any conclusions will be noted
in discussions at the end of each section. The first section is a
review of basic brain anatomy and organization. The second section
will examine the physiological substrata of consciousness and conclude
that human consciousness, characterized by self-awareness, is a
manifestation of processes which occur in the left hemisphere of
the brain. The third section will trace the systems involved in
cortical activation and note the existence of two mechanisms within
each hemisphere which have opposite effects on the form of cognitive
processes. The fourth section will examine the role of the aniygdala
and frontal lobes in the subjective experience of emotion and of
the right hemisphere in the expression of affect. The fifth section
will develop the basis for a theory of memory function and the role
of memory systems in organizing affective, arousal, and cognitive
processes. The sixth section will focus on the interactions of the
emotion, arousal, and memory systems and examine the role of bio
chemically mediated systems in coordinating these processes. In
the seventh section a model of four functional systems which
7

8
constitute the basic elements of the personality structure will be
proposed. The eighth section will describe the way in which the
basic elements are organized into a functional meta-system which
forms the infrastructure of personality and functions to assure the
emission of adaptive behavior. This model will be supported with
evidence concerning the psychological correlates of neurological
syndromes. In the final section the psychological phenomena
associated with various forms of psychopathology will be related
to neurological indices which reflect the operation of the lateralized
subsystems and their interaction within the functional meta-system.
It will be concluded that many psychodiagnostic entities may be
classified as hypo- or hyper-dominance spectrum disorders which
are functionally related to chronic, and maladaptive, under- or
over-utilization of the mechanisms in the left hemisphere of the
brain.
Review of Basic Brain Anatomy and Organization
Cortical Mechanisms
Although certain phylogenetically new areas of the cerebral
cortex are of special interest when discussing "higher mental
functions," these areas must be considered in their physiological
and evolutionary context. A brief review of basic brain systems
and anatomy will establish this perspective.
In man, as in all mammals, large portions of the cerebral
cortex are devoted to the more elementary functions of processing
sensory stimuli and the initiation and control of movements. The
brain structures subserving these basic functions are divided at the

9
central fissure (Rolando). Incoming somato-sensory, visual and
auditory nerve impulses are relayed, via the thalamus, from contra
lateral receptor surfaces to primary projection areas located in
the parietal, occipital and temporal lobes (see Fig. 1, areas 1, 2,
3; 41; 17), respectively (Noback & Demerest, 1972). In each case the
modally specific, somatotopic organization of nerve impulses in
the projection areas is transformed into functional information
(i.e., acquires meaning) in an adjacent secondary association
area (areas 5, 7, 18, 19; 42). With each new level of processing
there is increasingly complex synthesis of information and decreased
modal specificity (Luria, 1973a). Damage to a primary Drojection
area results in a loss of sensation (e.g., blindness) while lesions
of a secondary association area are likely to produce the inability
to recognize a stimulus in that modality (agnosia). Conversely,
artificial stimulation of a projection area produces a discrete
sensory experience while stimulation of a secondary association
area elicits a more elaborate sensory hallucination whose complexity
is related to the level within the hierarchy that is activated
(see Mullan & Penfield, 1959).
Progression within the hierarchy is reversed in the motor
systems. Specificity of control increases as the secondary (pre
motor) areas (areas 6 and 8) coordinate and fine tune their influence
on the pyramidal cells of the primary motor cortex (area 4) with
the assistance of continuous feedback from the sensory modalities.
Lesions of the primary motor cortex produce contralateral paresis
while stimulation elicits flexion of individual muscle groups.

10
Figure 1. Cytoarchitectural map of the lateral and medial
surfaces of the human cerebral cortex, with numbers repre
senting the areas of Brodman. (Redrawn from Noback & Demerest,
1972).

11
Stimulation of secondary motor areas produces smooth, coordinated
movements and ablation of these areas may result in the loss of
ability to perform skilled motor acts (apraxia) (Noback & Demerest,
1972).
The systems described thus far are typical of all mammals and
become progressively more elaborate and efficient in the higher
primates. Evolution proceeds generally by modifying and elaborating
existing hardware, allocating new functions to tissues which are in
some way pre-adapted to assume the new tasks (Campbell, 1974).
The development of higher mental functions in man is correlated with
bilateral anatomical expansion of two cortical areas which are
adjacent to the secondary association areas described above. These
regions subserve the highest level of organization in the hierarchies
and are called teritary association areas (Luria, 1973a). Both
areas are involved in what Penfield (1975) referred to as "trans
actions of the mind."
The inferior parietal lobule (I PL) (areas 23; 39, 40) lies at
the anatomical confluence of the secondary association areas in the
post-central cortex. This area is the "association cortex of
association cortexes" (Geshwind, 1979). Here processed information
from the surrounding sensory association areas is further integrated
and synthesized. The area is called "supramodal" because its
individual units can only be excited by the simultaneous stimulation
of two or more sensory modes (Luria, 1973a). Information processing
at this level is "abstract" in that it is independent of a particular
sensory modality (cf. Osgood, 1953). This ability allows the

12
simultaneous synthesis of information which permits the mental manipu
lation of the relationship between information units and as such is
a prerequisite for the high level mental functions that are charac
teristic of human beings (Luria, 1973a).
At the anterior pole of the brain, contiguous with the secondary
motor association areas, the prefrontal lobes (areas 9-12) are
dramatically enlarged and now represent up to one-fourth of the entire
cortical mass. The prefrontal lobes have extensive two-way connections
with all other parts of the cerebral cortex (Luria, 1973a). The
coordinating and control operations carried out by the lower level
(motor) systems in this hierarchy are evident in the functions of
the tertiary integration areas. The frontal lobes have been called
the "executive of the brain" (Pribram, 1973). They are the seat of
Luria1 s "unit for programming, regulation and verification of
activity" (Luria, 1973a). Lesions of the frontal lobes lead to
a defect in the patient's "capacity for planned initiative"
(Penfield & Evans, 1940), and to disturbances on impulse control
(Pribram, 1973).
Subcortical Systems
The limbic system consists of a group of interconnected
structures, situated between the midbrain and the neocortical
mantle, including the hypothalamus, amygdala, hippocampus, septal
area, and cingulate cortex (see Fig. 2). Because of their relation
ship to the olfactory bulbs, these structures were originally
thought to be concerned with that function and so this area of the
brain was designated the rhinecephalon ("nose-brain"). Although

Figure 2. Partially schematized representation of the limbic system. FR CTX frontal cortex;
CING cingulate cortex; TEMP CTX temporal cortex; CC corpus callosum; SEPT septal area;
AMYG amygdala; HPC hippocampus; HT hypothalamus; MB mammillary bodies; ANT anterior
thalamic nuclei; HAB habenula; ILTN intralaminar thalamic nuclei; IP interpenduncular
nucleus; RF reticular formation; DBB diagonal band of Broca; MFB median forebrain bundle;
SM stria medularis; ST stria terminal is; FNX fornix. (Redrawn from Isaacson et al, 1973)

14
lower animals depend on the sense of smell for such survival-
related activites as food-getting, the detection of enemies, and
mating (MacLean, 1949), the role of these subcortical structures
in motivational and emotional processes was not appreciated until
the 1930s. Since Kluver and Buey (1938) published their description
of altered emotional behavior following the bilateral removal of
the temporal lobes, including the amygdalae and hippocampi, limbic
system components have been the subject of intense scrutiny. The
massive literature which has accumulated is filled with confusing
inconsistencies and conflicting findings. This may be due in part
to the fact that the function of these structures appears to vary
according to enviromental circumstances (Olds, 1958). Although
a number of tentative models of limbic system function have been
offered (e.g., Papez, 1937; Gloor, 1956; Olds, 1958) none has
found general acceptance.
MacLean (1970) distinguishes three major types of systems in
the mammalian brain which correspond to stages in its evolutionary
development. He describes a protoreptilian core-brain, a paleo-
mammalian brain (the limbic system) and a neomammalian brain (the
neocortical areas). This triune brain conceptualization provides
a useful perspective on the hierarchical arrangement of anatomical
and functional systems and their relationship to behavior (Isaacson,
1974).
The protoreptilian brain represents the fundamental core of
the nervous system consisting of areas homologous to parts of the
upper brainstem and midbrain, hypothalamus, and basal ganglia.

15
In primitive organisms this brain produces a repertoire of instinctive,
stereotyped behaviors which are sufficient to insure the survival of
the individual and species. These unlearned, species-specific
behavior patterns relate to the elementary functions of obtaining
food and shelter, establishing and defending a home territory,
breeding, maternal behavior, etc. (Isaacson, 1974). Programs for
these behavior patterns are apparently stored in the brainstem
and triggered by the hypothalamus: complex behavior sequences such
as eating, drinking, sexual acts, aggression and many other types of
unlearned behavior can be elicited by electrical stimulation of the
hypothalamus at levels below those needed to activate motivational
systems (Isaacson, 1974). In protoreptilian animals the performance
of these acts would be coordinated by the basal ganglia. In these
forms the striatum is the highest center for sensory and motor
processing, these functions being subserved by structures corresponding
to the caudate/putamin and globus palladus, respectively (Schade'
& Ford, 1973).
Stimulus and response are yoked in the protoreptilian brain.
Since its repertoire of behaviors is unlearned (i.e., instinctive)
this neural system "reacts to changes in the environment by increasing
or decreasing the intensity of the predominant response sequence.
. . Suppression of response on the basis of non-reward or punishment
is difficult" (Isaacson, 1974, p. 273).
The development of the paleomammalian brain allows the suppres
sion of stereotyped ways of responding. The addition of the limbic
system circuitry permits the organism to adjust its behavior based

16
on new sources of information about the internal and external
environments and to utilize new and more efficient forms of learning
in organizing its responding. It is important to appreciate that
the final product of limbic system operations in the paleomammalian
brain is the inhibition of lower centers. However, these same
mechanisms later came to exert important influences on the neo-
cortical systems.
The cerebral cortex of the neomammalian brain developed in close
association with the limbic system and basal ganglia. New nuclei were
added to existing sensory and motor systems, culminating finally
in the arrangement now found in primates. More efficient information
processing in the neocortical additions permits more precise sensory
discriminations and rapid, fine-grain movements of the extremities
(Isaacson, 1974) but these new systems still operate in close con
junction with the older subcortical mechanisms (Schade1 & Ford,
1973).
The importance of the limbic system in human experience can
hardly be overstated. These structures are implicated in most,
if not all, forms of psychopathology. They are the probable sub
strate for the therapeutic effect of most psychotropic medications
(Broekkamp & Lloyd, 1981) and the targets for all forms of "psychia
tric surgery." Unfortunately, existing theory regarding the
functional significance of the limbic system is primarily descriptive
in nature or consists of generalizations so broad as to be of
little use to the applied psychologist.

17
The Interpretation of Neurology Literature
The neurological data which form the basis of this review were
gathered over many decades, from a variety of populations, by
scientists with widely divergent theoretical beliefs. The psycholgist
who is unfamiliar with this literature must become aware of its
inherent problems before venturing interpretations based upon it.
Those who come under the neurologist's care have, almost
invariably, suffered a catastrophic change in their lives and
personalities. Much of the data is drawn from individuals who have
incurred brain damage as a result of a head wound, cerebrovascular
accident (stroke), or cerebral tumor (neoplasm). While offering an
otherwise unavailable opportunity to study higher brain functions,
these devastating "natural experiments" invariably preclude the
rigorous application of scientific method to that study.
Although techniques continue to improve, it is unusually
impossible to define the parameters of a lesion accurately. Thus,
crucial independent variables cannot be precisely isolated and
controlled. Cause and effect interpretations are further
hampered by the fact that a patient's pre-morbid level of functioning
may be impossible to ascertain. Generalizability is also compromised
when the subject has a chronic brain disease, such as epilepsy,
or has undergone a surgical intervention to control that condition,
such as temporal lobectomy or cerebral commissurotomy. In these
cases, as with patients who have been subjected to "psychiatric
surgery," it is reasonable to suggest that the "pre-treatment"
psychological organization may have been grossly abnormal.

18
The human brain has remarkable ability to compensate for
injury by utilizing undamaged tissue. This recovery of function
in the time elapsed between injury and testing is a further
confounding factor.
Much of this literature is in the form of case reports and the
problem of individual variability is especially difficult to manage.
Research designs utilizing groups and statistical analyses are
becoming more frequent, but these studies invariably suffer from
the difficulties noted above.
The most difficult problems to be encountered here have to do
with the specification of dependent variables: "higher brain func
tions" tend to resist definition and to defy quantification.
Generally, only gross operational definitions have been available.
The physician/experimentor was often forced to resort to a judgement
as to whether a given (and hypothetical) function was "intact"
or "lost," with the attendant risks of experimentar bias and error.
Related to this is the understanding, which emerged slowly in the
development of this literature, that the function of a circumscribed
anatomical area cannot be inferred directly from a deficit which
follows the ablation of that area: one can only specify how the
brain functions in the absence of that tissue.
Human Consciousness
Right versus Left
There have been three basic approaches to the study of lateralized
cortical function: the comparison of patients with unilateral brain
damage; the lateralized presentation of stimuli or task to the

19
sensory receptor fields of normal subjects; and the study of patients
whose cerebral hemispheres have been surgically separated (Nebes,
1977). The early studies of split-brain subjects prompted a great
deal of speculation about the abilities of the right hemisphere
and a listing of the published works concerned with asymmetrical
hemispheric functioning is beyond the scope of this paper. Evidence
pertaining to the differences between the cognitive operations
performed by the left and right hemispheres has been ably reviewed
by a number of writers (e.g., Bogan 1969a, 1969b; Galin, 1974, 1977;
Gazzaniga, 1970; Gazzaniga & Ledoux, 1977; Nebes, 1974, 1977;
Sperry, 1968). Some of the more important differences will be noted
very briefly here.
The primary deficits seen after right hemisphere lesions involve
difficulties in perceiving, manipulating, and remembering spatial
relationships and in perceiving and remembering sensory stimuli
which resist verbal description (Nebes, 1977). The right hemipshere
is superior to the left at generating a percept of the whole from
fragmentary information and seems predisposed to notice complex
gestalts or patterns rather than the parts (Nebes, 1971, 1974,
1977). The right hemisphere tends to process input simultaneously
(in parallel) as opposed to the left hemisphere's preference for
sequential (serial) processing, Cohen, 1973). In spite of many
claims to the contrary, the evidence indicates that the right
hemisphere considers only the immediately perceived context and
performs its tasks in a reflexive, automatic fashion.

20
Language and the Left Hemisphere
The prominent Soviet geneticist, Theodosius Dobzhansky, observed
that . while all other organisms become masters of their envi
ronments by changing their genes, man does so mostly by changing
his culture, which he acquires by learning and transmits by teaching"
(Dobzhansky, 1964, p. 145). Cultural evolution was made possible
by the development of language, but language is more than a special
form of social communication by which culture is transmitted: it
is the mechanism which produces the adaptive behavior patterns
that released the species from the constraints of biological
evolution. Speech is the fundamental tool of intellectual activity.
Language processes are essential in the operations of abstraction
and generalization, the basis of categorical thinking, and the
vehicle for organizing and regulating mental processes and behavior
(Luria, 1973).
The clearest example of functional brain asymmetry is the
lateralization of language to the left hemisphere, a fact long
established by two observations: deficits in language functions
(aphasia) following damage to the left hemisphere and the retention
of language following right hemisphere injury (right handedness is
assumed throughout the text unless otherwise noted). The patterns
of language deficit following circumscribed lesions provide the
most concrete evidence that is available about the mechanics of
consciousness and its product, cognition. The focus of the following
review is not on language itself, but on the evidence which indicates
that the active operations of consciousness which produce

21
adaptive behavior are part of a functional system that is lateralized
in the left hemisphere of the brain.
Aphasia: Anatomy and Syndromes
Three general types of aphasia have been defined and traced
to different anatomical structures in the left hemisphere. Through
an analysis of these syndromes it is possible to deduce the outline
of a functional system in which these separate areas work together
to produce the complex function of language. The following summary
is based on reviews by Gardner (1975) and Zaidel (1978).
Broca's aphasia, resulting from damage to the inferior, post
frontal zones of the left hemisphere, is an expressive disorder.
Utterances are difficult to initiate and speech is painfully slow
and labored. Patients are usually able to name objects and to repeat,
having less trouble with words that are familiar and concrete.
Although they manage to convey their meanings in a peculiar,
"telegraphic" style, they are unable to produce a fully formed
sentence. Their productions consist almost entirely of substantives;
grammatical parts and forms are absent or impoverished. Nouns are
usually delivered in the singular and verbs appear in their simplest,
noninflected form. Parts of speech that are purely grammatical in
function (conjunctions, prepositions, articles, adverbs) are exceeding
ly uncommon. The same deficit pattern is evident in the patient's
reading and writing.
Although patients with Broca's aphasia may have difficulty in
unravelling complex grammatical relationships (e.g,, "The lion
was eaten by the tiger: which animal is dead?"), their comprehen-

22
of language is generally intact. The ability to understand and
utilize nonverbal symbolism (e.g., gesture, pantomime) is also
spared. Intellectual functioning is relatively unimpaired. Patients
retain the ability to reason logically, to abstract and to generalize,
and to respond to context appropriately. Their associational
processes are not loosened, tangential or pressured. They do not
produce paraphasias or confabulate. Finally, these patients are
acutely aware of their deficits; they may be appropriately depressed
and are prone to sudden, transient emotional outbursts.
Exactly the opposite clinical picture is evident when Wernicke's
area, on the lateral, convex surfaces of the left temporal lobe, is
damaged. Wernicke's aphasia is characterized by impaired language
comprehension with fluently articulated but nonsensical speech.
Unlike the Broca's aphasic, words are spoken clearly with normal
sounding cadence, intonation and melody (prosody). However, the
speech of the Wernicke's aphasic is lacking in content and may
consist almost entirely of semantic jargon which has little communi
cation value. These patients appear to have lost control of the
language mechanisms at all levels. It is as if the selection
thresholds for phonemes, words, and ideas were all lowered.
Repetition is poor and marked by paraphasic errors in which the
correct sounds may be present but emerge in the wrong order.
Patients are able to name only the most familiar objects accurately,
although the word produced may come from the same category as the
target. Prompting with the initial wordsounds seldom helps.

23
In spontaneous speech the key substantives are often missing
and the remaining parts of speech lack their organizing influence.
Grammatical parts and forms are used abundantly but incorrectly.
Adjectives,, adverbs, conjunctions and prepositional phrases are
strung together haphazardly and the result may resemble a schizo
phrenic's "word salad" (Luria, 1973). Nonsense syllables and neo
logisms are frequent. The guiding thought behind a verbal produc
tion may be evident, but it becomes obscured by tangential associa
tions and incomprehensible babbling.
The Wernicke's aphasic understands little of what is said and
seems to rely on nonverbal cues in order to respond to a situation.
They are also unable to comprehend nonverbal symbolism and so their
communication is vague and concrete at all levels.
Perhaps the most striking feature of Wernicke's syndrome is
the patients.' complete lack of awareness of and indifference to their
deficits. They appear unconcerned and will vehemently deny any
problems.
The same deficits are evident in reading and writing tasks.
The patient can read single words but does not seem to grasp their
meaning or relate them to him or herself. They may correctly
repeat a simple command written on a card but will make no attempt
to comply. Unable to formulate an acceptable sentence on their
own, they are able to arrange cards with words printed on them to
form a syntactically correct sentence. However, the key substan
tives are likely to be misplaced (e.g., "The man bit the dog.")
confirming again that the patient's facility is with granmar as
opposed to meaning.

24
There is less agreement about the third major language
disorder, known as anomic or amnesic aphasia, which results from
more posterior lesions located in the angular gyrus in the parieto
occipital area. Part of the confusion may stem from the fact that
its main symptom, the loss of the ability to name objects, is
common to all aphasic disorders. However, the anomic aphasia
syndrome is distinguished by the fact that the naming disorder is
accompanied by relatively intact comprehension of written and spoken
language and normal spontaneous speech. The ability to read and
to repeat are also spared in anomic aphasia.
The anomic aphasic has no difficulty using words in their
appropriate context but cannot find the word in isolation of
context; he is unable to divorce himself from the immediate
situation. The anomic aphasic cannot produce the name of objects
on demand even though he knows what they are. When an object is
designated the patient is unable to produce its name, and conversely,
given a name, the patient is not certain what it refers to.
Other language difficulties are evident. The patient's
spontaneous speech seems to be either too detailed or too general.
Thinking is very concrete; the patient will interpret proverbs
literally. The patient is aware of the diabilities and will often
develop strategies to compensate for them.
Lanugage, Symbolism, and Meaning
The theory of the functional organization of language develooed
by Wernicke in 1885 is still generally accepted today. According
to this model the underlying structure of an utterance arises in

25
Wernicke's area. It is then transferred via large fibre bundles
(the arcuate fasciculus) to Broca's area where it evokes a
detailed program for vocalization which is governed by the rules
of grammar and syntax. The program developed in Broca's area, the
linear scheme of the sentence, is supplied to the adjacent face area
of the motor cortex which in turn drives the muscles which produce
the vocalization. Thus, the content of speech originates in
Wernicke's area and finds its form in Broca's area.
Wernicke's area is also essential for the comprehension of
language. Auditory stimuli are relayed from the Organ of Corti
to the primary auditory projection areas in Heschell's gyrus in the
left temporal lobe. At this point, as with the other sensory projec
tion areas, the information is somatotopically organized and retains
its modal specificity; to be understood it must be transferred
to the secondary auditory association area (Wernicke's area)
where the somatotopical organization is converted into a functional
organization (Luria, 1973a). Here, the fundamental phonemic
characteristics of language are isolated and identified. Processing
by Wernicke's area is essential for both the encoding and decoding
of meaning. An intact Wernicke's area is also essential for the
expression and comprehension of meaning through symbolic gesture
and pantomime (Goodglass & Kaplan, 1973; Gionotti & Lemmo, 1976).
It is interesting to note in this regard that "illusions of
interpretation" emerge in consciousness after electrical stimulation
of the temporal lobe in the right hemisphere but not after stimula
tion of any other brain area (Mullan & Penfield, 1959).

26
Human Consciousness, Self-Awareness, and Thought
In the 1960s the general belief in cerebral dominance gave
way to the idea of cerebral specialization due, in large part, to
the "split-brain" studies conducted on the cerebral commissurotomy
patients of Drs. Vogel and Bogan. In 1969 Bogan took the progression
a step further when he revived Wigan's (1844) notion of "the duality
of mind." Wigan (noting the anatomical duality of the brain,
autopsy findings of hemispheric atrophy in patients whose personality
was apparently intact, and introspective evidence of concurrent,
opposing trains of thought) argued that if one hemisphere can
sustain a mind, "it necessarily follows" that a man with two hemi
spheres must have two minds (p. 271). Bogan endorsed this concept
in the third of his influential "Other side of the Brain" papers
(Bogan, 1969a) and concluded that
Pending further evidence, I believe (with Wigan) that
each of us has two minds in one person. . Various
kinds of evidence, especially from hemispherectomy,
have made it clear that one hemisphere is sufficient
to sustain a personality or mind. We may then
conclude that the individual with two intact hemi
spheres has the capacity for two distinct minds.
This conclusion finds its experimental proof in
the split-brain animals who can be trained to per
ceive, consider, and act independently. (1969,
pp. 156-157)
The "dual mind" concept implies two relatively equal but
functionally independent entities which act as opposed forces in
the process of determining behavior. Bogan contributed the hypoth
esis that the two hemispheres utilize different "modes of thought"
in this process: "propositional" on the left and "appositional"
on the right (1969, p. 160)- These appealing ideas were enthusiastically

27
embraced by laymen and professionals alike. They have been cited
to suDport all manner of theories concerning psychological, philo
sophical, and spiritual dualities (see the critique by Kinsbourne,
1982). However, a closer analysis of the data cited by Bogan
suggests that ushc conclusions are misleading.
Michael Gazzaniga, an author of the pioneering animal studies
referred to by Bogan as perhaps the most dedicated and prolific of
the "split-brain" researchers, complained about the "overpopulari
zation" of basic data produced by himself and his colleagues:
These popular psychological interpretations of "mind
left: and "mind right" are not only erronious: they
are inhibitory and blinding to the new students of
behavior who believe classic styles of mental activity
break down along simple hemispheric lines. (1977, p. 416)
There is no doubt that one cerebral hemisphere can, in the absence of
its counterpart, support high level intellectual activity if the
loss of the other hemisphere occurs early in development. Griffith
and Davidson (1966), for example, report that children show relatively
good recovery from hemispherectomy for infantile hemiplegia.
Smith and Sugar (1975) renorted on a 26 year old man who showed
superior intelligence (WAIS VIQ:126, PIQ:102, FSIQ:116) 21 years
after undergoing left hemispherectomy at age five and one-half.
However, it is improper to infer normal functioning directly from
grossly abnormal cases such as this, or from animal studies. While
it is clear that the right hemisphere may have the capacity to
develop higher mental processes, there is no evidence that it does
so normally, and considerable evidence to the contrary. Research

28
efforts in this area have been hampered by the lack of adequate
operational definitions (for such phenomena as consciousness, cogni
tion, and thought) which distinguish the hither mental processes
of humans from the brain functions of lower forms.
Cerebral dominance for consciousness has been investigated
using the Wada carotid amobarbital test, a procedure developed to
localize language functions prior to neurosurgery. The Wada test
involves injecting sodium amobarbital into one common carotid
artery and results in the anesthetization of only the cerebral hemi
sphere on the side of the injection. Terzian (1964) reported an
absolute and immediate arrest of any communication, both verbal and
nonverbal, in the first thirty to sixty seconds after the injec
tion of the drug into the carotid artery of the dominant side
which he interpreted as a transient loss of consciousness.
Serafetinides and his co-workers reported similar results and
noted that the phenomenon rarely occurred following barbiturization
of the non-dominant hemisphere (Serafetinides, Driver & Hoare,
1964, 1965a, 1965b). They concluded that unconsciousness, and by
implication consciousness, is in general linked with the function
of the hemisphere dominant for speech (Serafetinides et al., 1965a).
Rosadini and Rossi (1967) attempted to replicate these findings
using more strictly operationalized definitions of consciousness.
In one group (48 cases) the criteria consisted of an "analysis of
the capacity of the patient to keep in contact with the examiner
through verbalizations or movements, to react to noxious stimuli

29
and to describe at the end of the examination what happened during
the examination itself" (p. 103). In a second group the criteria
for consciousness consisted of "a simple stimulus-resoonse test"
in which "the patients were instructed to work a switch held in
the hand ipsilateral to the intracarotid injection [i.e., the
hand controlled by the unanesthetized hemisphere] any time they
heard a given sound or saw a flash of light" (p. 103). Behavioral
and clinical events indicating unconsciousness were required to
last more than one minute in order to "permit their safe detection."
They found that 47 of their 69 cases did not meet their criteria
for unconsciousness and the 22 cases which did occurred in roughly
the same percentage following left and right injections; aphasia
occurred in 16 cases (15 left, one right); only five of the 21
cases (three left, two right) evaluated with the stimulus-response
test failed to operate the switch held in the hand controlled by
the unanesthetized hemisphere in response to the signals used.
Although their results were complicated by existing neuropathological
and cerebrovascular abnormalities, these authors concluded that
"the existence of a cerebral dominance for consciousness is not
supported" (p. 111). The usefulness of this study appears to
be severely limited by the criteria used to evaluate consciousness
in the unanesthetized hemisphere: a large number of animals of
various species have demonstrated the ability to "work a switch"
in response to visual and auditory stimuli and to respond to noxious
stimuli, but these lower forms are not considered conscious in the
same sense that humans are. The authors were looking for "the

30
occurrence of signs revealing the capacity of the subject to keep in
contact with the external world" (p. 103). They acknowledged that
"the suppression of expressive and receptive speech functions make
such a task quite difficult with the patients receiving barbiturate
in the dominant hemisphere" (p. 109). It seems that it is only
the appearance of aphasiz (after dominant hemisphere anesthetization)
that is specific to human consciousness in this study, and these
findings are consistent with those of Terzian and Serafetini des
et al.
Rosadini and Rossi did not report the results of their test of
subjects' ability to recall what had occurred during the Wada
procedure but this question was addressed directly in an experiment
reported by Gazzaniga (1977). This author found that "information
encoded while the left hemisphere was anesthetized was uninterpretable
by the verbal system when the left hemisphere returned to normal
functioning . when information is encoded by other than the
verbal system the person is not consciously aware of the information"
(p. 150).
Another approach to the localization of conscious awareness
involved an analysis of the temporal discrimination for simultaneity
when two visual stimuli were presented separately to the left and
right visual half-fields, separated by a very brief interval.
Efron (1963a; 1963b) found that normal right-handed subjects reported
that the two flashes occurred simu"!taneously only when the
light flashed in the left visual half-field was presented several
milliseconds earlier than the light flashed in the right visual field.

31
Efron argued that this was because the "conscious comparison"
of the two flashes takes place only in the hemisphere dominant for
language, the time lag representing the extra neural steps involved
in relaying sensory information from the right hemisphere over the
corpus callosum:
It is only after sensory data have reached the left hemi
sphere that one is "conscious" of the occurrence of an
event. ... To be conscious of something is to be conscious
of something now. It is the thesis of this paper that
the "now" is the moment of arrival of sensory data in
the dominant temporal lobe. (1963b, p. 421)
The most convincing evidence of a correlation between human
consciousness and language ability emerged from studies of a
unique cerebral commissurotomy patient known as "case P.S."
P.S., a right handed boy, developed epilepsy following an injury
to his left hemisphere incurred at age two. He subsequently
developed language skills in both his right and left hemispheres.
At age 14 he underwent complete surgical section of his corpus
callosum to relieve his epilepsy. Following surgery it was found
that P.S.'s right hemisphere could spell, comprehend verbal commands,
process parts of speech and make conceptual judgements involving
verbal information. It was also discovered that his right hemisphere,
although unable to speak, could generate answers to printed ques
tions presented tachistocopically to his left visual half-fields.
He accomplished this by arranging Scrabble letters with his left
hand. These answers were often different from those given
verbally by his isolated left hemisphere. Gazzaniga and Ledoux
(1977) argued that P.S.'s right hemisphere possessed qualities
deserving of conscious status because

32
His right hemisphere has a sense of self, for it knows
the name it collectively shares with the left. It has
feelings, for it can describe its mood. It has a sense
of who it likes and what it likes, for it can name its
favorite people and its favorite hobby. The right
hemisphere in P.S. also has a sense of the future, for
it knows what day tomorrow is. Furthermore, it has
goals and aspirations for the future, for it can name
its occupational choice. . The fact that this mute
half-brain could generate personal answers to ambiguous
and subjective questions demonstrates that in P.S. the
right hemisphere has its own independent response-
priority determining mechanisms, which is to say, its
own volitional control system, (pp. 143-145)
P.S. is the only split-brain patient with advanced language skills
in his right hemisphere and the only patient to demonstrate double
consciousness. Ledoux, Wilson and Gazzaniga (1977) stressed the
fact that "in all other patients, where linguistic sophistication
is lacking in the right hemisphere, so too is the evidence for
consciousness" (p. 420).
The capacity for speech and conceptual thought is clearly innate
in homo sapiens; only the symbols themselves must be learned
(Campbell, 1974). Recent evidence has clarified the anatomical
substrate of this genetically transmitted specialization. The
development of language capabilities in the human species is
correlated with the anatomical expansion and interconnection of
the association areas in the left hemisphere (Campbell, 1974). The
posterior area of the planum temporale, which forms a part of the
secondary auditory cortex (Wernicke's area) is significantly larger
on the left side (Geshwind & Levitsky, 1968. The enlargement of
this area can be explained in terms of its distinctive cellular
organization (Galaburda, LeMay, Kemper & Geshwind, 1978) and the

33
incomplete development of this cellular architecture has been
related to language dysfunction (see Geshwind, 1979).
In her exhaustive study of hundreds of brain injured war
veterans, Semmes (1967) discovered that elementary sensory and
motor capacities were focally represented in the left hemisphere and
diffusely represented in the right. She proposed that this difference
indicates the mechanism of hemispheric specialization: focal
organization favoring fine control and the integration of similar
units (e.g., manual skills and speech) and diffuse organization
favoring multimodal coordination (e.g., the various spatial
abilities).
Gazzaniga and Ledoux (1977) observed that nearly every demonstra
tion of a right hemisphere advantage in split-brain patients has
involved manipulo-spatial activities and concluded that
[This advantage] exists so long as manipulative activities
are involved in either the stimulus perceptions or the
response production. . The probable neural substrate of
these manipulo-spatial acts involves the inferior parietal
lobule of the right hemisphere in humans. In the left
hemisphere, however, linguistic functions occupy the
inferior parietal lobule. . The superior performance
of the right hemisphere of split-brain patients on such
tasks does not reflect the evolutionary specialization
of the right hemisphere, but instead represents the
price paid by the left hemisphere in acquiring language.
. . Our view is not that the right hemisphere is
specialized in some unique way in man. Rather, it
continues to do what it does elsewhere in the phyla.
(pp. 420-421, emphasis added)
Campbell (1974) noted that "spatial relationships involving
depth and distance may appear to be predominately spatial concepts,
but they are not of space but about space; of themselves they are
spaceless and concerned with pattern rather than place" (p. 337).

34
It appears, then, that the left hemisphere extracts meaning from
the relationship of individual parts to each other, while the
right gathers meaning from the pattern of the whole.
Bogan (1969b) proposed that the right "mind" utilized a
different mode of thought which he characterized as aopositional to
denote the ability to appose, or compare, information. He contrasted
this with the propositional mode utilized by the left hemisphere. A
number of investigators have distinguished similar dichotomies of
information processing style. Luria (1973a) spoke of narrative
versus relational processes. Gal in (1974) suggested that the right
hemisphere solved problems through a process of multiple converging
determinants as opposed to a left hemispheric style which utilized
a single causal chain. Sechenov (quoted by Luria, 1973a) postulated
that the human brain utilizes two forms of integrative activity:
organization into simultaneous and primarily spatial groups, and
into temporally organized successive series. This is consistent with
Spearman's conclusion that intelligence comprises two components: the
eduction of correlates used in analogical reasoning and the eduction
of relations, the basis of abstract reasoning (see McFie & Piercy,
1952). Campbell (1974) noted that "abstraction means escaDe from
the present . what distinguishes man from animals is the length of
time through which his consciousness extends (p. 335). Finally,
Bogan (1969b) observed that the most important distinction between
the left and right hemispheric modes might be "the extent to which
the linear concept of time participates in the ordering of
thought" (p. 160).

35
There is a clear consensus recognizing two modes of informa
tion processing. However, the ability to process information is
not necessarily a sufficient condition for consciousness. Although
the notion of right brain "thought" has gained wide currency, to
date, there has been no conclusive evidence that any cognitive opera
tion occurring in the right hemisphere is directly experienced in
consciousness. A possiblity not considered by any of the above
authors is that one mode might be an ancillary resource utilized
by the other. The data reviewed thus far suggests that one must be
very careful to avoid anthropomorphizing when attempting to describe
right brain processes. However, some inductive conclusions may be
drawn.
Discussion
It is evident that human consciousness is inexorably linked
with the abstract symbolic processes associated with language. Perhaps
the most universally accepted characteristic of human thought is
self-awareness. Self-awareness (and its product, the self-concept)
requires the abstraction and appreciation of defining features which
are consistent over time and situation. Only the left hemisphere,
with its temporal acuity, can consider and aopreciate changed or
conserved relationships in different conditions or contexts. Thus,
only the left hemisphere can define itself. The resulting self-
awareness provides a reference point for all of the memories, feelings,
intentions and thoughts that are collectively known as the- mind
and which allow the individual, thus defined, to interact intelligently
with the environment. Lacking the temporal organizing skills to

36
construct such a consistent frame of reference the right hemisphere
is bound to the immediate context with only the influences of the
physiological status of the organism (and the left hemisphere) to
guide its processes. Complex motivations, therefore, cannot exist
in the right hemisphere. Likewise, so-called "pictorial thinking,"
if temporally ordered and goal directed, must be organized by the
left hemisphere. Gal in (1974) suggested that the context bound,
egocentric and impulsive nature of right hemisphere congition
resembled Freud's notion of primary process thinking. Higher mental
processes in the right hemisphere almost certainly qualify as
cognitions (i.e., a way of knowing) and may account for the phenomenon
of intuition (knowledge without awareness of the process by which
it was gained). However, the term "thought" seems misleading and
"information processing" might be preferable. As noted above, there
is no direct evidence that mental events occurring in the right
hemisphere are directly experienced in the conscious left hemisphere;
one is left to ponder the question of whether a tree falling in the
right brain would make a sound if the left wasn't listening.
The restrictions outlined above are in no way inconsistent
with the demonstrated role of the human right hemisphere in the
analysis of emotional communications and the modulation of affective
expression (e.g., Ross & Mesulam, 1979). The brains of lower
forms (and, apparently, the right brain in humans) are primarily
concerned with neuronal signals which represent the survival needs
of the organism within the immediate environment. Interaction with

37
the social environment is critically important to survival throughout
the phylum. Campbell (1974) noted that animal vocalizations and
signals are "emitted only in the presence of the appropriate stimu
lus" (p. 349) and warned against equating these vocalizations with
human speech: "the signals . are generated or motivated by the
phenomenon of emotion, and find their neurological origin not in the
cortex but in the limbic system of the brain" (1974, p. 348). The
cortical organization of these functions in the right brain of humans
appears to mirror that of language in the left hemisphere with
comprehension and expression utilizing anatomical areas homologous to
Wernicke's area and Broca's area, respectively (Ross & Mesulam, 1979).
Right hemisphere responses might achieve direct expression in circum
stances where control by Broca's area in the left hemisphere is
impaired or attenuated, a case in point being the clearly enunciated
emotional exclamations of the frustrated Broca's aphasic. Similarly,
poorly defined and undifferentiated emotionally generated behavioral
impulses (e.g., approach, avoidance) might also achieve motor
expression in the absence of adequate left hemisphere control.
Hughlings Jackson (1864) suggested that if the "faculty of
expression" was proven to be lateralized in the left cerebral
hemisphere it would then be reasonable to expect that its corre
sponding opposite, perception, might be lateralized to the right.
Although the concept of mental "faculties" has given way to an
appreciation of complex functional systems, the role of the right
hemisphere within those systems might, in a broad sense, be said to
conform to Jackson's prediction. While the information processing

38
style of the right hemisphere is not suited to solving problems or
making decisions, it is uniquely qualified to perceive the quality of
significance (as defined by the individual's experience) in complex
environmental stimuli. The evidence suggests that the human right
hemisphere attends to the overall pattern of stimuli, searches out
("apposes") associations which are correlated with important
stimulus configurations and collates them into percepts that have
meaning for the organism. Its associational processes are unen
cumbered by rules of logic and its perceptions uninfluenced by
expectations.
The thrust of Bogan's (1969b)"dual mind" thesis was a reaction
against the traditional concept of hemispheric dominance which
relegated the right hemisphere to the role of an "automaton" or
reserve organ (e.g., Henschen, 1926; Strong & Elwyn, 1943). A
basic assumption in the present work, however, is that evolutionary
pressures reguired in the development of an automatic environment
monitoring system in order to permit the transition from instinctual
to self-determined behavior. It appears that evolution solved
this problem by taking advantage of the fact that the human central
nervous system contains two relatively autonomous brains which could
be yoked together by the limbic system. Within this configuration
the left hemisphere may be seen as a problem-solving and response
generating system and the right hemisphere might be said to
function as the repository and librarian of the individual's
reinforcement history.

39
Cortical Mobilization: Attention,
Arousal, and Activation
Consciousness and cognition become possible only when minimum
levels of cortical tone are attained. These tonic levels, reflected
in the desynchronized EEG pattern, permit sensory discriminations,
motor acts and other cognitive operations to take place. Once
activated, the cortex has the ability to make phasic modifications
of its level of activity and to voluntarily direct its attention.
The fundamental systems which govern cortical tone and attention,
however, are automatic and capable of overriding voluntary controls.
It is clear that these systems, with their ability to control the
level and content of consciousness, will exert a significant
influence on the personality structure.
The Reticular Activating System and Tonic Arousal
The mobilization of the cortex is accomplished by the brainstem
reticular formation (RF). This structure is a network of highly
interconnected neurons which adjusts its level of activation by
integrating input from the sensory pathways, limbic system struc
tures, and the neocortex. Impulses from the reticular formation
lower the activation thresholds of the neurons it projects to.
When the reticular formation is relaxed, cortical tone is lowered
and the organism sleeps (Moruzzi & Magoun, 1949).
The reticular activating system (RAS) regulates the state of
activation of the brain in two ways: the ascending reticular
activating system (ARAS) affects the brain diffusely and sets the
generalized (tonic) level of arousal; descending influences direct

40
RAS impulses to accomplish localized (phasic) arousal of specific
areas of the brain. The ascending pathways of the RAS project
rostrally from the brainstem reticular formation (via the central
tegmental tract/medial forebrain bundle) to the hypothalamus, septal
area, and nonspecific intralaminar thalamic nuclei (ILTN). The
second path extends from the interpeduncular nucleus to the ILTN
via the habenula. The only direct ascending connections from the
RAS to the neocortex are projections from the nonspecific thalamic
nuclei (midline and ILTN) to the orbitofrontal cortex (via the
ventral anterior nucleus of the thalamus). Descending influences
are conveyed from the prefrontal neocortex to lower structures by
way of the thalamocortical radiations, corticoreticular fibers,
medial forebrain bundle, and thalamotegmental fibers. Hippocampal
output reaches the reticular formation via the fornix, mammillary
bodies, and mammillotegmental tract. The septal area has an addi
tional connection with the reticular formation by way of a stria
terminalis--habenul a--habenul ointerpeduncular tract--interpeduncular
nucleus pathway (Noback & Demerest, 1972).
Based on their analysis of some 200 experiments, Pribram
and McGuinness (1975) outlined two major subsystems in the brain
stem which control cortical mobilization and identified separate
forebrain mechanisms which modulate their functioning. These
systems initiate two different types of cortical activity. Diffuse
cortical "arousal," which is associated with the orienting response,
is based on the serotonergic brainstem median raphe' nuclei located
in the core of the reticular formation. Arousal is modulated by

41
a lateral-frontal-anrygdala lateral-hypothalamus fcilitory circuit
and an inhibitory orbitofrontal--amygdalamedial-hypothalamus
circuit. "Activation"is an attention focusing process involved in
perceptual expectancies and motor readiness to respond. This system
is based on the locus ceruleus, in the periaqueductal gray, which
supplies norepinephrine to the forebrain. Activation was thought
to be modulated by the ancient motor control system in the basal
ganglia. Together, these systems provide for appropriate attending
to novel or significant stimuli and prepare the organism to respond
cognitively and behaviorally. The hippocampus was seen to integrate
the functioning of these systems and to exert ultimate control over
cortical mobilization through a mutually inhibitory relationship with
the reticular formation.
Phasic Control Systems: The Frontal Lobes and Thalamus
A novel (possibly significant) stimulus elicits an orienting
response (OR) from the organism. The psychological phenomena
associated with the OR are familiar to all who have had experience
with "things that go bump in the night." The complete orienting
reaction includes
The suppression of ongoing behavior, the orienting of
the body and receptor towards the new stimulus, changes
in the peripheral autonomic nervous system, and, perhaps
less obvious, preparations for associating the new stimulus
with memories from the past and expectancies of the
future. (Issacson, 1974, p. 110)
Pribram (1973) noted that the stimulus sampling aspects of the
orienting reSDonse differed from the processes necessary to register
a stimulus in awareness and memory (which must be accomplished

42
before the organism can habituate to a stimulus). In contrast
to the indiscriminate arousal associated with orienting, these
latter processes require the focusing of attention.
The mobilization of selective attention ("activation") appears
to be reflected in the contingent negative variation (CNV) or
"expectancy wave" (Tecce, 1970). The CNV is a special form of
cortical evoked response which consists of a spreading wave of
negative potential that appears whenever there is a contingent
relationship between two stimuli. Negativity develops when brain
tissue is maintaining a readiness for processing (Pribram &
McGuinness, 1975). Thus, the CNV appears whenever the organism
is expecting to perform a perceptual or motor act. The negativity
becomes abruptly positive when that act is executed (Walter, Cooper,
Aldridge, McCall urn, & Winter, 1964). High amplitude CNVs are related
to greater efficiency of perceptual and motor responses; concentra
tion facilitates the CNV while inattention, boredom, or fatigue
decrease it (Cohen, 1974). Elithorn et al. (1958) postulated
that frontal lobe injuries somehow damaged the mechanism under
lying anticipatory sets. It is interesting, in this light, that
the CNV generally appears in the prefrontal lobes and sweeps
posteriorally over the post-central cortex.
Patients with frontal lesions are unable to sustain their
attention. While intelligence, as measured by standardized tests,
may be unimpaired, these individuals are highly distractable and
cannot carry out purposeful activity which is normally directed by
intentions (Luria, 1973a). Luria pointed out that patients with

43
lesions of the temporal, parietal, or occipital lobes may have
sensory, orientation, or intellectual deficits, but their attention
and concentration remain sustained and directed by intentions.
Luria and his co-workers suggested that the frontal syndrome reflected
the loss of this selectivity (Luria, Homskaya, Blinkov, & Critchley,
1967). An understanding of the functioning of thalamic systems
suggests a mechanism by which the frontal lobes might select or
"recruit" psychological operations in the post-central cortex.
The thalami are a pair of egg-shaped masses located beneath the
cortex in the center of the cerebral hemispheres. The thalamus
is the final processing point for cortical input and the central
integration station of the nervous system. The brief review of
thalamic anatomy and functioning presented below is based on reviews
by Noback and Demerest (1972) and Chusid (1976).
The ventral half of the thalamus contains the specific relay
of nuclei of the sensory-motor systems. The nuclei of the dorsal
tier are association nuclei which have reciprocal connections with
the association areas of the post-central cortex and no sub
cortical connections; the dorsolateral and posterolateral nuclei
are interconneected with the parietal lobe, and the pulvinar with
the temporal and parietal lobe.
The dorsomedial and anterior thalamic nuclei are association
nuclei involved in emotion and memory, respectively. The dorso
medial nucleus receives input from the amygdala and lateral hypo
thalamus and has reciprocal connections with the association areas
of the prefrontal lobe. The anterior nuclei of the thalamus receive

44
the output from the hippocampus and have reciprocal connections
with the cingulate cortex.
Lying between and separating the major thalamic association
nuclei, the nonspecific (intralaminar, midline and reticulate)
nuclei have only intrathalamic and subcortical connections. They
receive their main input from the brainstem reticular formation and
from the rostral end of the ascending reticular activating system
(ARAS). The intralaminar nuclei have the "remarkable property of
being able to exert a controlling influence upon the rhythmic
electrical activity of the entire cortex" (Jasper, 1949, p. 406).
Jasper noted that this system is in a position to provide a central
coordinating mechanism for cerebral activities:
A central integrative mechanism with ready access to all
afferent and elaborative systems of both hemispheres,
and closely related to autonomic spring of action; is
necessary to explain consciously directed thought and
behavior. It seems that the thalamic reticular system,
with its diffuse cortical projections, relations to
afferent and efferent systems, relations to mesencephalic
hypothalamic and striatal systems, is a good candidate
for this office. (Jasper, 1949, p. 419)
Discussion
The brain's mobilization systems with their brainstem, thalamic,
and forebrain components, regulate consciousness, unconsciousness,
and the differential consciousness of attention. Sensory signals
representing possibly significant stimuli cause the reticular
formation to initiate diffuse cortical arousal. When a stimulus
has been identified, cortical activation systems facilitate the
organization of cerebral activity to deal with the situation
appropriately. It appears that the frontal lobes direct this process

45
by recruiting psychological operations in the post-central cortex.
The frontal lobe may accomplish this through its influence on the
nonspecific thalamic nuclei which, in turn, control the phasic
activation of specific cortical systems.
It is important to note that the two biochemically mediated
subsystems which control the mobilization processes are duplicated
in both halves of the brain. Unilateral prefrontal lobe lesions
have been found to produce deficits which resemble those seen after
damage to post-central lesions on the same side: right frontal
lesions have been associated with disturbances of emotion and spatial
abilities while left-sided injuries lead to disorders of speech and
thought (Zangwill, 1966; Benton, 1968; Luria, 1973a).. Processes
which disrupt the biochemical balance between the two control
mechanisms have far-reaching psychological consequences which will
be reviewed in a later section.
Motivation: Emotion and Affect
The greatest risk involved in giving up instinct-based responding
in favor of self-determined behavior is the possibility that the
individual might fail to respond appropriately in survival-related
situations. The evolution of the species could not have occurred
if this problem had not been solved. The forces which motivate
adaptive behavior must, by definition, be the single most powerful
influence on the personality structure. The evidence indicates that
the source of these forces lies in the limbic system. It appears
that separate cortical mechanisms mediate their internal experience
and external expression.

46
Amygdala Circuits and the Prefrontal Lobes
The role of the amygdala in emotional processes established by
K1uver and Buey in 1938, has been assumed to be affected through this
structure's close relationship with the hypothalamus. The amygdala
seems to direct behavior toward biological goals (Halgren, 1981)
and is implicated in the control of species-specific behaviors related
to survival needs, including defensive and aggressive behaviors,
sexual activity, and feeding (Isaacson, 1974). In lower forms these
processes might depend on a simplified (instinctive) form of
memory in which stimulus and response are yoked (Pribram & McGuiness,
1975). In addition to mediating emotional states the amygdala is
involved in the analysis of reinforcement contingencies. Amygdala
lesions have been shown to produce impaired recognition of stimuli
associated with rewards (Weiskrantz, 1956; Schwartzbaum, Thompson
& Kellicut, 1965; Jones & Mishkin, 1972) and inability to resDond
appropriately to changes in the magnitude of rewards (Schwartzbaum,
1960).
Strong interconnections with the hypothalamus (via the stria
terminalis and ventral amygdalofugal fibers) give the amygdala
immediate access to information concerning the internal status of
the organism (Price, 1981). The amygdala also receives processed
sensory information from all of the secondary sensory association
areas (Van Hoesen, 1981). Mishkin and Aggleton (1981) noted that
this arrangement places the amygdalae in a position to integrate
external events with their internal consequences, which would
permit the attachment of emotional and motivational significance

47
to sensory stimuli. Kessner (1981) reported experimental evidence
that demonstrated the essential role of the amygdala in encoding
and retrieving the positive and negative attributes of a specific
memory. In lower forms the identification of a motivationally
significant stimulus might result in the release of species-specific
behaviors, but in humans behavior is self-determined. Halgren
(1981) concluded from his amygdala stimultion studies with humans
that "the amygdala helps organize the discharge of emotional tension
into consciousness" (p. 404) and noted that this would allow the
directing of consciousness toward biological goals. The amygdala's
input to the neocortex is directed to the entire prefrontal lobe
both directly, via the uncinate fasciculus, and indirectly, by
way of the dorosomedial thalamus (Noback & Demerest, 1972; Price,
1981).
While damage to the dorsolateral area of the prefrontal lobes
has been associated with intellectual disturbances, lesions of the
orbito-frontal cortex (and orbital undercutting, which disconnects
this area from the amygdala) result in emotional changes (Lewin,
1961). Eli thorn, et al. (1958) concluded that this type of damage
produced a "generalized impairment of the ability to form appropriate
emotional responses" (p. 250), including the ability to elaborate
on the affect appropriate to the concepts present in consciousness.
In contrast to the planning deficits, loss of energy and interest,
and affective dullness seen after dorsolateral frontal damage, orbito-
frontal injuries often lead to euphoria, impulsive (disinhibited)
behavior, and the appearance of "greediness, selfishness, and

48
tactlessness" (see Lewin, 1961). Faust (1966) noted that such
patients resemble psychopaths in that they are unable to profit
from experience and are in constant conflict with their environ
ment and the law. Zangwill (1966) pointed out that the tactlessness
conrnon in frontal lobe patients does not result from a loss of
knowledge of social conventions, but from the failure to regulate
behavior in accordance with those standards.
Disconnecting the orbito-frontal cortex from the aniygdala
(orbital undercutting) has been reported to be the most effective
psychosurgical operation for relieving the symptoms of anxiety
and depression (Elithorn et al., 1958; Lewin, 1961; Levinson &
Meyer, 1965). Elithorn et al. (1958) noted that this procedure
increased reactions of "a hysterical type" and is contraindicated
for those conditions. It is interesting to note that drugs that
reduce anxiety and produce euphoria (e.g., the barbiturates) have
been found to exert an uncoupling effect between the frontal lobes
and the limbic system (Heath & Galbraith, 1965).
Affective Expression
The right hemisphere plays an essential role in both the
comprehension of emotional Communications and the expression of
affect. Patients with right hemisphere lesions showed impaired
recall of stories with emotional content versus neutral stories
(Wechsler, 1973). Hielman, Scholes and Watson (1975) demonstrated
that judgements of the emotional mood of a speaker (sad, happy,
angry, indifferent) made by patients with right temporoparietal
lesions were significantly impaired relative to patients with left
sided lesions and controls. This finding was replicated by Tucker,

49
Watson and Hielman (1976) who showed also that right hemisphere
patients were impaired in the vocal expression of emotion. The
efforts of patients with right temporoparietal damage to impart a
sad, happy or angry tone to their voices were rated as incorrect
significantly more often than controls. Ross and Mesulam (1979)
presented case studies of two well-educated patients who had com
parable damage in the right supra-sylvian area, which is homologous
to Broca's area on the left side. Both patients showed flattened
affect and had completely lost the ability to laugh, cry, or
otherwise express any emotion in their speech. Their ability to
experience and comprehend emotions was unchanged. The authors
noted that the organization of emotion in the right hemisphere
seems to mirror that of language in the left: the area homologous
to Wernicke's area being essential for comprehension, and to Broca's
area, for expression.
Gazzaniga and Ledoux (1977) suggested that right hemisphere
functioning in humans is distinguished only by contrast to the
left; it continues to perform its functions in the same manner
as elsewhere in the phylum. Although it should be noted that the
human right hemisphere is in possession of tertiary association
areas and so would perform those tasks more efficiently, the
data reviewed above are not inconsistent with Gazzaniga's interpre
tation. The brains of lower forms (and, apparently, the right
brain in humans) are primarily concerned with neuronal signals
which represent the survival needs of the organism within the
immediate environment. Interaction with the social environment

50
is critically important to survival throughout the phylum.
Campbell (1974) noted that animal vocalizations and signals are
"emitted only in the presence of the appropriate stimulus" (p. 349)
and warned against equating these vocalizations with human speech:
"the signals . are generated or motivated by the phenomenon
of emotion, and find their neurological origin not in the cortex
but in the limbic system of the brain" (1974, p. 348).
Discussion
It appears that basic human emotional exnerience is an
emergent property of the functioning of mechanisms that originally
served to regulate the emission of SDecies-specific behaviors
which were elicited directly by releasing stimuli in survival-
related situations. The functional system which evolved in humans
decouples stimulus and response. Thus, in humans, it is the emotional
experience evoked by a stimulus--rather than the stimulus itself--
that is the primary motivating factor (reinforcer) which ultimately
determines behavior. Further, this emotional experience might be
most properly considered to be a part of the experiencing person's
environment, since that experience is involuntary and has the power
to condition the person's response. These processes appear to
have their functional impact in the left hemisphere, where the
formulation of behavioral responses occurs.
The physiological mechanisms which motivate adaptive behavior
in humans are centered on the amygdala which integrates information
from the internal and external enviroments in order to attach
emotional significance to stimuli. This structure is involved in the

51
encoding and retrieval of this information in memory and forwards
its signals to the prefrontal lobes where they are experienced as
subjective emotions. The prefrontal lobes appear to utilize these
signals in the process of forming intentions to direct adaptive
behavior. This amygdala-prefrontal pathway appears to be the sub
strate of anxiety and depression. When the amygdala-frontal connec
tion is severed surgically, or uncoupled pharmacologically, the
neocortex experiences euphoria, but fails to behave in an adaptive
manner.
Memory Functions
It is evident that the functional brain systems which form the
infrastructure of personality include separate cognitive, affective
and arousal components. It appears that these subsystems evolved
to take full advantage of a form of learning which utilizes reinforce
ment and emotional experience in determining behavior. The product
of any learning experience is memory. The importance of memories
(or "associations") in the organization of cognitive operations is
obvious and most, if not all, arousal and affective processes must
depend on the ability to discriminate personally relevant stimuli.
Clearly, memory is fundamental to all aspects of personality function,
but the material substrate of memory remains a complete mystery
(e.g., Lashley, 1950; Luria, 1973a); our concepts regarding it are,
of necessity, only abstract descriptions. Before reviewing the
physiological organization of memory systems it will be necessary
to define and delimit, as far as possible, those abstract

52
concepts of memory phenomena that are pertinent to the interests
of the applied psychologist.
Experimental psychologists have traditionally approached
the study of memory by subdividing it into registration, retention,
and recall, attempting to isolate and measure these aspects and
the variables which affect them. Rapaport (1961) criticized this
methodology as artificial, insisting that these functions are in
extricably related and that such experiments merely demonstrate
how memory can_ function under given laboratory conditions, Working
from a psychoanalytic perspective, Rapaport preferred to treat
memory as an aspect of cognition. He argued that "actual memory
phenomena are encountered only in the context of thought processes;
at best the classical memory experiments could ignore this fact
and make us ignore it, but they could not produce memory phenomena
outside this context" (Rapaport, 1961, p. 6). He acknolwedged
the difficulties in determining the relation of indistinct entities
such as emotion and memory and attempted to clarify the psychoanalytic
viewpoint by suggesting that "memory is a motivated behavior
phenomenon and . emotions are motivating factors" (Rapaport,
1961, p. 8). This statement, however, appears to beg the question;
if memories are activated by emotions, then what initiates arousal
and affective processes?
The interaction of cognition, affect and memory in the etiology
and cure of psychopathology were central themes in the work, pub
lished in 1893 by Josef Breuer and Sigmund Freud, which gave
psychoanalysis its start. The emphasis on the significance of

53
memory phenomena in psychoanalytic literature (e.g., slips of the
tongue, forgetting, false remembering, repression) can be traced
to this seminal paper in which the authors concluded that "hysterics
suffer mainly from reminiscences." In this work, Breuer and Freud
made a crucial distinction regarding the memory processes operating
in psychoneurosis which may have sowed the seed from which the
notion of unconscious causation of psychological phenomena germinated:
. . the causal relation between the determining psychical
trauma Ian experience which calls up distressing affects
such as those of fright, anxiety, shame or physical pain]
and the hysterical phenomenon is not of a kind implying
that the trauma merely acts like an agent provocateur
in releasing the symptom, which thereafter leads an
independent existence. We must presume rather that the
psychical traumaor more precisely the memory of the
traumaacts like a foreign body which long after its
entry must continue to be regarded as an agent that
is still at work. (Breuer & Freud, 1974, p. 355)
The authors became aware of this "highly remarkable phenomenon"
and its relation to affective processes in the course of their
experimental treatment of hysterical conversion symptoms:
[We found] that each individual hysterical symptom
immediately and permanently disappeared when we had
succeeded in bringing clearly to light the memory of
the event by which it was provoked and in arousing
its accompanying affect, and when the patient had
described that event in the greatest possible detail
and had put the affect into words. Recollection
without affect almost invariably produces no result.
(Breuer & Freud, 1974, p. 355)
Hillix and Marx (1974) have suggested that "it was necessary for
Freud to invent the psychic apparatus and much of his psycho
analytic theory just to account for what he and Breuer had already
observed" (p. 352). It is to be hoped that recent evidence will
make a more parsimonious accounting possible.

54
The implicitly verbal form of memory referred to by Rapaport
seems to be qualitatively different from the "foreign body" which
Breuer and Freud assumed to be the culprit in hysterical neurosis.
They referred to the latter type of memory by the less formal term
"idea" and indicated that symptom removal depends on the transforma
tion of this "idea" into a more formal thought process so that its
associated affect can be abreacted:
[the therapeutic procedure] brings to an end the opera
tive force of the idea which was not abreacted in the
first instance, by allowing its strangulated affect to
find a way out through speech; and it subjects it to
associative correction by introducing it into normal
consciousness, (p. 356).
This special type of memory would seem to merit a more detailed
description. It is evident that we are concerned here with a
subset of memories which have significanee for the individual.
By definition, these are memories that are associated with reinforce
ment and/or emotional experience. They are experiential (nonverbal)
and may be isolated from consciousness. It may be noted that this
subset of memories will define the relationship between the individual
and his or her environment and might be the organism's most important
survival resource. A concept from social learning theory seems to
encompass this type of memory comfortably and provides a more
operational definition.
Julian Rotter (1966) theorized that "a reinforcement acts to
strengthen an expectancy that a particular behavior will be followed
by that reinforcement in the future" (p. 2). Further, "when an
organism perceives two situations as similar, then his expectations

55
for a particular kind of reinforcement, or class of reinforcements,
will generalize from one situation to another" (Rotter, 1975, p. 57).
Rotter distinguished two types of "generalized expectancies" (GE).
The first has to do with the nature of the reinforcement: expectations
for a particular kind of reinforcement in a given situation. The
second type deals with other properties of situational stimuli and
has to do with the perception of control that one can exercise to
change or maintain the situation: the kind of behavior that is
likely to produce or terminate reinforcement. The first type is
designated with a subscript _r for reinforcement (GEr). The second
type is designated a problem-solving generalized expectancy (GEps).
Striking insights into the nature and mechanics of this sort of
experiential memory were afforded by Penfield's observations of
certain psychical phenomena elicited by direct electrical stimulation
of the conscious brain (see Mullan & Penfield, 1959).
Wilder Penfield1s data were collected from patients undergoing
radical brain surgery, with local anesthesia, for the relief of intrac
table epilepsy. His observations consist of spontaneous reports from
these patients following applications of a mild electric current to
the exposed cortex from the tip of a unipolar electrode. The responses
to such stimulation which are of interest here fall into three
categories:
1. The emergence in consciousness of vivid and coherent
experiential hallucinations which appeared to be recollections of
(or abstractions from) the subject's past experiences.
2. Changes in a patient's subjective experience of his or her
relationship with the immediate environment.

56
3. "Illusory" emotional experiences.
Based on his analysis of the data, Penfield (1975) postulated the
existence of two related brain systems: a "mechanism of recall,"
and a "mechanism of interpretation." The latter involved the temporal
cortex (exclusive of the speech areas) and was referred to by Penfield
as the "nonverbal concept mechanism." Penfield comoared its func
tion with nonverbal concepts to the operation of the speech cortex
with verbal concepts.
Somehow [this mechanism] seems to analyze the components
of sensation, compares them with previous experience,
and by that analysis and comparison, transmits into
consciousness their present and immediate significance
... [an emotional response] is a signal that rises
into consciousness as a result of an interpretation of
what the present situation may bring the subject in
the immediate future.. (Mullan & Penfield, 1959, p. 283)
It appears that the cognitive products of these mechanisms fit
the criteria for generalized expectations. Penfield's evidence
assists in understanding the different types of memory referred
to by Rapaport and by Breuer and Freud and indicates the neural
substrate of these processes. (Penfield's data and conclusions
will be reviewed and evaluated in detail presently.)
It has been assumed here that at the base of personality there
are mechanisms whereby cognitive operations and affective processes
are appropriately activated by significant memories. The present
task is to describe neurological evidence which accounts for the
memory phenomena reviewed above in terms that fit the criteria
for "functional systems" as defined by Luria, and satisfy the
evolutionary imperatives outlined at the beginning of this chapter.

57
The neuropathological correlates of human amnesia syndromes and data
from related animal studies will be reviewed. It will be hypothe
sized that human beings possess two autonomous, lateralized memory
systems, centered on the hippocampi, each of which is intimately
associated with its own emotional and cortical activating system.
Human Amnesia Syndromes
A number of terms are used to describe different aspects of
memory function and dysfunction. Immediate memory, usually measured
by digit span, probably reflects the ability to hold information
in the primary or secondary sensory cortex as long as voluntary
attention (directed by the nonspecific thalamic nuclei) is focused
upon it (Smithies, 1966). The terms short-term ("recent") and
long-term ("remote") memory indicate recollection over increasingly
greater periods of time, but both are almost certainly subserved
by the same physiological processes (Brierley, 1977). The term
retrograde amnesia refers to a period of time before an accident or
illness for which the patient's ability to recall is diminished
or lost. Anterograde amnesia is an inability to retain in memory
events that occur after such an injury or illness.
The combination of a severe retrograde amnesia and a debilitating
anterograde amnesia is the hallmark of the Wernicke-Korskoff
syndrome, the most common form of memory disease. This illness is
most frequently seen as a result of brain lesions brought on by
dietary (thiamine) deficiencies in chronic alcoholics, although the
lesions and illness may be produced by a number of toxic or disease
processes (see the excellent review by Brierley, 1977). This illness

58
was described independently in the 1880s by Karl Wernicke (whose
work with aphasia was reviewed earlier) and S. S. Korsakoff, a
Russian psychiatrist. Wernicke focused on the acute stage of the
illness during which the patient is usually depressed, fearful and
anxious; often paranoid; and always severely confused and disoriented
(Wernicke's encephalopathy). Patients who survive this acute stage
become stabilized in the phase known as Korsakoff's psychosis.
This chronic state is characterized by severe memory disorders and
profound changes in the patients personality, motivation, and affect.
Against a background of retained intellectual skills and intact
remote memory, the victim of Korsakoff's psychosis suffers a retro
grade amnesia for periods of up to several years before the onset
of the illness, and an almost total inability to recall any new
information once his or her attention is distracted from it. Conse
quently, these patients live virtually in the immediate present and
are always disoriented as to time, place and situation. These
patients are, at best, only vaguely aware of their inability to learn
new material. They often produce confabulations to cover gaps in
their recollection and fuse or combine ("reduplicate") experiences
from different periods in their lives (Gardner, 1975). In both
cases they believe that their statements reflect reality. In most
cases the patient's affect is blunted, although some patients have
exhibited chronic euphoria (e.g., Remy, 1942). They show reduced
spontaneity and initiative and a "lack of desire for alcohol, sex,
and other traditional reinforcers"'(Gardner, 1975, p. 11188).

59
(Such indifference to alcohol is especially interesting in light of
the fact that the brain damage in most of these patients was caused
by years of sustained, heavy drinking.)
Post-mortem examination of the brains of persons who suffered
from Wernicke-Korsakoff disease reveals lesions of certain anatomical
structures in the limbic system associated with the well-known
circuit of Papez. Such damage almost invariably includes, and may
be confined to, injury to the mammillary bodies, the relay for
hippocampal output on its way to the anterior thalamic nuclei
(Brierly, 1977).
A pure form of this memory disorder was the unfortunate consequence
of bilateral removal of the hippocampi in humans. In the 1950s
Scoville performed a series of experimental operations designed to
relieve the symptoms of chronic schizophrenia without the undesirable
side-effects of a complete frontal lobotomy. The surgical procedure
involved the resection of the medial surface of the temporal lobes
from 5.0 to 8.0 cm posterior to the tip of the lobe combined in
some cases with orbital undercutting. Thirty severely deteriorated
schizophrenics had undergone the operation, with slight improvement
in their conditions, when a purely temporal resection was performed
on a nonpsychotic epileptic patient whose seizures were unresponsive
to medication. When this patient, "case H. M.," recovered from
the operation it became apparent that he had developed a severe
amnestic disorder which resembled Korsakoff's psychosis and which
persisted at 14 years (Milner, Corkin & Teuber, 1969). Scoville
and Milner subsequently examined eight of the psychotic patients

60
who had undergone the operation and who were able to participate in
formal testing. They discovered "some generalized memory disturbance
in all patients with removals extending far enough posteriorly
to damage portions of the hippocampus and hippocampal gyrus (Milner,
1958, p. 112). The degree of memory impairment was more or less
proportional to the amount of these structures removed. Bilateral
resection of the uncus and amygdaloid nucleus alone did not result
in amnesia (Brierly, 1977), nor did removal of the gyri of the outer
aspects of the temporal lobes (Bailey, 1946). It has been concluded,
therefore, that "the structures necessary for normal memorizing are
the hippocampal formations within the temporal lobes, the manrnillary
bodies and, possibly, certain thalamic nuclei within the diencephalon"
(Brierley, 1977, p. 221); that is, the hippocampi,, their output
pathways and related projection sites.
There has been, as yet, no definitive explanation of either the
nature of the amnesic deficit described above or of its underlying
mechanisms. The fact that remote memory seems to be intact in
these patients has led many to believe the the hippocampal-
diencephalic structures are not involved in the process of recall,
although Brierley (1977) pointed out that such a conclusion is
unjustified in the absence of adequate pre-post evaluation of this
function. Milner (1966, ch. 5) attempted to account for the pairing
of a period of retrograde amnesia with an inability to learn new
material by hypothesizing that the establishment of a permanent
memory trace requires an extended period of "consolidation," which
is somehow disrupted in this syndrome. In Milner's view the deficit

61
represents a failure to transfer sensory impressions into long-term
store. The adequacy of the consolidation hypothesis is called into
question, however, by demonstrations that amnesic patients are in
fact able to recall new information under certain conditions. The
most convincing evidence comes from experiments using the technique
of "cued recall" in which a subject is given partial information
about a stimulus (e.g., a previously presented word or picture)
and asked to identify the whole item. Under these circumstances the
performance of amnesic subjects was not significantly different
from that of controls (Weiskrantz & Warrington, 1970). This
suggested to the authors that the amnesic deficit involved problems
with mechanisms of retrieval rather than those of acquisition or
retention.
Weiskrantz (1979) reviewed a number of experimental paradigms
in which normal learning has been demonstrated in amnesic subjects
and underscored the fact that, in each case, the patients themselves
persistently failed to acknowledge the fact that their performance
was based on specific past experience (or that they had been
confronted with the task before). Thus, amnesia victims do not have
access to their memories on a conscious level, nor is such awareness
necessary for that memory to be demonstrated objectively. Weiskrantz
pointed out that this "striking dissociation between the subjects'
commentaries and their objective performance . suggests a
dissociation between levels of processing rather than a failure on
any particular level" (p. 385).

62
Levels of processing in memory are the subject of a theory
(summarized by Gaffen, 1972) which is based on arguments by Tall and
(1965) and supported by experimental evidence (Peterson, 1967;
Kintsch, 1970). Briefly, the theory postulates that the process
of recall consists of two separate and autonomous stages: retrieval
(or search) and recognition. The retrieval process "proceeds at
several levels . each being terminated by an implicit act of
recognition" (Talland, 1965, p. 304). The recognition stage is
based on a record from which the past cannot be read directly,
but which can assign a particular response in a particular context
a value of "familiarity--unfamiliarity" (the correct response being
the most familiar in that context). Thus "[in the retrieval stage]
various responses are generated (but not emitted); when finally the
correct response is generated, it is recognized as such by the
recognition stage, and is then emitted" (Gaffen, 1972, p. 328).
The theory postulates that amnesic subjects (animal and human)
lack the faculty of discriminating familiarity. This basic deficit
is manifested in the premature termination of search cycles, resulting
in an incorrect match. These formulations are not inconsistent
with those of Butters and Cermack (1974), who concluded from their
experiments that increased sensitivity to proactive interference,
subsequent to inappropriate encoding of information, was the
critical factor underlying the amnesic disorder. Finally, it is
interesting to note that modern theories regarding amnesia seem
to have arrived at the point at which they began: Korsakoff (1889),
in keeping with the associationist doctrine of his time, believed

63
that his patients were deficient in making associations among new
ideas and in connecting past and present experience.
The suggestion of bilevel processes in memory noted above are
particularly interesting given that language and conscious awareness
have been associated with temporal lobe structures of the left
hemisphere, and Penfield's (1975) report that electrically induced
"illusions of recognition" were elicited only by stimulation of
temporal structures of the right hemisphere.
Memory and the Neocortex
As noted earlier, Penfield's experiments with electrical stimula
tion of the cortex in conscious patients led him to postulate the
existence of two separate memory systems: a "mechanism of recall"
and a "mechanism of interpretation." The existence of the former
was suggested by the fact that, following stimulations of the exposed
cortex, some of his patients reported vivid auditory and/or visual
experiences ("flashbacks"); it seemed to the patients as if they
were reliving prior experiences in their lives, although they retained
their awareness of the operating room environment. Since many of
these sensory sequences were trivial, yet perceived as familiar,
Penfield concluded that his electrode was tapping a "continuous
record of conscious experience." Although widespread areas of the
cortex were exposed and explored, these "experiential responses
come only from the temporal lobe, never from any other part of the
brain" (Penfield, 1975, p. 31).
Niesser (1967) presented persuasive arguments refuting Penfield's
claim that "nothing is lost . the brain of every man contains an

64
unchanging ganglionic record of subjective experience" (Penfield,
1954, p. 67). Niesser suggested that "most of the cases described
by Penfield seem more like generic and repeated categories of events
rather than specific instances" (p. 168). (It will be noted that
Niesser's formulation is congruent with the notion of a "generalized
expectation," as defined earlier.)
Penfield's second mechanism was suggested by another category
of electrically induced phenomena which consisted of the "misrepre
sentation or altered interpretation of present experience" (Mullan &
Penfield, 1959, p. 269). Prominent among these were "illusions of
recognition" during which "present experience seemed familiar,
strange, altered, or unreal" (p. 270). These "illusions of compara
tive interpretation" were associated with stimulation of the temporal
cortex in the hemisphere that was minor for handedness and speech.
The authors believed that "in normal life, these are signals that
rise into consciousness, signals that depend on subconscious
comparison of past experience with the present" (p. 283).
In 1951 Penfield proposed that portions of the temporal lobes
be called "memory cortex" in the belief that his electrode had
activated a neuronal record which was stored there. He was obliged
to revise this theory in 1958 because of a new understanding of the
physiology of electrical brain stimulation. When an electrode passes
a current into the cerebral cortex, the current completely disrupts
the patient's normal use of that gray matter (e.g., stimulation
of the speech areas produces momentary aphasia). Therefore, any
positive responses are produced by axon-conduction and the functional

65
activation of a distant, secondary ganglionic station (Penfield,
1975, ch. 7). In his later formulations then, Penfield referred
to those temporal structures as the "interpretive cortex" and postu
lated that his electrode had activated a final common pathway to
a secondary center which in turn produced the illusions of
comparative interpretation. Since the temporal lobe forms the
principal source of input into the hippocampus (which was known to
be related to memory), Penfield assumed that this was the secondary
center in question. He suggested that
The hippocampi seem to store keys-of-access to the
record of the stream of consciousness. With the
interpretive cortex, they make possible the scanning
and the recall of experiential memory. (Penfield,
1975, p. 36)
Penfield's finding that illusions of familiarity were associated
with activity of the temporal lobe in the right hemisphere is
complemented by Kimura's (1963) evidence that the right temporal
lobe appears to be more involved in the analysis of unfamiliar stimu
li. Kimura presented familiar and unfamiliar visual stimuli to the
right and left visual fields of patients with lesions of the right
or the left temporal lobe. The right temporal group was impaired in
the perception of the unfamiliar stimuli but not the familiar.
Kimura interpreted her results in terms of the verbal i dentifiability
of a stimulus:
It seems clear that a frequent (though not a necessary)
concomitant of familiarity in a perceptual sense is the
possibility of verbal identification. Where increased
familiarity with a stimulus object, or class of objects,
is associated with the repeated naming of the object,
the ability instantly to attach a name to it represents
an important step in the development of a concept. It

66
seems probable that in such cases a large part of the
increase in permanent neural representation which is
assumed to correlate with familiarity will take place
in the language centers, that is, in the dominant
hemisphere. (Kimura, 1963, p. 269)
Thus, unfamiliar stimuli, which are not represented in verbal
memory by a permanent neural model (e.g., a name or concept) are
more likely to be processed by the right hemisphere, which Kimura
suggests is more important than the left in the establishment of
such "cell assemblies." Since all of the material in the memory store
are initially unfamiliar it follows that many, if not all, verbal
concepts (in the left hemisphere) might be based on neural models,
or gestaltan, which were assembled (and are stored) in the right
hemisphere. Such an hypothesis is supported by evidence from
split-brain studies that the right hemisphere is far superior to
the left in the discrimination of part-whole relationships (Nebes,
1974).
The appearance of material-specific amnesia syndromes following
unilateral temporal lobectomies suggests that the isolation of
language in the left hemisphere extends to verbal learning also, and
is thus virtually complete. Milner (1971) reports that
A comparison of left and right anterior temporal lobec
tomy in epileptic patients has revealed certain specific
memory defects that vary with the side of the lesion.
These material-specific disorders are to be distinguished
from the global amnesia that follows bilateral damage
in the hippocampal zone (Milner, 1958). Thus, left
temporal lobectomy, in the dominant hemisphere for
speech, selectively impairs the learning and retention
of verbal material (Meyer & Yates, 1955; Milner, 1958).
. . Conversely, removal of the right, nondominant
temporal lobe leaves verbal memory intact but impairs
the recognition and recall of visual and auditory patterns

67
that do not lend themselves easily to verbal encoding.
. . Thus within the sphere of learning and memory
there is a double dissociation between the effects of
these two lesions. (Milner, 1971, p. 274)
Butters and Cermack (1974) focused on the specifically verbal
aspects of the amnesic disorder in Korsakoff patients and noted
that, during learning tasks, these patients did not react to
changes in semantic categories on successive lists. They concluded
that the amnesic deficit was due to the patient's
inability to encode verbal information along semantic
or meaning dimensions. . Korsakoff patients do not
spontaneously employ semantic encoding strategies,
but rely on basic acoustic and associative categorizations.
If the Korsakoff patient is instructed to encode semanti
cally, he will do so, but in an impaired manner.
(pp. 74-75)
Butters and Cermack assumed that the Korsakoff patients'
deficient utilization of "meaning" in learning tasks was a specifically
verbal (i.e., left hemisphere) phenomenon. However, Gazzaniga and
his colleagues produced evidence which suggests that the right
hemisphere may play an important role in imparting "meaning" to
verbal memory processes. These authors tested patients with partial
or complete section of the cerebral commissures for recall of two
lists of paired-associate nouns. On presentation of the second
list each patient was instructed to "form a 'picture in his mind'
of the two items interacting in some unusual or amusing way"
(Gazzaniga, Risse, Springer, Clark & Wilson, 1975, p. 12). Patients
with partial sectioning of the cerebral commissures showed marked
improvement with the imagery instructions but none was seen where
there was complete section of the hemispheric interconnections.

68
Discussion
The evidence reviewed thus far suggests that the central problem
in the amnesia syndromes involves mechanisms which normally facili
tate access to stored memory traces. The problem might occur at
the point of encoding and deposition, or retrieval, or both.
The hippocampus appears to be critical to these coding and decoding
operations; this structure may normally provide the memory "cues"
which must be externally administered in its absence.
The evidence is supportive of Niesser's (1967) proposal that
(rather than tapping a "continuous memory strip'1) "Penfield's
electrode may have touched on the mechanisms of perceptual synthesis"
(p. 169). There can be little doubt that Penfield's "final common
pathways" in the temporal lobes are related to hippocampal afferents.
However, it is now clear that the hippocampus cannot be the only
secondary ganglionic station which must be activated before a signal
indicating familiarity, originating in the right hemisphere, can
"rise into consciousness." Studies of split-brain patients have
shown conclusively that the ability to give a verbal account of
events occurring in the right hemisphere is dependent on the
integrity of the cerebral commissures (e.g., Sperry, 1968).
The general layout of the commissures is such that a specific area
in one hemisphere is connected via commissural fibers to the homolo
gous area in the contralateral hemisphere. The temporal lobes have
their own private interconnection in the anterior commissure, which
also connects the two amygdalae (Gray, 1977). Studies of patients
with only partial sectioning of these commissures have indicated

69
that highly processed information might be even more "transferable"
than elementary sensory information and may be able to utilize any
commissural pathway that is available (Gazzaniga. et al., 1975).
It appears that there are two separate and autonomous memory
systems in the brain which are specialized as to their function:
a verbal system, lateralized in the left hemisphere, and a nonverbal
(experiential) system lateralized in the right. The evidence
suggests that memory functions might be conceptualized as having
both vertical and lateral dimensions: the scanning (or search)
within each system may involve a temporal-hippocampal interaction
and the phenomenon of recognition may be a function of right-left
temporal lobe interaction.
A voluntarily initiated search of memory would, most certainly,
begin in the left-hemisphere system, but an environmental stimulus
might activate an initial nonverbal (right-hemisphere) scan and
analysis. In either case, the result of these processes would seem
to be the emergence in conscious awareness of a signal indicating
the "familiarity" (or "strangeness") of the stimulus and, finally,
the facilitation of generalized expectations and verbal associations
related to that stimulus. The implications of such a formulation
for the understanding of psychopathological processes and the practice
of psychotherapy will be discussed in later sections, following
an evaluation of the limbic system's role in experiential and emo
tional memory.

70
The Limbic System, RAS, and Memory
Little distinction is made in the human amnesia literature
between the verbal, experiential, and emotional aspects of memory.
The verbal manifestations of the disorder are the most obvious, the
most amenable to description and testing, and so have become the
primary focus of scientific attention. Although it is seldom
emphasized, most case studies of amnesia victims relate anecdotal
evidence of emotional dysfunction (e.g., blunted or flattened
affect; euphoria). There are also more or less vague, but consistent,
references to what might be termed disturbances of tonic arousal
("decreased spotaniety," "lack of initiative," "indifference,"
"passivity"). These emotional and arousal difficulties appear to
be associated with systems centered on the amygdala and RAS,
respectively. Both of these will be described below following a
consideration of the nonverbal, predominantly unconscious mechanisms
of experiential memory.
Before an organism can respond to a stimulus on any level
(emotional, verbal, or behavioral) its meaning must be ascertained.
At the most elementary level the organism's survival depends on
its ability to make appropriate decisions about whether to invest
neural energy in attending to and further analyzing a particular
stimulus (orienting) or to ignore it (habituation). Such a decision
demands a judgement as to the apparent novelty, possible significance,
or lack of these qualities in the stimulus configuration, a process
which requires access to a patterned memory trace or "neural
model" (Sokolov, 1963). Such a process must begin with sensory

71
input and end with afferents which are capable of modulating the
activity of the brainstem RAS.
A determination of novelty or significance might be made at
the level of the secondary sensory association cortices where raw
receptor impulses are converted into functional (i.e., "meaningful")
information, although such duplication of effort in all the modalities
would be cumbersome and inefficient. Further, the relative signifi
cance of a stimulus may depend on the internal state of the organism
(e.g., satiation, the presence or absence of certain hormones, etc.).
The fact that information related to this added consideration is
most readily available in subcortical structures is another argument
favoring a centralized location for the mechanisms involved with the
decision to orient or habituate. The hippocampus meets all of the
criteria specified above.
Luria concluded that "many nuerons in the hippocampus and
connected nuclei do not respond to modality-specific stimuli of
any sort, but serve to compare present stimuli with traces of
past experience; they react to every change in the stimulus and
thus play to some extent the role both of 'attention neurons' and
of 'memory neurons'" (1973a, p. 289). According to Luria the
hippocampus provides for the "elimination of responses to irrelevant
stimuli and enables the organism to behave in a strictly selective
manner" (1973a, pp. 271-272). It appears that the hippocampus
accomplishes this complex task by coordinating the activities of
the cortical and subcortical mechanisms which are directly in
volved in the processes of attention, memory and learning.

72
Early investigators were puzzled by the fact that cortical
activation (EEG desynchronization) was accompanied by synchronous
slow-wave activity (4-8 Hz theta rhythms) in the hippocampus
(e.g., Green & Arduini, 1954). The most constant behavioral corre
lation of hippocampal theta activity in animals of different species
is orienting towards, and attending to, stimuli in the environment
(Isaacson, 1974). Cortical activation, such as that seen in the
orienting response, is accomplished by the brainstem RAS in conjunction
with the nonspecific thalamic nuclei. The hippocampus appears to
regulate the process of involuntary attention by performing switching
functions through a mutually inhibitory relationship with the reticu
lar formation (Smithies, 1966). The Soviet neuropsycholgist Vinogradova
provided important insights into the mechanics of this process.
By observing unit activity with microelectrodes Vinogradova
(1970) determined that all of the neurons in the hippocampus monitor
incoming stimuli, habituating to repetition and dishabituating to
any change in the stimulus configuration. Such responsiveness
requires constant matching of the stimulus with a related neuronal
model (Sokolov, 1963). That these models exist in the cortex, and
not in the hippocampus itself, is established by evidence that the
quality of sensory information is almost completely erased in
hippocampal neurons (Gloor, 1961).
Vinogradova distinguished two types of neurons in the hippo
campus: A-neurons (30-40%) which are activated by a stimulus and
I-neurons (60%) which are inhibited. She went on to proDOse a
mechanism whereby the hippocampus is able to modulate the processes
of attention and learning:

73
The hippocampus exerts a tonic inhibitory influence upon
the reticular formation, blocking activatory processes
through the tonic discharge of its I-neurons when
novelty is absent and registration [a change in the
neuronal model] is not needed. But when a stimulus which
is not registered in the memory system appears, this
inhibitory control is blocked (I-neurons become silent],
arousal occurs, and the process of registration starts.
(1970, p. 114)
Vinogradova's hypothesis is supported by observations of electrical
activity in the brains of animals in classical conditioning paradigms.
Theta rhythms (associated with activity of Vinogradova's A-neurons)
are found in the early stages of learning but disappear when the
response has become well established (Isaacson, 1974). During
conditioning the time course of the theta rhythm and the orienting
response are matched. As the latter is replaced by the stabilized
conditioned response, theta dies out (I-neurons become active again)
and the hippocampus resumes its inhibitory controlover the RAS,
thus ending the orienting response and thereby allowing the fully
developed conditioned response to materialize (Smithies, 1966, p. 90).
The hippocampal-cortical interaction was apparent in an
analysis of the characteristics of hippocampal theta rhythms in
situations requiring different types of cortical information
processing. Bremner (1970) investigated the effects of orienting,
simple conditioning, discrimination, and discrimination reversal
tasks on various parameters of the theta rhythm using the habituated
organism (rat and man) as a baseline. He found that theta power
(amount of energy) increased in the presence of stimuli which elicited
orienting and arousal and decreased in the interval preceding a
response in the conditioning situation; the range of energy distribution

74
around the peak frequency narrowed during discrimination; and the
location of the peak shifted in discrimination reversal procedures.
In summary, the hippocampus appears to be able to monitor
incoming stimuli and match them against (cortical) neuronal models
which represent the past experience of the organism with related
stimuli. In addition to facilitating appropriate access to these
memory traces hippocampal activities have been directly associated
with the triggering of cortical processes which permit further
analysis of a stimulus, emotional and behavioral responses as
warranted, and/or alterations in the neuronal model itself (i.e.,
learning).
Papez Circuit and Memory
The hippocampus forms part of a continuous pathway within the
limbic system which Papez (1937) believed to be the substrate of
emotion. Papez was aware that destruction or stimulation of limbic
structures produced major alterations in emotional behavior and
believed that emotional expression depended on the integrative action
of the hypothalamus. He was also convinced that subjective emotional
experience required the participation of the cerebral cortex.
Papez outlined an anatomical circuit through which he thought
emotion might arise in either of those two centers. Thus
Incitations of cortical origin would first pass to
the hippocampal formation and then down by way of
the fornix to the mammillary body. From this they
would pass upward through the mammillothalamic tract
... to the anterior nuclei of the thalamus and thence
by the medial thalamocortical radiation (in the
cingulum) to the cortex of the gyrus cinguli . .
Radiations of the emotive process from the gyrus
cinguli to other regions in the cerebral cortex

75
would add emotional coloring to psychic processes
occurring elsewhere. (Papez, 1937, pp. 304-306)
Papez believed that sensory input to the system originated in the
thalamus and was communicated via the subthalamus to the hypothalamus.
The circuit is completed with the connection of the cingulate
gyrus to the hippocampus by way of the cingulum bundle.
It is now known that other limbic structures are more actively
involved in the specifically emotional processes. However, Papez's
anatomical concepts might be rehabilitated if their context were
changed from emotion to memory. Input to the hippocampal circuits
would be seen as processed sensory information and its output as
memory indexing information capable of "cueing" associations and
generalized expectations related to the input stimulus. It remains
to place the role of the hippocampus in the context of the functional
system it subserves and to examine the other components of that
system.
Fornix. Hippocampal output makes its way via direct and
indirect pathways to the thalamus (anterior, dorsomedial, and intra
laminar nuclei), the hypothalamus, and the midbrain reticular
formation. Its main efferent fiber system, the fornix, is composed
of axons from pyramidal cells in the body of the hippocampus.
These fibers converge in the fimbria, traveling backwards within the
temporal lobe, and then arch forward under the corpus callosum as
the crura (posterior pillars) of the fornix. Here a number of
a
fibers cross to the other side, forming the hippocampal commissure.
The two crura then join to form the body of the fornix which

76
continues to arch forward, following the course of the lateral
ventricle to the rostral edge of the thalamus. Here the bundles
separate again to form the anterior columns of the fornix which
curve downward in front of the intraventricular foramen and above
the anterior commissure (AC). Approximately half of the fibers
descend behind the AC as the postcommissural fornix; the other half,
in a less compact bundle, pass in front of the AC as the precommissural
fornix. Postcommissural fibers pass through the hypothalamus to
the mammillary bodies, giving off fibers to the thalamus on their
way. Some of the precommissural fibers distribute to the septal
area and others join with septal fibers and continue into the same
areas as the postcommissural fornix. Approximately one-third of the
fornix fibers reach the mammillary bodies (Daitz, 1953); the anterior
nuclei of the thalamus receive as many direct fibers from the fornix
as they do from the mammillothalamic tract (Truax & Carpenter, 1969,
ch. 21).
Liss (1968), working with the rat, found that hippocampal
and fornix lesions had analogous effects on learning and behavior
in passive and active-avoidance tasks. In the monkey, fornix
lesions led to impaired learning of a spatial reversal task that
was "functionally similar" to the deficit seen after hippocampal
removal (Mahut, 1972; Mahut & Zola, 1973). Gaffen (1972) reported
a series of six experiments in which rats with fornix lesions were
shown to have a defect in "recognition memory" that the author
argued was eguivalent to anterograde amnesia in humans. In Russel's
(1971) survey of brain wound cases, the eight patients who showed

77
a Korsakoff-type of memory disorder were thought to have had their
fornices damaged by metal fragments. Four of the five cases with
typical amnesic syndromes in Jarho's (1973) study of brain-injured war
veterans were considered to have bilateral interruption of the fornix
or mammillothalamic tract.
Sweet, Talland, and Ervin (1959) reported the case of a woman
in whom the anterior columns of the fornix were sectioned to facilitate
the removal of a colloid cyst of the third ventricle. This patient
showed a rapid recovery of old skills, but developed a "severe loss
of memory for recent events [which persisted at two years] ... a
retrograde amnesia of at least several weeks and a subjective
complaint of amnesia for specific events of four or five years past"
(p. 76), coupled with intact remote memory. In the discussion
following the presentation of this patient, Brenda Milner reported
on a similar case operated on by Welsh in 1954. This patient showed
a gross initial memory disorder but, at one year, was able (with
effort) to effect some compensation for his defect: "Whenever he
deliberately sought associative links he was able to improve his
performance considerable" (Sweet et al., 1959, p. 79). Milner
concluded that
I think that one can only account for the paradoxical
diversity of data from the fornix cases, as contrasted
with the consistent and severe memory loss in the hippo
campal cases (and maybe in the mammillary body cases
also), by supposing that you are only interfering
with a part of the system by fornix section. Thus,
you are most apt to see a temporary disruption of dis
turbance of memory with minimal residual loss. (p. 79)

78
Data on the fornix is relatively scarce, and there have been
reports of negative findings. In his influential review, Brierly
(1977) took a very conservative stance on this subject:
The most discrete link between the hippocampal and
diencepahlic regions is the fornix. It is surprising,
therefore, that with the exception of the case reported
by Sweet, Tall and and Ervin (1959), bilateral division
of the fornix (usually in the region of the intra
ventricular foramen) has not resulted in a disorder
of memorizing (Dott, 1938; Cairnes & Mosberg, 1951;
Garcia-Bengochea and his colleagues, 1954). This
finding suggests that the two groups of structures
linked by the fornix cannot be regarded as a unitary
system subserving the process of memorizing, at least
until major interconnections other than the fornix
have been identified, (pp. 221-222)
Other interconnections are available. Smithies (1966) describes
a "massive direct hippocampal-hypothalamic pathway" that runs
diffusely through the subthalamus and which he suggests might be
"quite able to carry on hippocamoal and limbic circuit function in
the absence of [the fornix]" (p. 122). A seond look at the reports
cited by Brierley, however, allows the possibility that his
conclusions are premature.
Sweet et al. (1959) emphasized that their patient's "conversa
tions, social amenities, and general demeanor gave little evidence
of [her] severe deficits unless they were specifically looked for"
(p. 76). They also noted that she lacked spontaniety, made little
effort to converse, and was apparently indifferent to her deficits
(cf. Korsakoff's syndrome). The very brief report of Bengochea,
De la Torre, Esquival, Vieta and Fernandez (1954), after a "short
follow-up" of their patients (whose fornices were severed in an
experimental operation to relieve intractable epilepsy) did not

79
mention any attempt at quantification of behavior. They simply
stated that "so far, in none of the 12 surviving cases there has
been [sic] any unfavorable neurological or psychiatric sequela"
(p. 177). It seems possible that these authors may have missed
subtle symptoms in their apparently superficial evaluation.
Wilder Penfield (quoted in Sweet et al, 1959) underscored the
fact that "patients who have [bilateral hippocampal lesions] do
not forget their skills. Two of them were able to carry out most
complicated skills learner previous--glove cutting and engineering
drawing" (p. 81). Unfortunately, the only behavioral measure
reported by Cairnes and Mosberg (1951) involved a return to work.
These authors noted that some of their pateints (who had incurred
fornix damage in the course of surgery to remove colloid cysts
of the third ventricle) showed initial confusion, loss of memorizing,
and amnesia for the period surrounding the operation, but: "after
operation all [but one of their nine surving cases] returned to
work, and . showed no disturbance of emotion or intelligence"
(p. 564). Thus
Four of the five young women . are doing normal
housework; three have borne children. The other
young woman is in regular work as a clerk, and is
free from complaints. . Two older women . .
are also doing their housework [although one has a
'slight impairment of memory']. ... Of the two men,
one is working regularly as a doIicemen, (p. 568)
The extent of the lesions in these patients is unclear. The authors
report only that "each had . partial or complete division of
the anterior columns of the fornix" (p. 564). (It seems possible
that these surgical fornicotomies spared the precommissural fornix.)

80
It should be noted that a colloid cyst of the third ventricle
tends to produce confusion, dulling of attention and memory, and
sometimes a progressive dementia prior to its surgical removal.
These factors would make a pre-post evaluation of memory function
very difficult. Still, the scantiness of the reported date in the
studies reviewed above is unfortunate. It is evident that injuries
to different parts of this system result in different expressions
of the disorder. It is reasonable to conclude, however, that
damage to the fornix has adverse effects on memory function which
vary as to the quality and degree, and may leave a greater possibility
of recovery of function.
Mammillary bodies and mammillothalamic tract. The mammillary
bodies are a collection of nuclei at the posterior boundary of the
hypothalamus. They form a major relay station for hippocampal
output on its way to the thalamus (via the mammillothalamic tract)
and to the midbrain reticular formation (by way of the marnmillo-
tegmental tract).
In humans, damage which is apparently limited to the mammillary
bodies has resulted in the full Korsakoff amnesic syndrome (Remy,
1942; Delay & Brion, 1951; Gruner, 1956; Symonds, 1966), although
Victor (1964) suggested that additional damage to the thalamus
was necessary to produce the disorder. In the rat, lesions of the
mammillary bodies or of the mammillothalamic tract impaired the
ability to perform a spatial discrimination in a T-maze in order to
avoid footshocks (Thompson, Langer & Rich, 1964). Krieckhaus
(1962, 1964) found that complete or partial destruction of the

81
mammiliothalamic tract in the cat reduced the retention of a two-way
active avoidance task and produced a less striking deficit in the
retention of a one-way active avoidance task. These findings were
later replicated in the rat (Krieckhaus, 1965). Thomas, Frey,
Slotnick and Krieckhaus (1963) studied the post-operative acquisition
of the two-way active avoidance learning task and reported mixed
results: four of their eleven cats were completely unable to learn
the task, while seven of them mastered the problem within the
number of trials required by normal animals. (The significance of
the distinction between acquisition and retention will be discussed
in the following section.)
Cingulate cortex. The cingulate cortex lies above the
corpus callosum on the medial side of the hemisphere, separated
from the neocortex above by the cingulate sulcus. It merges with
the hippocampal gyrus posteriorly and with the neocortex of the
frontal lobe anteriorly. As noted by Papez, the cingulate gyrus
receives its main afferent supply from the anterior nuclei of the
thalamus and projects to the hippocampus via the cingulum bundle.
Stimulation of the cingulate cortex also produces activity in the
prefrontal and orbitofrontal regions of the neocortex (Dunsmore &
Lennox, 1950). There are reciprocal connections with the anterior
and other thalamic nuclei (including the dorsomedial). A strong
projection to the interior parietal lobule (IPL) in the post-central
neocortex has been demonstrated in the monkey (Mesulam, Van Hoesen,
Pandya & Geshwind, 1977). These authors, using the horseradish
peroxidase technique, found that "the cingulate gyrus contained one

82
of the heaviest concentrations of labeled neurons in most cases"
following injection of that substance into the IPL (p. 324).
The literature documenting the efforts of physiological
psychologists to define the role of the cingulate cortex in learning
and memory is confused somewhat by varying interpretations of the
data by those authors (an objective review is available in Isaacson,
1974). Several experimenters have ascribed the learning deficit
which follows cingulate lesions to an enhanced fear response. However,
Kimble and Gostnell (1968), using two different behavioral measures,
failed to find any support for this hypothesis. Lubar, Perachio,
and Kavanagh (1966) suggested that incidental damage to the visual
cortex could account for the deficits following cingulate lesions,
but adequate control lesions (e.g., Kimble, 1968) and lesions which
avoid damage to the visual cortex (Trafton, Fibley & Johnson, 1969)
are adequate proof of the cingulate's role in the observed impairments
(Isaacson, 1974). The essential findings on the effect of cingulate
lesions on memory in the rat and cat are straightforward: such
lesions impair the ability of these animals to acquire active
avoidance conditioning (Peretz, 1960; McCleary, 1961; Thomas &
Slotnick, 1962; Lubar & Perachio, 1965; Kimble & Gostnell, 1968;
Trafton et al., 1969).
A number of relatively complex deficits have been noted after
cingulate damage which resemble symptoms seen in human amnesia
syndromes. These include disruption of the orderlysequencing of
behaviors (in nest building and maternal behavior, Stamm, 1955;
Slotnick, 1967), deficits in the temporal ordering of responding

83
(in runway problems and bar-press alternation: Barker & Thomas, 1965,
1966; Barker, 1967), and the failure to exhibit behaviors which were
indicative of opium addiction (Marques, 1971). The last, especially,
is supportive of the hypothesis voiced by some authors that cingulate-
lesioned animals are unable to anticipate the emotional consequences
of their behavior for both rewards and punishments (Glass, Ison,
& Thomas, 1969; Isaacson, 1974).
Anterior cingulectomy has been termed the psychosurgical
"operation of choice" for the treatment of severe obsessional and
anxiety disorders (Lewin, 1961; Whitty, 1966). Whitty noted that
one of the long-term effects of cingulectomy was "relative neglect
of the impact of external events." Finally, Pechtel, McAvoy,
Levitt, Kling & Massermann, (1958) concluded that lesions of the
cingulate gyrus in humans resulted in, among other things, "amnesia
for previous learning" and "impairment of new learning skills."
Discussion
The failure to transfer learning between hemispheres seen in
cerebral commissurotomy preparations (animal and human) indicates that
the storage of memories is a neocortical function (Geshwind, 1965).
Generalized memory disorders, on the other hand, are only produced
by bilateral damage to the diencephalic structures of the hippocampal
system. It is evident that these structures in the circuit of
Papez form part of a functional system which monitors the environ
ment, matches incoming stimuli with representations of the organisms
previous experience with similar stimulus configurations, activates
the organism in the presence of potentially significant stimuli,

84
and finally, causes pertinent information concerning that stimulus
to be presented to conscious awareness. It appears that there are
two such systems in the human brain, each specialized to deal with a
different type of information. The first, lateralized to the right
hemisphere, performs the functions listed above with experiential
data. The second, operating in the left hemisphere, deals with
language and verbal concepts which may be based on gesta!ten that
are assembled and stored in the contralateral system. All of the
functions noted above are impaired, to a greater or lesser extent,
in various manifestations of the amnesic syndrome.
The evidence suggests that some learning does not take place in
amnesia victims, that is, memory traces are stored. However, the
amnesic subject has difficulty gaining access to those memory
traces when they are needed. More specifically, they are unable to
recognize and select the appropriate memory trace in a given situation
from the set of available traces. Access to the proper trace is
facilitated with adequate cueing. It appears that the function of
the hippocampal system is to assure the activation of appropriate
memory traces based on the requirements of the situation. In the
present formulation, significant (experientially based) memory traces
have been designated by the superordinate term "generalized expecta
tions." It appears that the hippocampal system is responsible
for the generalizing of these expectations from one situation to
another, similar, situation.
In lower forms, where survival is dependent on instincts, the
identification of a significant stimulus may culminate in the release

85
of species-specific behaviors that are organized at a subcortical
level. The elicitation of unlearned behavior sequences (related to
feeding, fighting, fleeing and reproduction) by hypothalamic
stimulation is well established and seems to be related to programs
stored in the midbrain or brainstem (Isaacson, 1974). In humans,
however, cortical functions are more important in determining the
behaviors that insure survival.
Papez's circuit is an "anatomical identity which connects the
temporal and cingulo-frontal cortex of both hemispheres" (Barbizet,
1963). Given the recently identified massive interconnection of
the cingulate gyrus with the inferior parietal lobule (IPL), Papez's
circle may be seen as forming a part of a larger cortical circuit
(see Fig. $). As such, it is in a position to directly modulate
the activity of the two areas of the brain that are identified with
the highest levels of information processing: the specifically
human tertiary association areas of the prefrontal lobe and the IPL.
Information flow within this cortical-hippocampal circuit
might be seen as follows: raw sensory data enter the system in the
primary projection areas and are processed in the secondary association
areas of the post-central cortex; as this information acquires meaning
it is passed on to the temporal cortex and into the hippocampal
circuit; here the meaningful bits are translated into a code that
emerges in the cingulate cortex as memory indexing information; this,
in turn, is referred to the IPL where it facilitates associations
that fine-tune the continuing input into the system. This process
continues until an adequate match is made and recognized at the

t
TEMP CTX
HPC MB
Ant. Thai. CING
>IPL
00
CT>
Papez's circuit
Figure 3. The position of Papez's circuit within a larger cortical circuit. (TEMP CTX -
temporal cortex; HPC hippocampus; MB mammillary body; Ant. Thai anterior nuclei of the
thalamus; CING cingulate gyrus; IPL interior parietal lobule.

87
level of the temporal cortex. This match would take the form of a
finished "gestalt" in the experiential system and a formal "concept"
in the verbal system. With the appearance of such a match the
temporal component would terminate the search process and relay the
product to the cdntralateral system. Thus, the recognition of
a gestalt might initiate the search for a related verbal concept or,
conversely, a verbal formulation may trigger a scan for pertinent
experiential associations. The description of these two complementary
systems seems to provide adequate explanation for the qualitatively
different types of memory which were described by Breuer:.and Freud
and by Rapaport, respectively as noted at the beginning of this
section.
Substantial support for the model described above may be found
in a situation where hippocampal activity is induced from within
the limbic system (rather than from the neocortex, as was the case
in Penfield's experiments). Such is the case when amygdala stimula
tion produces after-discharges in the hippocampus. Halgren (1981)
reviewed the effects of amygdala stimulation in conscious humans and
noted that such stimulations sometimes produces "complex formed
hallucinations, sometimes complete scenes as in a dream or vivid
recollection and sometimes more vague, apparently similar to an
intruding thought . and illusions of familiarity (deja vu)"
(p. 345). Halgren suggested that it is the activation of distant
normal tissue subsequent to amygdala stimulation which produces the
resulting mental phenomena. He cites evidence that

88
Amygdala stimulation seldom evokes a mental phenomenon
unless it also evokes an after-discharge . most
amygdala stimulations, even if they evoke an after
discharge (AD), do not evoke any reported mental phenome
non . thus, there is no direct connection between
amygdala activity and hallucinatory experiences. . .
Simultaneous recordings from multiple brain areas
indicate that amygdala ADs seldom remain localized.
Initial spread is to the ipsilaterla hippocampus and
hippocampal gyrus. AD may remain confined to these
structures, in which case no mental phenomenon is
necessarily evoked. Further spread is to the ipsi-
lateral limbic cortex (orbital, insular and cingular)
and diencephalon (especially the anterior nucleus of
the thalamus, but also to the centre median, pul vinar
and dorsomedial nuclei). ADs seldom spread to the
neocortex, which may however desynchronize." (pp. 396-
397, emphasis added)
These findings are congruent with the present formulation, which
would interpret the appearance of "complex formed sensory hallucina
tions" following the stimulation described above as the result of
hippocampal output evoking experiential memory traces in the posterior
association cortex by way of the anterior thalamus and cingulate
cortex. The appearance of identical "experiential hallucinations"
following temporal lobe stimulation and limbic system ADs supports
Penfield's contention that final common pathways in the temporal
lobe may produce activity in the hippocampal system which utlimately
results in the appearance of mental phenomena.
The hippocampal system is directly involved in the mechanics
of learning. Hippocampal activity seems to assure that the organism
attends to novel stimuli and sets the conditions which permit changes
to be encoded into the neural models which form the centeral represen
tations of those stimuli. Pribram and McGuinness (1975) suggested
that such changes in neural represenations "may be conceived as

89
changes of state, set, or 'attitude'" (p. 132). Such experiential ly
based alterations in the disposition of the organism relative to a
stimulus object, situation, or event provide an operational definition
for the generic term "generalized expectation" as used in the present
formulations. This type of memory appears to be the "idea" which
Breuer and Freud believed to be of central importance in the etiology
of psychoneurosis. (It will be suggested that it is precisely these
generalized expectations, in this memory system, which will provide
the answer to the question posed in the introduction: "What is to be
changed in the process of psychotherapy?")
In addition to their mnemonic defects, amnesia victims with
damage to the hippocampal systems also present anomalies in their
affect and arousal. The latter may be traced to the failure of the
hippocampus to trigger cortical activation (via the RAS) as it
normally does in conditions of uncertainty or significance. The
former seems to reflect the disruption of hippocampal modulation
of the emotion/motivation processes by way of its interactions
with other limbic structures and with the prefrontal areas (via the
dorsomedial thalamus).
Interfaces and Interactions of the Monitoring
Motivating, and Mobilization System
Pharmacological research with animals and humans has implicated
the two classes of biogenic amines in the modulation of fundamental
brain processes. The catecholamines dopamine (DA) and its metabolite
norepinephrine (NE) have been associated with cognitive function and
dysfunction (e.g., schizophrenia) while affective disorders seem to

90
involve an interaction between NE and the indolamine serotonin
(5-HT). Histological researchers have identified six major mono
amine systems in the rat brain (Fxe, 1965; Fuxe, Hamberger &
Hokfelt, 1968).
The niqro-striatal DA system originates in cell bodies in the
substancia nigra whose axons extend through the lateral hypothalamus
to terminate on cells in the caudate nucleus. This tract is known
to regulate the activity of the ancient extrapyramidal motor
control system.
The meso-limbic DA system originates in the ventral tegmental
area of the pons and projects mainly to the septal area and related
nuclei in the limbic forebrain. Dysfunction in this system is
assumed by many to be the source of schizophrenic disorders.
The meso-cortical DA system is less well defined but appears
to include projections from the ventrotegmental to the frontal
cortex and from the substancia nigra to the anterior cingulate
cortex (see Meltzer, 1979).
The ventral NE system originates in the reticular formation
(medulla and pons) and ascends through the median forebrain bundle
(MFB) to terminate on cells throughout the hypothalamus and amygdala.
Stein, Wise and Berger (1972) suggested that this system primarily
regulates motivational activities.
The dorsal NE system arises in the locus ceruleus in the pons
and may supply up to 70% of the NE in primate brains (Redmond,
1979). The axons in this system ascend through the MFB and give
off branches to the hypothalamus, the hippocampus and amygdala,

91
the septal area (which receives the bulk of these terminals), the
anterior-ventral thalamus, the cingulate gyrus and the neocortex.
Stein and Wise (1971) suggest that this dorsal NE system is involved
in regulating cognitive activities.
The ascending serotonin (5-HT) system originates in the median
raphe' nucleus which is situated in the core of the reticular
formation and receives collateral branches from the sensory nerves.
5-HT cells in the raph^; have terminations throughout the central
gray of the midbrain. Axons from the raphe' also rise in the MFB and
distribute to the same areas as the dorsal NE system, with the
addition of terminals in the basal ganglia and more widespread
distribution in the neocortex. This system seems to be most
directly involved in regulating tonic arousal.
Brodie and Shore (1957) suggested that NE and 5-HT exert
opposing effects that modulate a variety of CNS functions (e.g.,
sleep, appetite, sexual drive). It appears that these parallel
systems control behavior by reciprocal action in balanced systems.
While functional specificity is determined by the neural circuitry
involved at a given anatomical level, the cumulative effects of
specific functional outcomes produced a preponderance of one or
the other transmitter seem to be consistent and to result in
coordination across levels. It appears that a preponderance of
the indolamine 5-HT, for example, results in the release of tonic
mobilizing energy at the brainstem level, the suppression of
ongoing behavior at the hypothalamic level, the identification of
"negative reinforcement" at the limbic system level, and

92
cognitive patterns consistent with the goals of orienting at the
neocortical level. An opposite pattern is evident when the catechola
mines predominate. The specific systems involved in these functions
will be described in the following sections.
The Biochemistry of Emotion, Motivation, and Learning
The fundamental prerequisite for adaptive behavior is a system
to sort environmental inputs and pair them with appropriate responses.
The more primitive life forms must rely on the experience of their
species, accrued over evolutionary history and trasmitted genetically,
to accomplish these tasks. The adaptability of such forms is
strictly limited by the repertoire of sensory discriminations and
motor response sequences they are programmed to perform. Kety (1972)
noted that adaptability would be greatly enhanced if the organism
could learn to determine and preserve those sensory, evaluation, and
response patterns which, in the individual's own experience, pro
vided the ultimate survival advantage. He suggested that neuronal
and neurochemical mechanisms which utilized constant reference to
a small number of inborn values or states would allow the nervous
system to become increasingly elaborate and effective in performing
these functions. Kety postulated that only three such inborn states
should be required: arousal, pain, and pleasure. He proposed
that these mechanisms would interact to produce an emotional state,
consisting of increased arousal and the activation of particular
circuits, with a resulting drive to approach or avoid the confronting
stimulus. Arousal would accompany both pleasure and pain but might
occur in the absence of either in response to novel stimuli or to

93
inputs either genetically or experientially endowed with significance
to the organism. The emotional state would be accompanied by the
"intercerbral release of a trophic neurogenic substance . .
[which would permit] transcription into more permanent form . .
to present and immediately preceding states of neuronal activation
where the outcome had been significant for the organism" (Kety,
1972, p. 71).
Simply put, the results of the organism's adaptive efforts (i.e.,
positive or negative reinforcement) might produce a characteristic
biochemical signature within these circuits which reflected the
consequent affective status of the organism in that situation. These
data could then be encoded along with the memory of the situational
context. If that same stimulus configuration was confronted again
at a later time the encoded emotional information would cause the
biochemical pattern to be replicated in these circuits, the effect
of which would be to recreate the original emotional state to
motivate the organism appropriately based on the original experience.
Extensive work by Stein and his colleagues indicates that the
circuits involved include a noradrenergic median forebrain bundle
reward/behavior-release system which is antagonized by the seroto
nergic periventricular system (PVS) which functions as a punishment/
behavior-suppression mechanism. Stein, Wise and Berger (1972)
summarize experimental evidence which implicates these systems in
the release and suppression of unlearned behavior:
Briefly, electrical stimulation of the median fore
brain bundle . elicits species-typical consummatory
responses, such as feeding and copulation, which

94
produce pleasure and permit the satisfaction of basic
needs. Similar effects are produced pharmacologically
by potentiation of the noradrenergic (or blockade of
the serotonergic) activity of the median forebrain
bundle. On the other hand, electrolytic lesions of
the median forebrain bundle cause severe deficits in
goal-directed behavior and the loss of consummatory
reactions; again, similar effects are produced by
pharmacological blockade of noradrenergic function or
potentiation of serotonergic function. (Stein et al.,
1972, p. 82)
These authors suggest further that the NE system produces positive
feedback which facilitates ongoing behavior that was rewarded in the
past and the 5-HT system produces negative feedback which terminates
behavior that was unsuccessful or punished. Experimental evidence
supports their contention that these systems also mediate learned
behavior. Wise, Berger, and Stein (1970) demonstrated that drugs
which deplete central stores of serotonin, (and drugs which produce
blockade of serotonergic receptors) significantly reduced a shock-
induced elevation of brain serotonin levels significantly increased
the conditioned behavioral suppression. None of the drugs effected
the behavior of unshocked control animals. The complementary role
of NE was evident in a study where monkeys received bilateral
stimulation or lesions of the locus ceruleus (the source of the
dorsal NE system). Redmond (1979) reported that bilateral locus
ceruleus stimulation elicited behaviors which were associated with
increased fear. Animals with bilateral locus ceruleus lesions
subsequently rose in their social hierarchies because they lost their
fear of previously dominant animals (and humans): they showed
increased motor activity, increased social aggressiveness, and
decreased retreat from threat from pre-lesion levels while retaining

95
normal responses to actual pain. Arbuthnott, Fuxe, and Ungerstedt
(1971) presented evidence that all of the "positive reinforcing"
(self-stimulation) points are within the ventral NE system. Stein
and Wise (1969) demonstrated (with permanently implanted push-pull
cannulae) that "rewarding" self-stimulation of the MFB caused
increased synthesis and turnover of NE and a marked increase in the
release of radioactively labeled ME into perfusates of the amygdala
and hypothalamus.
Isaacson (1974) suggested that the almost total'lack of fear
and aggression resDonses in animals after bilateral amygdalectomy
reflects a decrease in the ability of enviromental stimuli to
elicit reactions from the organism. Increased 5-HT activity has
been associated with increased fear and decreased aggression while
decreased NE activity has been associated with decreased fear and
increased aggressiveness. It appears that in lower forms these
systems mediate the release or suppression of instinctive behavior
at the level of the hypothalamus. It follows from all of the data
presented above that the ability to learn from experience, a
prerequisite for self-determined behavior, involves an integration
of emotional experience and memory which is based on an interaction
of NE and 5-HT in the amygdala.
Emotion, Amygdala Circuits, and Memory
The role of the amygdala in emotional processes, established
by Kluver and Buey in 1933, has been assumed to be affected through
this structure's close relationship with the hypothalamus. The
amygdala seems to direct behavior toward biological goals (Halgren,

96
1981) and related to survival needs, including defensive and aggressive
behaviors, sexual activity, and feeding (isaacson, 1974). In lower
forms these processes might depend on a simplified (instinctive)
form of memory in which stimulus and response are yoked (Pribram &
McGuinness, 1975). In addition to mediating emotional states the
arnygdala is involved in the analysis of reinforcement contingencies.
Amygdala lesions have been shown to produce impaired recognition of
stimuli associated with rewards (Weiskrantz, 1956; Schwartzbaum,
Thompson & Kelli cut, 1964; Jones & Mishkin, 1972) and inability to
respond appropriately to changes in the magnitude of rewards
(Schwartzbaum. .1960).
The more recent discovery of strong anatomical interconnections
between the amygdala and the neocrotex (Aggleton, Burtin & Passingham,
1980; Herzog & Van Hoesen, 1976; Turner, Mishkin & Knapp, 1980)
allows for the involvement of this structure in higher levels of
mental processing and memory. Kessner (1981) suggested that long
term memory "consists of a set of bundle of traces, each representing
some attribute or feature of a learning experience." He postulated
that "the amygdala mediates the encoding, storage, and retrieval
of specific emotional attributes often associated with reinforcement
contingencies in specific situations" (p. 332). In a test of this
hypothesis Kessner and Conner (1974) showed that five seconds of
bilateral subseizures level electrical stimulation of the amygdala
(which disrupts the normal functioning of that structure) following
footshock in a passive-avoidance paradigm resulted in amnesia for
that training experience. Implanted and unoperated controls without

97
such stimulation demonstrated excellent memory for the learning experi
ence. Kessner interpreted this result as the failure to encode the
negative affect associated with the experience.
In order to discriminate the relative contribution of the
amygdala and the hippocampus to these phenomena Baker, Kessner,
and Mi chai (1981) took advantage of the fact that suitable reminder
cues often serve to facilitate existing memories and thus enhance
recall. Rats with electrodes implanted in the amygdala or hippo-
capmus and unoperated animals received a single training trial with
footshock (FS) while licking a water tube in a goal box. Retention
was evaluated 24 hours later as an increase in latency to enter the
goal box and lick the tube at least ten times. Following this first
retention test four of the groups received a reminder cue (non
contingent FS) in a different enviroment, followed immediately by
hippocampus or amygdala stimulation or no stimulation. Twenty-three
hours later all groups were tested a second time for retention of
the passive-avoidance learning in the original apparatus. Operated
and nonoperated controls that received a reminder cue but no brain
stimulation exhibited a marked increase in latency on the second
(relative to the first) retention test, demonstrating that the
reminder cue effectively enhanced retention of the aversive experience.
Conversely, operated and nonoperated controls which did not receive a
reminder cue showed a decrease in latency on the second test,
indicating that some extinction had taken place due to re-exposure to
the original apparatus during test number one. Animals whose amygdala
function was disrupted by electrical stimulation following the reminder

98
cue showed a decrease in latency on the second retention test as compared
to the first, and thus resembled the groups that never received a cue.
By contrast, animals which received hippocampal stimulation following
the reminder cue exhibited an increase in latency, similar to the
control animals which benefited from the reminder FS. Kessner (1981)
concluded from these results that "amygdala stimulation disrupted the
encoding of negative affective attributes associated with the reminder
FS negating its efficacy in enhancing subsequent retention. In
contrast, hippocampal stimulation had only a small effect and did
not totally disrupt the efficacy of the FS reminder" (p. 336). It
seems, therefore, that the amygdala is selectively involved in the
memory of emotional experience, at least for the negative affect
associated with passive-avoidance learning situation.
Kessner (1981) reviewed several studies which suggested that
the amygdala was not involved in the encoding of positive affective
attributes. However, he cites evidence that amygdala stimulation
does appear to disrupt retention when there is a magnitude of reward
effect (i.e., fewer errors for the expectation of a greater magnitude
of reward). Kessner concluded from the studies reviewed above that
"the amygdala encodes, stores, and retrieves both positive and
negative emotional attributes of a specific memory, providing the
input is of sufficient intensity to elicit a relatively strong
emotional reaction. . Different neural systems (e.g., hippo
campus) would encode other attributes (e.g., enviromental context)
of the same specific memory" (1981, p. 340).

99
Mishkin and Aggleton (1981) reviewed data which demonstrate
that the amygdala receives highly processed sensory information
from all of the secondary and polymodal association areas of the
posterior neocortex in addition to its reciprocal connections with
the hypothalamus. They suggested that these connections provide a
mechanism for complex stimuli to become intergrated with, and
later, to evoke emotional responses which are organized at the
level of the hypothalamus.
Smythes (1966) suggested that the hippocampal system may lay
down memories and the amygdala system may determine what memories are
to be laid down. While this may overstate the case somewhat, it
is clear that these two functional systems work synergistically to
provide the organism with pertinent information from its past
experience which is caoable of motivating and facilitating appropriate
responding based on the requirements of the situation. The septal
area appears to play an essential role in the interaction of the
amygdalar and hippocampal systems and may integrates their activity
with that of the brain's arousal systems.
The septal area is interposed anatomically and functionally
between the amygdala and hippocampus (via the stria terminal is
and fornix, respectively) and has a reciprocal relationship with
both (Swanson & Cohen, 1976). The septum is connected with the
brainstem reticular formation (via the MFB) and is in a position
to modulate the activity of the ARAS through its connection with
the habenula (via the stria medularis), which is a station between the
RF and ILTN. (All of these anatomical relationships are illustrated

100
in Fig. 2). The septo-hippocampal axis has received the most
attention to date.
Based on their micro-electrode recording studies Edinger and
Siegal (1976) suggested that
Activity in adjacent groups of hippocampal efferent
pyramidal cells could result in the discharge of a
septal neuron. The activation of this neuron will
excite the inhibitory interneuronal network [within
the septal area] to block activity in other portions
of the septum. The septum therefore acts as a
filtering mechanism to Dermit transmission of informa
tion only from that portion of the hippocampus whose
pyramidal neurons are most active at that point in
time. Information transmitted from less active areas
of the hippocampus will be suppressed by the inhibitory
network of the septum, (p. 249)
The septal area appears to have a similar effect on the interneuron
population of the hippocampus. Lynch, Rose and Gall (1977)
conclude from their anatomical investigations that "septal contacts
on [hippocampal] interneurons modulate their activity and thus
allow the 'biasing,' in effect, of the response of the hippocampal
neurons to their excitatory inputs . the septal projections may
serve to modulate the response of the hippocamous to the activa
tion of its massive afferent input from the entorhinal cortex"
(p. 17). Thus, the septal area is in a position to integrate the
activity of the brain's memory, emotion, and arousal systems and
the available evidence supports the suggestion of such a role
for this structure. This arrangement would permit the amygdala,
acting through the septal area, to focus the hippocamoal system's
memory search on associations which have immediate (emotional)
significance for the organism.

101
The identification of emotionally significant stimuli in
phylogenetically lower forms may result in the release of species-
specific responses at the level of the hypothalamus, but in higher
forms the neocortex is more involved in motivating behavior. It is
assumed here that the more recently developed systems which provide
the substrate for higher mental processes must be programmed in
some way to insure survival. It appears that biochemically mediated
mechanisms may have evolved in parallel over the more ancient control
circuits to perform this function. Stein and Wise (1971) suggested
that the dorsal NE system was involved in the regulation of cognitive
processes. Several lines of evidence, taken together, strongly
suggest that the cingulate cortex and NE participate in the integra
tion (encoding and decoding) of emotional experiences with memory
to facilitate adaptive functioning at the highest levels:
1. Based on the review in the previous section it was hypothe
sized that the cingulate cortex is the final station in a hippocampal
memory circuit which performs memory indexing functions.
2. The cingulate cortex is interposed anatomically between the
prefrontal lobe and the inferior parietal lobule (I PL) and is
thus in a position to coordinate the actions of these two highest
centers of information processing.
3. Animals with cingulate lesions are unable to acguire
avoidance learning (see previous section). Ison and Thomas (1969)
concluded from their studies that cingulate lesions "dissociated
conditioned emotional goal responses from instrumental performance"
(p. 17). Ward (quoted in Isaacson et al., 1971, p. 232) reported

102
that monkeys with cingulate damage treat their companions like
inanimate objects, which he interpreted as a "loss of social con
sciousness."
4. fiiesser (1967) noted that the content of the complex
hallucinatory experiences reported by Penfield's subjects a'fter
electrical stimulation of the right temporal cortex was not
surprising: "What is surprising is only the vividness of the
imagery" (Ntesser, 1967, p. 169). Whitty (1966) reported that,
for one to three days postoperatively, patints who had undergone
anterior cingulectomy experienced "increased vividness of thoughts
and images so that a subjective difficulty is experienced in
distinguishing between thoughts and exterior happenings" (p. 404).
5. The cingulate cortex receives projections from the brain
stem ascending 5-HT and dorsal NE systems and has the highest
density of NE terminals in the cortex (Fuxe et al., 1968).
6. Redmond and his colleagues (see Redmond, 1979) reported
that monkeys with bilateral locus ceruleus lesions showed an
"absence of emotional responses to threat" (p. 158). Encephalitis
lethargica is a disease which affects the brainstem and the peri
aqueductal gray area (which contains the locus ceruleus) in
particular. Penfield (1975) noted that postencephalic patients
often developed an obsessive-compulsive syndrome: such patients
were "preoccupied with compulsive thoughts, compulsive utterances,
and compulsive behaviors to an extent that often caused severe
disability" (p. 98).

103
7. In obsessive-compulsive illness certain thoughts and/or
actions acquire an inappropriate level of significance for the
individual and social functioning is impaired. Cingulectomy is
the psychosurgical "operation of choice for the classical syndrome
of obsessive neurosis;" after surgery "the obsessive thoughts
retreat gradually into the background and the drive behind them
lessens" (Lewin, 1961). Humans who have undergone cingulectomy
show some disinhibition of behavior (but decreased aggressiveness)
and seem to neglect the emotional impact of external events
(Whitty, 1966).
Discussion
It is evident that the cingulate cortex is involved in the
coordination of emotional, attentional and cognitive processes.
It appears that this structure modulates the infusion of emotional
significance into ongoing perceptual and/or associational operations
and influences the allocation of neural mobilizing energy, thus
permitting normal emotional reactivity and appropriate inhibition
of behavioral and cognitive activity. While it is not possible at
present to sort out the complexities of 5-HT/NE interactions
across brainstem, cortical, and right/left dimensions, the available
evidence suggests that the cingulate gyrus evolved as a cortical
extension of those brainstem control mechanisms. In the context
of functional systems, the cingulate cortex is in a position to
encode emotional data from the amygdala-orbitofrontal circuit into
neural models being "indexed" by the hippocampal memory system.

104
Biochemical and Electrophysiological Aspects of Cortical Mobilization
Processes
It appears that the NE and 5-HT systems interact to control the
energizing output of the reticular formation. At the brainstem
level stimulation of the raphe' (5-HT) nuclei produce calmness
and EEG patterns similar to those of normal sleep, while damage to
these nuclei result in increased motor activity (see Fawcett, 1975).
The raphe' nuclei are thought to be the primary site of action in
morphine-induced catalepsy (see Broekkamp & Lloyd, 1981). Kety
(1972) summarized pharmacological evidence which implicates NE in
the modulation of arousal and sedation. Evidence to be reviewed
below suggests that, at the neocortical level, the 5-HT system
supports a pattern of indiscriminate cortical arousal which facili
tates gross sensitivity at the expense of complex analysis. Another
system mediated by DA ssems to permit the focusing of attention and
may be the means by which the frontal lobes recruit and organize
cognitive activities in the post-central cortex.
Surgical ablation of the prefrontal lobes is sometimes performed
to relieve otherwise intractable pain. This operation is not a
specific cure for pain, however, but a cure for "suffering,"
the source of the pain and its threshold remain unaltered. It is
the person experiencing the pain that is altered by this psycho-
surgical intervention. Petrie (1952; 1960; 1967) found that the
changes in pain tolerance were paralleled by striking differences
in the subjectively perceived intensity of sensation other than
pain. She suggested that the personality and perceptual tendencies

105
of a person before and after prefrontal lobotomy exemplify the
extremes on a continuum of "perceptual reactance" which can be
seen in the normal population. Petrie (1960; 1967) administered her
kinesthetic figura! aftereffects test to a variety of normal and
abnormal populations and demonstrated that, after repeated tactile
stimulation of the fingertios, some people consistently overestimate
(augment) the size of a standard stimulus while others consistently
underestimate (reduce) its size. Reducers tolerate pain well and
augmenters do not. Petrie argues that augmenters and reducers are
displaying basic differences in their methods of perceiving the
environment.
Several investigators studying visual evoked potentials to
light flashes have noted that some subjects showed a reduction in
the amplitude of these cortical responses as stimulus intensity
increased, a phenomenon which Kamphuisen and van Leeuwen (1968)
called "paradoxical diminution." Buchsbaum and Silverman (1968)
reported a significant correlation between flash visual evoked
responses and perfromance on Petrie's kinesthetic figura! after
effects test: augmenters showed an increasing cortical response as
stimulus intensity increased while reducers showed a leveling off
or decrease in the amplitude of their cortical responses. This
finding was replicated by Spilker and Callaway (1969) using
different stimuli and different electrode placement, von Knorring,
Espvall and Perris (1974) demonstrated empirically that reducers
classified by visual averaged evoked responses had significantly
higher pain thresholds and pain tolerance levels than augmenters.

106
By altering the cortical evoked response paradigm so that the
second of a pair of responses could be expressed as a function of
the first Pribram (1967) was able to make imoortant inferences
about the organization that is imposed on incoming information
When a double click or a double flash is used to evoke
a neural response, the amplitude of the second of the
pair of responses serves as an indicator of the duration
over which a part of the system is occupied in processing
the first of the pair of inputs. A depression in the
amplitude of the second of the responses thus indicates
a longer recover--a longer processing time for a signal
within the channel. Such an increase in processing
time effectively desynchronizes the channel to repetitive
inputs: fewer fibers are available for processing
any given signal in the series. Prolongation of re
covery thus reduces redundancy in the channel. At any
moment more information can be processed. . Thus
the rate of information processing is enhanced. (Pribram,
1967, p. 460)
Augmented redundancy in information processing channels would mean
that more neurons were available to fire in a stimulus-linked
fashion. Thus, the sensitivity of the organism would be increased
(e.g., decreased pain tolerance, increased subjective experience
of pain). Since information tends to be "chunked" in this mode
there would be a decrease in the focusing of attention on details
and decreased articulation of experience. Conversely, reduced
redundancy in information processing channels would leave fewer
neurons available to fire in a stimulus-linked fashion, thus
reducing the sensitivity of the organism; bits of information would
tend to be separated in this mode permitting increased focusing
of attention on details and increased articulation of experience.
It will be noted that the augmentation mode would favor the performance
of cognitive tasks associated with the orienting response (i.e.,

107
stimulus identification) while the reduction mode would facilitate
the sequential information processing and abstraction of detail
associated with problem-solving and language processes.
A novel stimulus (information plus uncertainty) "disinhibits
and gains access to large number of neurons indiscriminantly
throughout the neocortex" (Kety, 1972, p. 71). Pribram and
McGuinness (1975) reviewed evidence which indicates that the source
of the generalized cortical arousal associated with orienting
is to be found in the serotonergic nuclei of the brainstem. The
searching and sampling of the orienting process is terminated
when the stimulus is identified and registered in memory. At
this point uncertainty is resolved. If the situation registered
in the neuronal model of the identified stimulus has no built in
"demand" characteristics the organism may habituate to the stimulus
and proceed to ignore it. However, if the neuronal model generates
important expectations related to the stimulus, then cortical
activation systems which facilitate perceptual and motor readiness
to respond will be engaged. In contrast to the indiscriminate
cortical arousal associated with orienting and perceptual augmenta
tion, these activation systems produce a highly organized pattern
of cortical activity. Pribram and McGuinness (1975) reviewed
evidence which indicates that the mechanism and pathways involved
in controlling these activation systems are dopaminergic, and that
their operation is reflected in the phenomenon of the contingent
negative variation, or "expectancy wave."

108
The characteristics of the contingent negative variation
(CNV) were reviewed by Cohen (1974). Briefly, the CNV is a special
form of cortical evoked response consisting of a gradually developing
wave of negative potential which originates in the frontal lobes and
sweeps posteriorally over the cortex whenever there is a stimulus-
response contingency or expectancy built into a situation. This
wave becomes abruptly positive when the required perceptual or
motor response is executed. The amplitude of the signal is directly
related to the reinforcement ratio, with peak amplitude at 100%
reinforcement. Reaction time is shortest following high amplitude
CNVs. Perhaps the most interesting characteristic of the CNV is the
fact that it responds to verbal instructions: the waves disappear
when the instruction which produced the expectancy is countermanded,
and fail to appear when the subject is told in advance that the
contingent signal will not be forthcoming. The association between
perceptual reduction processes, the CNV, and dopamine is strengthened
by the finding that evoked potential reducers have increased levels
of homovanillic acid (a dopamine metabolite) in the cerebral spinal
fluid (Gottfries et al., 1974).
The Functional Elements of the Personality Structure
It has been assumed here that the phylogenetic transition from
instinct-based responding to self-determined behavior required the
evolution of new mechanisms to insure the survival of the individual
and the species. Specifically, it was hypothesized that automatic
neural systems were needed to (1) Monitor the environment for
significant stimuli (b) Motivate the organism to respond in the
presence of such stimuli-, and (c) Mobi 1 ize the appropriate

109
psychological operations to determine the form of that response.
It was postulated that these mechanisms would be arranged in
functional systems (as defined by Luria, 1973a) and that these
central organizing factors would form the infrastructure of
personality. The brain mechanisms which appear to satisfy these
requirements will be outlined below. It will be proposed that
these systems form the basic functional elements of the personality
structure. Supporting evidence from animal studies of learning
and memory will be presented at the end of this section.
The Problem-Solving /Response-rGeneratinq System
It is evident that the psychological operations which generate
considered responses in a given situation take place in the neural
tissue of the association areas in the post-central cortex of the
left cerebral hemisphere. These mechanisms normally have access
to, and employ the products of, mechanisms lateralized in homologous
areas of the right hemisphere. Both of these areas solve problems
through a process of categorization, but they arrive at their
solutions in different ways. The right hemisphere is specialized
to perceive overall patterns in input; it solves problems through a
convergence of information, synthesizing basic units into meaningful
gestalts. The left hemisphere shares these spatial abilities to some
extent, but they have been superseded by more complex skills. The
dominant hemisphere has been genetically prepared to analyze
information along a temporal dimension and this ability allows
mental activity to escape from the bounds of the inmediate spatial
context. This freedom, in turn, permits the mental manipulation

no
of externally provided symbols and aids (e.g., language and rules
of granmar, mathematical tables and formulas). The sequential
operations involved in abstract reasoning, logical analysis, and
propositional speech are made possible by the temporal acuity of the
left hemisphere. The orderly selection (programming) of these
operations depends on the integrity of the tertiary association
zones in the dorsolateral area of the prefrontal lobe.
Three types of information are required at these neocortical
levels in order to guarantee that they will perform their problem
solving tasks appropriately in survival situations. This informa
tion is provided by three functional systems that were outlined
above. Each of these systems has limbic, thalamic, and cortical
components (see Fig. 4).
The Memory System
Input to the hippocampal memory system consists of processed
sensory data and/or the products of ongoing cognitive activity.
This information is passed from the post-central association areas
to the temporal lobe into the entorhinal cortex. Hippocampal
output emerges in the cingulate cortex as memory indexing information
which is passed on to the I PL where it facilitates access to memory
traces which are associated with the input stimulus. These associa
tions finally emerge in consciousness as formal percepts or concepts
which provide context information and generalized expectations to
the problem-solving/response_generating systems. Signals repre
senting emotional information are passed from the amygdala to the
septal area where they modulate or focus hippocampal memory

Key to the schematic representation of the proposed mode) of the physiological substrate of personality. Most connections are reciprocal; arrows
indicate pathways emphasized in the text. Heavy black lines indicate memory indexing information from the hippocampal Monitoring System. Dashed
lines indicate emotion/reinforcement information from the amygdalar Motivating System. Dotted lines indicate activating information from the
reticular formation Mobilizing System. AMYG--amygdala; aniyg--contralateral amygdala; DM--dorsomedial thalamic nucleus; Pre-FR CTX--prefrontal
cortex; MOTmotor cortex; BG basal ganglia; TEMP CTX--temporal lobe cortex; AIS--autonomic nervous system; HT--hypothalamus; SEPT--septal area;
RF--reticular formation; ILTN--intralaminar thalamic nuclei (includes all nonspecific thalamic nuclei; Post-ASSH CTX--postcentral association cortex
(includes secondary association areas and inferior parietal lobule); SENS--primary projection sensory cortex; HPC--hippocampus; hpc--contralateral
hippocampus; MB--manmi 11 ary body; ANT--anterior thalamic nuclei; ClNG--cingulate gyrus; CC--eorpus callosum; AC--anterior comnissure; ufuncinate
fasciculus; vaf--ventral aipygdalofugal fibers; st--stria terminalis; sm--stria medullaris; sm-hab--stria medullaris via habenlua; mfb--medial
forebrafn bundle; fnx--fornix; MIT--mamillothalamic tract.
Figure 4. Schematic representation of the proposed model of the physiological substrate of
personality.

112
coding/decoding operations through a two-way septal-hippocampal inter
action. The memory indexing process is further modulated or pro-
grammed by processed motivational input from the prefrontal lobe
to the cingulate cortex. This system accomplishes three principal
tasks in the course of its operation: (a) it monitors incoming
stimuli and matches them against cortical representations of the
organism's previous experience with similar stimulus configurations;
(b) when a potentially significant stimulus configuration is
encountered the system activates the organism by switching on the
Mobilization System; and finally, (c) it causes pertinent informa
tion related to those stimuli to be presented to conscious awareness.
It appears that there are two such systems in the human brain,
each specialized to deal with a different type of information. The
first, lateralized to the right hemisphere, performs the functions
listed above with experiential data. The second, operating in the
left hemisphere, deals with language and verbal concepts. It is
probable that environmental stimuli trigger an initial search in
the experiential (right hemisphere) system. (It may be noted
that these right hemisphere processes would be unconscious and
that their products might be excluded from consciousness.)
The Motivating System
The Motivating System is centered on the amygdala. This structure
receives processed sensory information from all of the sensory
modalities and is interconnected with the entire prefrontal lobe
both directly (via the uncinate fasciculus) and indirectly (by
way of the dorsomedial thalamic nuclei). It also has strong

113
reciprocal connections with the hypothalamus (via the stria terminal is
and the ventral amygdalofugal fiber system) and with the septal
nuclei (via the stria terminal is).
The amygdala is involved in the integration of information
from the internal and external enviroments which relates specifically
to the survival of the individual and its ability to reproduce.
Based on such an integration, the amygdala attaches motivational
significance to previously neutral stimuli by associating those
stimuli with their internal consequences. The amygdala produces
the signals which permit the positive and negative attributes of
an experience to be encoded in memory through an interaction with
the hippocampus (via the septal area). When that memory trace is
reactivated on a later occasion the amygdala responds to its encoded
emotional and reinforcing attributes, computes the reinforcement
value of the present stimulus in light of the immediate internal
status of the organism, and participates in focusing the hippocampal
systems on significant memory traces. Qualitative and quantitative
motivational information is relayed from the amygdala to the orbital-
frontal cortex where it is translated into a subjective experience
(.e.g, fear, anger, anxiety, etc.) of appropriate intensity. The
tertiary association area in the Tdorsolateral prefrontal cortex
responds to this experience and will attempt to resolve it by
programming the post-central problem-solving mechanisms to formulate
a behavioral response to the stimulus. The amygdala may also
initiate activity in the sympathetic and parasympathetic control
mechanisms in the hypothalamus to prepare the organism physically

114
to deal with the stimulus and might activate the pituitary-adrenal
"rapid alarm" system.
The Mobilization System
The cortical mobilizing system is based on the brainstem
reticular formation (RF). Signals from this mechanism lower the
activation threshold of neurons it projects to. The activity of
the RF is modulated by biochemically mediated forebrain systems
whose descending influences reach the RF through the hippocampus
(HPC) and amygdala (AMYG). Ascending RF pathways pass through the
hypothalamus and septal areas and terminate on the nonspecific
thalamic nuclei (e.g., ILTN).
Descending influences from the prefrontal lobe modulate the
activity of the thalamic reticular system and in this way direct
the activation of specific cortical systems. This is accomplished
through an interaction between the nonspecific thalamic nuclei and
the thalamic association nuclei. In this manner the frontal lobes
are able to recruit and program cognitive operations in the post-
central cortex in accordance with intentions.
The Memory, Motivating, and Mobilizing systems supply the neo
cortex with memory indexing information, qualitative and quantitative
emotional data, and activating impulses, respectively. Each of these
categories of information is essential for normal functioning.
Damage within any one of these functional systems results in a
different syndrome of specific clinical deficits. The manifestation
of these deficits may vary with the location of the damage within
the system.

115
Each of these systems evolved from more primitive mechanisms
which regulated the emission (in the presence of releasing stimuli)
of unlearned (instinctive) behavioral sequences related to the
survival functions of feeding, fighting, fleeing, and reproduction.
In their evolved form these systems allow the decoupling of stimulus
and response, thereby permitting the neocortex to determine behavior,
but they continue to insure that an adequate response is emitted.
Learning and Memory: Animal Studies
The model indicates specific functional/anatomical mechanisms
which could account for the sometimes puzzling deficits seen after
different limbic system ablations in experiments performed by
physiological psychologists.
Animals with hippocampal lesions are sometimes the same and
sometimes different from normal animals in the performance of
tasks for appetitive rewards (Isaacson, 1974, ch. 5). Such
conflicting results have made it difficult to define a specific role
for the hippocampus in memory functions. Studies of human amnesia
victims (reviewed in the previous section) indicate that their
amnesic deficit reflects an inability to select the appropriate
memory trace from the set of available traces, although memories
continue to be laid down in the cortex. A naive animal (with hippo
campal damage) in a strictly controlled laboratory situation may
store only one memory trace (or "hypothesis") for that situation
and not be troubled by competing memory traces. There have been
many reports that animals with hippocampal damage are able to
learn a simple discrimination problem (Isaacson, 1974, ch. 5).

116
However, when the situation requires that the animal select from
more than one possible hypothesis, as in successive or simultaneous
discrimination problems, hippocampal animals are impaired relative
to controls (Kimble, 1961, 1963). Similarly, naiv hippocampal
animals can learn to bar-press for rewards under continuous rein
forcement (CRF) conditions but are impaired when they are transferred
to a partial or intermittent schedule (e.g., the DRL). Schmaltz
and Isaacson (1970) demonstrated that it is the change in schedules
that is debilitating: Hippocampal animals trained from the outset
using only the DRL contingencies were not impaired on that task.
Winocur and Mills (1970) found that previous experience impaired
the performance of hippocampal animals only when the preceding task
was related in some way to the new problem; unrelated training
did not interfere. Thus, similarity between situations actually
hinders hippocampal animals rather than helping them. Isaacson
(1974) interpreted such results as the inappropriate transfer of
strategies or the fixation of predominant behavioral dispositions,
but the present formulation would stress the hippocampal animals'
inability to select between competing memory traces because they
do not receive memory indexing information ath the cortex.
Five experimental paradigms have been used extensively in
investigations of the affect of limbic system manipulations on
learning and memory. In the passive avoidance task an animal is
punished for making a response (e.g., receives a shock while
drinking from an electrified water tube) and must learn to avoid the

117
punishment by withholding that response (Isaacson, Douglas, Lubar
& Schmaltz, 1973, note that the inhibition of a response is actually
an active process). In the one-way active avoidance problem the
animal must learn to actively avoid punishment (e.g., footshock
[FS]) by making a specified response (e.g., jump over a hurdle in
response to a conditioned stimulus [CS1 which signals an impending
FS). In the two-way active avoidance task the animal must jump
between the two compartments of a shuttle-box on alternate trials
in response to a CS in order to avoid punishment. Isaacson et al.
(1971), point out that this situation produces a conflict between an
active- and a passive-avoidance response: on any given trial the
animal must leave the compartment it is in to avoid the signaled
FS (active-avoidance) and must enter the compartment in which it
was shocked on the previous trial (passive-avoidance). In the
differential reinforcement of low rates of responding (DRL)
situation the animal must learn to withhold responses for a pre
determined amount of time after the last response before a response
will be rewarded. On the fixed interval (FI) reinforcement
schedule normal animals learn to stop responding in the first half
of the interval following a reinforced response and to slowly
increase their rate of responding in the second half of that
interval. Continued responding in the first half of the interval
is considered to be a perseveration error; higher than normal rates
of responding in the second half of the interval are considered
anticipatory errors.

118
Animals with hippocampal or septal area damage show identical
alterations in their performance of all of these tasks (Isaacson
et al., 1971). They are impaired in the acquisition of passive and
one-way active avoidance problems; they perform better than controls
on the two-way task; they show perseverative over-responding on the
FI operant schedule and fail to obtain reinforcements in the DRL
situation because of this over-responding tendency (Isaacson, 1974;
Grossman, 1976). Each of these will be considered in turn.
It was noted in the last section that electrically induced
disruption of amygdala function immediately after training in the
passive avoidance (PA) paradigm resulted in a learning deficit
for that task which mimicked the deficit found after amygdalectomy.
It was hypothesized that the deficit reflected the failure to encode
the negative attributes of that situation in memory. The evidence
indicates that the septal area and hippocamous participate in this
encoding process. Animals with septal lesions also show a deficit
in this task (McCleary, 1961). Grossman and his colleagues have
succeeded in selectively transecting the various connections of
the septum using a specially designed retractable wire encephalotome
(see the review by Grossman, 1976). They found no PA deficit
following transection of the fornix (FNX), the medial forebrain
bundle (MFB), or the stria medullaris (SM), but cutting the stria
terminalis (ST) duplicated the PA deficit seen after septal (and
after amygdalar or hippocampal) lesions. Thus, it appears that it
is the isolation of the amygdala from the septo-hippocampal inte
grating mechanism and the subsequent failure to encode the negative

119
attributes of an experience in memory which produces the PA
deficit (rather than "perseveration" or a "loss of inhibitory
control" as assumed by many authors, e.g., McCleary, 1966). That
the hippocampus (HPC) is essential in this encoding process is
indicated by the finding that animals trained on the PA task before
bilateral HPC ablation were not impaired in the retention of the
learning after surgery (Wishart & Mogunson, 1970), while such lesions
impair the post-surgery acquisition of that problem (Isaacson &
Wickelgren, 1962).
Transection of the FNX, but not of the ST or MFB, duplicated
the pattern of responding seen on the FI schedule after septal and
HPC damage, and partially replicated the disinhibitory affects of
such lesions in the DRL paradigm (Grossman, 1976). Grossman believed
that these results were "due specifically to a disruption of septo-
hippocampal connections" (1976, p. 405). The present formulation
would emphasize the role of the septum in modulating hippocampal
control of the Mobilization system via the RF. Septal lesions
block the theta rhythms normally seen in the HPC following and
alerting stimulus (Smithies, 1966). Failure of the HPC to inhibit
the RF would be expected to result in increased tonic mobilization
and "over-responding," even when such behavior is inefficient.
The puzzling superiority of animals with HPC or septal lesions
over controls in acquiring two-way active avoidance learning
(Isaacson, Douglas & Moore, 1961; King, 1958) is probably due in
part to the peculiarities of the shuttle-box situation which gives
an advantage to animals with a PA deficit. Grossman (1976)

120
noted that rats with FNX cuts or septal lesions were hyperactive and
avoided mainly by shuttling spontaneously between compartments,
expending a great deal of energy in the process. The fact that
animals with MFB cuts were not hyperactive and learned to respond
approriately to the CS suggested to Grossman that the altered
responding by HPC and septal animals in this paradigm was due to
"an interruption of reticulo-septo-hippocampal interconnections"
(1076, p. 395).
Grossman (1976) reported that sectioning the FNX, ST, or MFB
failed to mimic the deficit in acquiring the one-way active avoidance
task seen in animals with amygdalar, septal, and hippocampal damage.
As noted earlier, an animal should be able to encode and retrieve
one strategy in a particular situation (as long as no competing
responses were available) if the cortex has access to amygdala
generated emotional information during training. An alternate
pathway for this amygdala information is available via a branch from
the ST to the SM (see Fig. 2). The one-way deficit was reproduced
by sections :of the SM (which appear to be downstream from the ST
branch) performed by Ross in Grossman's laboratory (Grossman,
1976).
A Functional Meta-system
Human beings differ from lower forms in that a large part of
human brain development and organization occurs postnatally.
The special and evolved integrative functions which distinguish homo
sapiens are related to specialization of the secondary and specifically
human tertiary association areas which do not reach functional

121
maturity until relatively late in ontogeny. The progression in
species from dependence on primitive perceptions through secondary
and tertiary integration is also evident in the development of brain
function in human individuals ('ontogeny recapitulates philogeny1).
The primary projection areas are operational at birth; secondary
association areas are programmed for function (via "learning") after
birth (Penfield, 1975); the tertiary integration areas do not become
fully operative until the seventh year of life (Luria, 1973a).
(This developmental sequence appears to parallel Piaget's descriptions
of sensory-motor, concrete operational, and formal operational
stages.) During ontogeny, the lower systems in the hierarchies become
subordinate to the higher levels: elementary perceptions are fit
into learned schemes and, in adults, are coded into logical systems
(Luria, 1973a). Inadequate development at any one level will affect
functioning at higher levels.
The development in humans of manual dexterity and, later, of
speech required the specialization of certain secondary and tertiary
association areas. A tendency emerged for these operations to be
organized in one hemisphere of the brain (Luria's law of the
progressive lateralization of functions). At this point the functions
of the hemispheres began to differ radically.
Both hemispheres possess advanced circuitry with the capacity
for high level information processing. However, they are relatively
isolated from each other and the left is genetically prepared to
deal with language. The primary projection areas in the two hemi
spheres have identical roles, but the secondary and tertiary

122
association areas, responsible for an increasingly complex synthesis
of incoming information, becomes specialized to perform different
tasks in different ways. The mode of information processing in the
two hemispheres begins to diverge as speech comes to dominate the
functional organization of association areas on the left, leaving
the high level integration of nonverbal sensory input to their
counterparts on the right. In the course of ontogenetic development
the tertiary zones within each hemisphere begin to control the work
of the secondary areas, which become subordinate to them (Luria,
1973a, ch. 2). In this manner the basic perceptual processes are
altered to conform to the needs of the higher centers. The secondary
association areas on the left become specialized to perceive semantic
data which can be coded into logical systems. The left tertiary area
analyzes this input sequentially, abstracting relevant details and
associating these with verbal symbols (see Nebes, 1977). The secon
dary areas on the right specialize in analyzing concrete structural
data and relationships which are organized into complex wholes by
the tertiary association area on that side.
Evidence (to be detailed below) indicates that during develop
ment these two increasingly specialized cortical information
processing systems (each with its own subjacent memory, emotion,
and arousal subsystems) become organized into a single functional
meta-system in which the cortical components are yoked together by
the limbic system. This meta-system optimizes the utilization of
its complementary components in the service of producing adaptive
behavior and assuring the survival of the species. Within this

123
meta-system the right hemisphere performs the functions of an
environment "monitoring" system and the left hemisphere constitutes
a problem-solving and response-generating system; the amygdalae
and the left prefrontal lobe form an emotional "motivation" system;
and the reticular activating system; (with their brainstem, thalamic
and prefrontal lobe components) compose a cortical "mobilization"
system.
While the differences between the cognitive operations performed
by the left and right cerebral hemispheres may have been over
popularized little, if any, attention has been paid to the fact
that the older medio-basal and subcortical structures (which sub
serve the processes of emotion and attention) are also bilaterally
represented. It is common practice to refer to "the" limbic system
when in fact the components of this system are duplicated within
each half brain. It is reasonable to expect that these lateralized
structures are also specialized as to function, or reflect the
specialization of the neocortical mechanisms which they subserve.
In the context of the functional meta-system, motivation and arousal
must normally be activated by the right (monitoring) system and
responded to by the left (problem-solving and response-generating)
system. If this were not the case then the automatic nature of the
overall system (which is guaranteed by the relative isolation of
its components) might be defeated with the result that adaptive
behavior would not be assured in survival situations. A basic
assumption here is that mother nature would not relinquish instinct-
based responding and permit self-determined behavior without firm

124
guarantees that the species would continue to survive. (The
psychological correlates of emotion and arousal activated by the
left hemisphere will be discussed in the section on psychopathology,
below!)
The normal chain of events within the meta-system might be
traced as follows (see Fig. 4): neuronal signals, originating
in the sensory receptors, which reflect novel or potentially
significant environmental stimuli excite the brainstem raphe'
nuclei by way of sensory nerve collaterals. The raohe' nuclei activate
the reticular formation resulting in generalized cortical arousal
(orienting). (1) The right hippocampal system responds to environ
mental stimuli by causing the (cortical) memory traces which
represent the organism's previous experience with similar stimuli
to be activated; (2) if any of these memory traces contain coded
emotional attributes the right amygdaloid como!ex is activated by
them; (3) the amygdala, (a) acting via the septal area, focuses
the hippocampal memory search on the significant memory trace with
the secondary effect of releasing hippocampal inhibition of the
reticular formation and (b) influences the reticular formation
directly to release mobilizing energy and (c) relays its activation
to the left amygdala via the anterior commissure; (4) left amygdala
activation is then forwarded (via the dorsomedial thalamic nucleus
to the left orbitofrontal cortex where it achieves conscious
awareness as subjective emotional experience; (5) the dorsolateral
area of the left prefrontal lobe utilizes this motivational informa
tion to formulate intentions and will attempt to resolve the

125
emotional experience by programming cognitive operations in the left
post-central association areas to analyze the situation (aided by
context information and generalized expectations obtained from the
right experiential memory system in steps two and three) and formulate
a behavioral response; (6) the behavioral response alters the
situation, allowing the termination of motivating and mobilizing
systems activity.
Evidence concerning the proposed model will be organized and
presented below in terms of its relevance to the arousal, emotional,
and cognitive components of the meta-system.
Lateralized Mobilization Processes
The functional meta-system model suggests that the initial
identification or categorizaton of an environmental stimulus will
normally take place in the right hemisphere which will then trigger
the mobilization of the left half of the brain to deal with that
stimulus. Substantial support for this postulate is provided by
data from investigations utilizing different experimental approaches.
Asymmetrical reaction time to laterally presented stimuli.
Hielman and Van Den Able (1977) tested the effect of neutral warning
signals (WS) presented to the left or right visual half-field
on right hand reaction time (RT) to a centrally presented light.
They found that a WS presented to the right hemisphere reduced
RTs of the right hand more than a WS presented to the left hemisphere.
Bowers and Hielman (1976) examined between-hand RT differences
to a binaurally presented tone which was randomly preceded by a
verbal or nonverbal WS stimulus presented visually to both hemispheres

126
simultaneously. They found that a nonverbal WS reduced RTs of
both hands equally, no RT asymmetries occurred when the verbal WS
merely forewarned the subject (simple RT condition), right hand
RTs were faster than left following a verbal WS only when a response-
linked decision process was required (the go/no go condition in
which the WS both forewarned and dictated whether or not a response
should be made). The results of these two studies suggested to the
authors that
The right hemisphere may activate the left hemisphere
via interhemispheric pathways, wherease the extent to
which the left hemisphere can activate the right hemi
sphere is considerably less. Essentially, a 'one-way
street' is proposed in which the right hemisphere
activates the left more than the left hemisphere acti
vates the right (i.e., asymmetric interhemispheric
activation). (Bowers & Hielman, 1976, p. 7)
Altered GSR following unilateral brain injury. Hielman,
Schwartz and Watson (1978) studied arousal responses in patients
with left hemisphere damage (an aphasia syndrome), right hemisphere
damage (with the neglect/indifference syndrome), and no brain
damage, by electrically stimulating the forearm ipsilateral to
the brain injury and measuring GSR from the fingers on that side.
They found that the right hemisphere group showed singificantly
less GSR than either the left hemisphere or control groups (five
of the seven right hemisphere pateints had no measurable GSR at
all). The left hemisphere group had an exaggerated GSR relative
to the controls. These results were not attributable to differences
in sensory input or lesion size. The authors concluded that patients
with right hemisphere injuries and neglect have defective arousal.

127
They suggested that this hypoarousal might be a factor in the
indifference and other mental status anomalies that are common in
these patients.
Asymmetrical biochemical and electrophysiological processes.
Evidence reviewed in an earlier section delineated two separate
cortical mobilization systems which co-exist in each hemisphere.
A diffuse arousal system is based on the serotonergic (5-HT)
neurons of the median raphe nuclei located in the core of the
reticular formation. An attention focusing activation system
utilizes the catecholamines NE and DA which are derived from the
more laterally placed locus ceruleus and nuclei in the peri
aqueductal gray. The diffuse, indiscriminate arousal of cortical
neurons produced by the 5-HT system results in increased stimulus
reactivity and other phenomena which facilitate the primary objective
of orienting processes (i.e., stimulus sampling and identification).
Conversely, the focusing of attention produced by the NE-DA
activation system is essential to the processes of analysis and
response generation. The proposed model would suggest, therefore,
that 5-HT related mobilization processes would predominate in
the right hemisphere and NE/DA related processes would be more
evident in the left. Gottfries, Perris, and Roos (1974) found that
increased levels of the serotonin metabolite 5-hydroxyindoleacetic
(5-HIAA) in the cerebrospinal fluid was significantly and positively
correlated with the amplitude of auditory evoked potentials from
the right, but not the left hemisphere in a group of mixed
psychiatric patients. On the other hand, increased levels of the

128
NE/DA metabolite homovanillic acid (HVA) in the CSF was significantly
and positively correlated with the amplitude of evoked potentials
from the left, but not from the right hemisphere. These findings
indicate that an identical stimulus triggers mobilization processes
associated with orienting in the right hemisphere and with problem
solving/response-generating in the left.
Bilateral Interaction in Emotion and Cognition
The model outlined above specifies that the quality of signifi
cance is encoded along with context information in neuronal
models (memories) which are stored in the neocortex. When such a
model is activated by an environmental stimulus this coded informa
tion causes the amygdala to generate an emotional response which
will motivate the organism to respond appropriately. This cortical-
limbic system interaction was apparent in a unique experiment
performed by Doty (1973) in which a macaque monkey was prepared
with a unilateral amygdalectomy combined with sectioning of the
optic tract on the opposite side, thus blinding the hemisphere with
the intact amygdala. This was followed by sectioning of all the
forebrain commissures except for the posterior one-third (the splenium)
of the corpus callosum. The splenium was ensnared by a ligature
which was left protruding through the skull. After recovering from
surgery the animal was placed in a large enclosure. When a man
entered the enclosure the animal responded with normal fear and
fled. However, when the commissural transection was completed
(by pulling the snare) the animal's behavior was dramatically
different; it became docile and would actually approach and

129
nuzzle the man. Prior to the completion of the neocortical
disconnection the hemisphere that was blind but capable of feeling
fear received adequate information from the sighted side to activate
the intact amygdala and motivate the organism appropriately. However,
when this blind but fearful hemisphere was isolated the animal's
behavior was determined by the sighted but fearless side. (The
animal responded with normal fear and aggression when touched on
the trunk or limb.) The result was, in effect, a functional
bilateral amygdalectomy with regard to visual stimuli (Puccetti,
1977), demonstrating that without adequate cortical input to the
amygdala the organism cannot behave in an adaptive manner.
When a visual stimulus is presented tachistoscopically to
one hemisphere (by way of the left or right visual half-fields)
in a split-brain patient, the opposite hemisphere is "blind"
with respect to that stimulus. Case P.S. differs from earlier
split-brain patients in that his anterior commissure (which connects
the amygdalae and temporal lobes) was left intact. The combination
of these factors led to an important discovery in the course of
a standard experiment with P.S.:
On the verbal command test . where a word was
lateralized to the right hemisphere and P.S. was
instructed to perform the action described by the
word, his reaction to the word kiss proved revealing.
Although the left hemisphere of this adolescent boy
did not see the word, immediately after kiss was
exposed to the mute right hemisphere, the left
blurted out, "Hey, no way, no way. You've got to
be kidding." When asked what it was that he was
not going to do, he was unable to tell us. Later,
we presented kiss to the left hemisphere and a
similar response occurred: "No way. I'm not going
to kiss you guys." However this time the speaking

130
half-brain knew what the word was. In both instances,
the command kiss elicited an emotional reaction that
was detected by the verbal system of the left hemi
sphere, and the overt verbal response of the left
hemisphere was basically the same, regardless of
whether the word was presented to the right or left
half-brain. In other words, the verbal system of the
left hemisphere seemed able to accurately read the
emotional tone and direction of a word seen by the
right hemisphere alone. (Gazzaniga & Ledoux, 1977,
p. 151)
Thus, P.S.'s left hemisphere appears to have experienced a direction
ally specific emotion in the absence of a cognition, and the
required information must have been obtained from the right
hemisphere via the AC. In this case it is not possible to differen
tiate whether the response was made possible by information passed
between his amygdalae, or by cognitive information passed from the
right temporal lobe to the left, or both. The interpretation of
this phenomenon is further complicated by the fact that P.S.
possessed language in his right hemisphere. The finding demonstrates,
however, that the interaction of the hemispheres in emotion as
proposed in the present model is tenable. The likelihood of such
an interaction is supported by evidence concerning differences
in the emotional responsiveness of split-brain patients and adults
who have had their right hemispheres removed because of fast
growing tumors.
The most surprising initial observation concerning split-
brain subjects is still the most significant: there were no
readily observable changes in the intelligence, behavior, or
personalities of these patients following the disconnection of
their cerebral hemispheres (Sperry, 1968). The model being

131
considered here would suggest that this is because the normal inter
actions between the hemispheres which involve motivation and arousal
take place at a subcortical level. Sperry pointed out that the
right hemisphere of these patients demonstrated "appropriate emotional
reactions" but that "apparently, only the emotional effect gets
across [to the left], as if the cognitive comDonents of the
process cannot be articulate through the brainstem . the
affective component gets across to the speaking hemisphere, but
not the more specific information" (1968, p. 732).
The proposed model specifies further that the right hemisphere
in humans specializes in analyzing and recording experiential data
and thus becomes the repository of the reinforcement history of the
individual. The emotional responsiveness and behavior of adults
whose right hemispheres were removed (because they had been invaded
by life-threatening, fast growing tumors) is markedly different
from the split-brain patient. Although normal intelligence
is generally retained, such people show a lessened capacity for
adaptability (Glees, 1961) and "suffered a loss in terms of
personality values . defects in judgement and . impairment
in insight, in emotional control, in initiative, and in perseverance"
(Gardner, Karnosh, McClure & Gardner, 1955, pp. 500-501). These
individuals, whose left hemisphere was deprived not only of sensory
and cognitive information from the right (as was the case with
cerebral commissurotomy patients), but also of that hemisphere's
motivational and mobilizing inputs, show a clear pattern of deficits:
their affect tends to be labile, inappropriate, and poorly modulated;

132
they have difficulty sustaining mental activity; they become ego
centric, shallow, and seemingly unaware of the social consequences
of their behavior (Gardner, 1933; Rowe, 1937; Bell & Karnosh,
1949; Mensh, Schwartz, Matarazzo & Matarazzo, 1952; Gardner et al.,
1955; Austin & Grant, 1962; Bruell & Albee, 1962). It may be deduced
from the above that normal, adaptive, responsiveness to the environ
ment depends on the products of right hemisphere processes. The fact
that the emotional impact of a situation is available to the left
hemisphere in split-brain patients but not in right hemispherectomy
patients indicates that the right cerebral hemisphere in humans
is responsible for making determinations regarding the significance
of environmental stimuli. The contrast between the cerebral dis
connection and right hemispherectomy syndromes further suggests
that relatively normal emotional resDonsiveness, judgement, and
adaptability can be maintained as long as the motivating and mobilizing
products of the right hemisphere's cognitive operations can be
transmitted to the left (i.e., can activate the left amygdala and
ARAS). In the absence of the right hemisphere's modulating input
the left hemisphere is subject to erratic and inappropriate emotional
experience because the neocortex on that side is not adequately
prepared to mediate these processes.
Patients with lesions in their left cerebral hemispheres
typically have a catastrophic/depressive reaction while those with
right hemisphere damage show a characteristic indifferent/euphoric
response (e.g., Gionotti, 1972a). Transient, but similar emotional
reactions are commonly seen during recovery from unilateral

133
hemisphere anesthetization using the Wada technique (e.g., Pern's,
Rosadini & Rossi, 1961) and have been reDorted after unilateral
ECT (Deglin & Nikolaenko, 1975). Such findings have prompted
speculation that each hemisphere tends toward a different emotional
state.
The fundamental problem with the lateralized emotional valance
theory is the fact that the actual direction of the valance cannot
be specified. Tucker (1981) notes that the phenomena observed
following the unilateral loss of cortical function might result
from the disinhibition of the emotional tendency of the opposite
hemisphere or from the release of subcortical processes on the
same side.
Dimond, Farrington, and Johnson (1976) reported experimental
evidence suggesting that the right hemisphere has a negative
valance. They presented three short films to normal subjects'
left or right visual half-fields (using a special arrangement of
spectacles and contact lenses) and to free-viewing controls. The
right hemisphere group rated the films as significantly more
unpleasant and horrific than did the left hemisphere and control
groups (who did not differ from each other). Dimond et al.,
concluded that the right hemisphere tends toward a negative
emotional appraisal of incoming stimuli similar to the "characteristic
perception of the depressed patient" (p. 691). This finding would
seem to support the contralateral disinhibition interpretation of
the emotional phenomena seen after unilateral brain injury.
However, the contralateral disinhibition notion is discredited by

134
the fact that the emotional responses seen with unilateral hemi
sphere anesthetization occur only as the effects of the anesthetic
are clearing: they should be most pronounced when the contralateral
hemisphere was completedly incapacitated (see Tucker, 1981).
The indifference/euphoric reaction seen after right hemisphere
injury is often accompanied by neglect for the left half of the
body and for the left extrapersonal space (Gionotti, 1972b).
This neglect syndrome is usually associated with damage to the
right IPL, although it has been reported after lesions of the right
frontal lobe (Hielman and Valenstein, 1972) which presumably programs
the operations of the right post-central areas. Hielman and
Valenstein note that the mechanism of neglect remains unknown but
observe that
The inability of sensory stimuli to excite or alert an
organism (neglect) cannot be completely explained by
defect in sensory synthesis. The lesions that pro
duce neglect must also interfere with centrifugal
or descending pathways that normally would permit
integrated sensory stimuli to excite or alert the
organism. (1972, p. 663)
These authors suggest that neglect following parietal lobe lesions
may result from disconnection between sensory association areas and
the limbic system. What is most puzzling, and perhaps most
significant, is the fact that neglect is rarely, if ever, seen
following left hemisphere lesions.
The puzzles of lateralized emotional valance and unilateral
neglect resolve themselves if the basic premises of the proposed
functional meta-system hypothesis are accepted. This model
suggests that the primary role of the amygdala is to generate

135
emotional signals which motivate the organism to respond appropriately
in life or death situations (e.g., fight or flight). These signals
must be transmitted via the left amygdala to the left frontal
lobe before they can achieve conscious status. The subjective
experiences which accompany these "primitive" emotional states are
intrinsically negative, and must be in order to insure a prompt and
vigorous response. The task of evaluating the significance of
environmental stimuli ("monitoring") is relegated to the right
hemisphere in humans, the left being occupied with language functions
which require an incompatible mode of information processing. If
the right hemisphere is damaged and the right amygdala receives
little or no information, then no motivating emotional signal is
generated to be passed on to the left side. In the absence of
these signals the conscious left hemisphere is indifferent to the
stimuli it perceives. If the cortex is mobilized in the absence
of motivational inputs the resulting subjective state might be
characterized as euphoria. Conversely, if a damaged left hemisphere
is receiving emotional input from a normal (or disinhibited) right
hemisphere, and the damage diminishes the left side's ability
to resolve or cope with such input, the resulting experience might
be characterized as catastrophic. In the case of damage to the
right hemisphere's highest, and final, level of information
processing (the IPL), the left would not only fail to receive
emotional/alerting signals but would be deprived of processed
sensory data about the left half of the body and space as well,
from the left hemisphere's point of view these stimuli would cease
to exist.

136
Springer and Deutch (1981) suggested that the left hemisphere
"assumes that what it sees encompasses everything there is" (p. 176).
This interpretation is supported by the fact that the speaking left
hemisphere of split-brain subjects is unaware of having lost half of
its sensory fields (Gazzaniga, 1970). Similarly, the subjects
in the experiment reported by Dimond et al. (1976), who viewed
films presented to only one hemisphere, were not aware that they
were blind in one visual field. In addition, they experienced
the stimuli as "centered" when in reality it was presented from
the left or right of the midline. Finally, it is worth noting in
this context that Korsakoff's psychosis (in which the patient is
unable to gain access to recent memory traces) is commonly referred
to as the "amnestic-confabulatory syndrome." The Korsakoff patient's
speaking left hemisphere produces false accounts when memory fails
to provide the facts. This material "is presented without awareness
of its distortions or of its inappropriateness, and ... is moti
vated in no other way than factual information based on genuine
data" (Talland, 1965, p. 50). In short, when deprived of expected
memory data, the left hemisphere believes that what it thinks is
true. Considerable evidence of confabulation by the isolated left
hemisphere in split-brain patients was reported by Sperry (1974).
Mental Health and Psychopathology
The model developed in the preceding sections defines the basic
functional elements of the personality structure. It remains to
describe the ways in which these interact to produce mental health
and psychopathology.

137
According to the model normal, adaptive functioning requires that
the problem-solving/response-generating system receive adequate
information from the monitoring, mobilizing and motivating systems.
Mental health would require, and result from, the integrated
functioning of these basic personality elements. Conscious left
hemisphere processes must be mobilized appropriately by emotional
experience and "channelized" by generalized expectations which are
congruent with reality (cf., Kelly, 1963). Such integrated
functioning was implicit in Rogers (1963) description of the
"fully functioning person":
This person would be open to his experience . .
every stimulus, whether originating within the
organism or in the environment, would be freely
relayed through the nervous system without being
distorted by a defensive mechanism . the
self and personality would emerge from experience,
rather than experience being translated or twisted
to fit a preconceived self-structure . since he
would be open to his experience . this person
would find his organism a trustworthy means of
arriving at the most satisfying behavior in each
existential situation. (1963, pp. 18-20)
Any process that interfered with the operation of an individual
element, or prevented adequate interaction among them, would diminish
the adaptability of the organism. The system is vulnerable to
interference at several levels.
The model suggests that, at a cognitive level, generalized
expectations (GEs) are the most potent influence on behavior.
These significant experiential memories are assembled and stored
in the right half of the neocortex and are accessed by the hippocampal
system lateralized in that hemisphere. Such GEs are here defined as

138
experientially-based alterations in the disposition of the organism
relative to a stimulus object, situation, or event. The cognitive
information contained in the GE can only pass to the left hemisphere
over the neocortical commissures. This exchange permits the verbal
system to symbolize its reinforcement history in a given situation.
It is possible that the more facile mechanisms in the left hemisphere
might be conditioned to somehow prevent this exchange of information
to avoid the evocation of psychological pain. In this case, the
individual would experience the arousal and emotion appropriate
to the stimulus but remain ignorant of its nature, and consequently,
impotent in his or her efforts to resolve the subjective experience.
At least two separate mobilizing systems co-exist within each
half of the brain: a serotonergic system produces indiscriminant
cortical arousal which facilitates the functioning of the monitoring
system in the right hemisphere; a dopaminergic activation and attention
focusing system is essential to the cognitive operations of the left
hemisphere's problem-solving/response-generating system. Any
process which altered the balance between these two systems (within
or between hemispheres) would disrupt normal functioning. The
biochemical mechanisms which form the neural substrata of the
motivating and mobilizing systems are modulated by descending
influences from the prefrontal lobes (via the amygdalae) and from
the temporal lobes (via the hippocampi). The normal functioning of
these modulating pathways might thus be subverted by conditioning
processes (in the interest of "defending the ego") at the expense
of normal, integrated functioning. In the absence of integrated

139
functioning the system which has greater access to the motivating and
mobilizing system would have the greatest impact on cognitive and
behavioral responding. This conclusion is perhaps best exemplified
in the psychological sequelae of unilateral temporal lobe epilepsy.
The Psychopathological Correlates of Unilateral Temporal Lobe
Epilepsy
In all members of the phylum, biologically important experiences
elicit an emotional resonse which facilitates the learning of
biologically important behavior (Campbell, 1974). Destruction of the
amygdala, or disconnection of the amygdala from the sensory cortices,
results in the dissociation of affective qualities from sensory
stimuli (e.g., Kluver & Buey, 1938). Sensory-limbic association
depends on a neural pathway which extends to the amygdala via the
ventral temporal cortex (Bear, 1979).
Patients with temporal lobe epilepsy often show progressive
personality changes which culminate in patterns which are indis
tinguishable from certain psychodiagnostic entities. Bear (1979)
reviewed evidence suggesting that these changes are produced by
chronic over-stimulation of the amygdala. In contrast to the
consequences of temporo-1imbic disconnection, he postulated that
the alternations in behavior, emotion, and thought observed in this
disease stem from a process of temporo-1imbic hyperconnection:
a "progressive overinvestment of perception and thought with affective
significance" (p. 359).
Bear and Fedio (1977) compared the self-reported and observer
ratings of personality traits in patients with left and right temporal
lobe epileptic foci. Factor analysis revealed two factors which

140
differentiated between the two groups. On an emotive-ideative
dimension, patients with right temporal foci were distinguished by
externally demonstrated affect while the left group was characterized
by ideational/ruminative traits. On a normal-severe dimension the
left group of patients reported more severity compared to the right
and endorsed more socially disapproved traits (e.g., paranoia,
aggression, dependence) but the rater evaluated the right group
as more severely disturbed. Bear (1979) observed that both groups
displayed a characteristic high intensity of affect but differed
in verbal awareness of the emotions. The major difference between
the groups was in the way they interpreted and reported their abnormal
affective experience. Bear suggested that "strong affect expressed
as mood excess differs from cognitive elaboration, often verbal or
logical, of specific relationships between stimuli and affect"
(1979, p. 370). This difference might account for the fact that
the right hemisphere epileptic foci have been associated with mood-
affective disorders and left foci with cognitive-paranoid psychosis
(Flor-Henry, 1974). Bear and Fedio (1977) speculated about the
ways in which a patient might interpret the enhanced affective
associations to previously neutral events or concepts which result
from chronic temporo-limbic hyperconnection:
Experiencing objects and events shot through with affec
tive coloration engenders a mystically religious world
view if a patient's immediate actions and thoughts are so
cathected, the result is an augmented sense of personal
destiny. A felt significance behind events that others
dismiss constitutes a seed bed for paranoia or may
confirm the feeling that the patient is a passive
pawn in the hands of powerful forces that structure
the world. Feeling fervently about rules and laws

141
may lead to action in which the patient "takes the law
into his own hands." Sensing emotional importance in
even the smallest acts, he performs these ritualisti-
cally and repetitively. Since details bear the
imprimatur of affective significance, they may be
mentioned in lengthy, circumstantial speech or
writing, (o. 465)
Discussion: Hyper- and Hypo-dominance Spectrum Disorders
The proposed functional meta-system model specifies that, in
survival situations, emotion and arousal are normally triggered
by the right ("monitoring") system and are resolved by the left
("problem-solving") system. The right memory system also provides
appropriate context information and generalized expectations (via
the cerebral commissures) to assist in this process. When the
verbal left hemisphere initiates limbic system processes directly
(e.g., by triggering its own amygdala) there would be a tendency
toward a vicious circle: the increased attentional/emotional
activity would exacerbate the focus on the initiating cognitive
theme (rather than eliciting an objective response to environmental
requirements). In normal circumstances this reverberating unilateral
process might account for mood (the maintenance of affective tone
in the absence of external stimuli). If abnormally exacerbated,
the process might produce the rigidified cognitive phenomena
associated with various pathological syndromes (e.g., anxiety,
depression, paranoia, obsessions). In addition, the overactive
verbal system would systematically initiate searches in the contra
lateral memory system for experiential data which were congruent
with its abnormal cognitive state. Finally, memories encoded with
"artifically" induced emotional significance would, by definition,
tend to deviate from reality.

142
Cognitive activity in the right hemisphere is limited by
its inability to manipulate time. The right half of the brain
is context-bound, both internally and externally; its products are
reflexive, not considered. While the right hemisphere is responsible
for mediating the external expression of immediate affect the
appreciation of subjective emotional experience is a left-brain
function. Right brain activities are seen here as ancillary processes
which are utilized by the left. Contrary to recent theories
which tend to conceptualize psychopathology as emerging from the
disruption of a balance between right and left brain functions
(e.g., Bogan, 1969b; Flor-Henry, 1979; Tucker, 1981) the present
model emphasizes the need for integration of the influences which
converge on the left hemisphere. Any process which interfered with
the integrated functioning of the meta-system would be, by
definition, pathological. A process which culminated in the relative
overactivation of the left hemisphere would diminish the right
hemisphere's modulation of behavior and, conversely the right
hemisphere's contribution would be exaggerated when activation of the
left was reduced or disrunted. Such processes would be exacerbated
by the effects of reciprocal transcallosal inhibition (Flor-Henry,
1979). Since considered, "self-determined" responding is coordinated
by the left hemisphere such processes would lead to pathological
conditions of hyper- and hypo-dominance, respectively.
In the following paragraphs standard psychodiagnostic entities
will be related to neurological indices which reflect the operation
and interaction of elements within the proposed functional meta-system.

143
the evidence suggests that these entities may be divided into hyper-
and hypo-dominance spectrum disorders. Such a grouping should have
important theoretical implications for the treatment of these
disorders.
Schizophrenia and the Affective Disorders
The neurological correlates of schizophrenia and the affective
disorders have been reviewed by Tucker (1981) and Flor-Henry (1979)
The evidence suggests that these major disorders involve a severe
disruption of intrahemispheric functioning in addition to the
disturbances in interhemispheric information flow which is seen
here as contributing to neurotic and characterological illnesses.
The finding, noted earlier, that left temporal lobe epilepsy
often evolves to a schizophreniform pattern of symptomology is
paralleled by evidence of left hemisphere EEG abnormalities in
schizophrenia (see Tucker, 1981; Flor-Henry, 1979). The evidence
suggests that this abnormal cortical mobilization is mediated by
the dopaminergic activation system. Disruptive overactivation
of the left hemisphere, and right-ear deficiencies in temporal
discrimination tasks observed in schizophrenics were both normalized
following administration of chlorpromazine (Serafetinides, 1973;
Gruzelier & Hammond, 1976), a drug which blocks dopamine receptors.
Bruder and Yozawitz (1979) reported that patients with affec
tive disorders showed left-ear deficiency on dichotic listening tasks
which were correlated with the level of symptomology, indicating
right hemisphere dysfunction. This is consistent with findings
that unilateral right ECT and bilateral ECT were superior to left

144
ECT in relieving the symptoms of depression (haliday, Davidson &
Brown, 1968; Cronin, Bodly & Potts, 970).
While the dopaminergic activation system appears to predominate
the mobilization of the left hemisphere, the serotonergic arousal
system seems to be more imoortant in modulating the right hemisphere's
activity (see Gottfrieds, Perris & Roos, 1974). In their pharmaco
logical investigation, Mandell and Knapp (1979) found that lithium
treatment significantly reduced serotonin hemispheric asymmetry.
They hypothesized that the phenomenon of mood may be an emergent
property of asymmetrical serotonin regulation, and suggested that
varying degrees of serotonergic asymmetry accounted for the phase
being manic or depressive in bipolar affective disorders. (They
did not speculate on the direction of the asymmetry.)
Flor-Henry (1979) reported on a complex analysis of EEG data
taken from bipolar patients engaged in performing verbal and spatial
tasks (WAIS Vocabulary and Block Design subtests) and in neutral
conditions. He found abnormally high right parietal activity and
variability in depression which became bilateral in mania and
schizoaffective disorders. He also noted that, during spatial tasks,
depressives showed an increase in left temooral activity (versus
neutral conditions); they also showed an increase in right parietal
activity during verbal tasks. He suggested that these changes
indicated that comolex shifts of lateralized hemispheric specializa
tion were taking place. If this is the case, it is tempting to
speculate that such shifts might be caused by a reversal of the
normal inter- and intrahemispheric pattern of interaction between
the DA and 5-HT mobilization systems.

145
Anxiety Disorders, Obsessive-Compulsive Illness, and Paranoia
On the WAIS, a verbal I.Q. that is significantly higher than
performance I.Q. is especially characteristic of patients with
anxiety neurosis (see evidence summaried by Ogden, 1967). This
relationship indicates a relative predominance of left hemispheric
processes in this disorder. Tucker and his colleagues (1978)
reported two experiments which linked anxiety with left hemisphere
overactivation and dysfunction. Subjects reporting high anxiety
showed performance decrements on tasks lateralized (via the visual
half-fields) to the left, but not the right, hemisphere. High
trait anxiety was also associated with a right-ear attentional bias
and a low incidence of left lateral eye movements (Tucker, Antes,
Stenslie & Barnhardt, 1978).
Anxiety patients show lower than normal CNV amplitude. The CNV
develops slowly and irregularly in acute anxiety states, extinguishes
rapidly during deconditioning, and reappears slowly, if at all,
on reconditioning trials (Cohen, 1974). Since the CNV has been
related to the dopamine-mediated activation system, the elevated
levels of psychological activity and reduced cognitive efficiency
seen in anxiety states appears to be related to excessive serotonin-
mediated arousal processes in the left hemisphere. By contrast,
obsessive-compulsive subjects have an exaggerated CNV amplitude which
shows slow resolution, less decrement with partial reinforcement, and
fails to habituate (Cohen, 1974). Thus, the excessive (verbal)
cognitive activity which is characteristic in these individuals
seems to be related to overactivation of the dooamine-mediated

146
activation system. (The association between NE-mediated mobilization
processes involving the locus ceruleus and cingulate cortex and
the "hyper-indexing" of memories in obsessive-compulsive illness
was noted in an earlier section.)
Amnhetamine increases the amount of dopamine available at
synapses. In animals, increasing amounts of dopamine activation
results in a decrease in the ranges of behaviors emitted and an
increase in the frequency of a few behaviors, leading to stereo
typed motor sequences at high doses (Iverson, 1977). In humans,
chrnoic amphetamine abuse has been reported to result in stereotyped
repetitive behaviors, increased attention to detail, compulsive
disassembly of objects, ruminative preoccupation with intellectual
ideation, and features of hypervigilance difficult to distinguish
from paranoid schizophrenia (Ellinwood, 1967).
Witkin (1965) developed the concept of a field-dependence-inde
pendence dimension in perception. In the field-dependent mode, per
ception is dominated by the overall organization of the field, the
parts of which are experienced as fused. In field-independent
perceiving, parts of the field are experienced as discrete from
organized background (cf., perceptual augmenting and reducing).
Field-dependence and independence were dramatically related to
right and left hemisphere functioning, respectively, in an experiment
utilizing unilateral ECT as the independent variable and rod-and-
frame test scores as the dependent variables. Twenty-four subjects
were administered the rod-and-frame test shortly after admission to
the clinic and were randomly assigned to receive right or left ECT

147
(for the treatment of depression) within 48 hours. A second rod-
and-frame test was administered five hours after the treatment.
Twelve subjects whose first treatment was rescheduled for non-
clini cal reasons served as controls and were retested five hours
after the missed first appointment. The results were highly
significant; all twelve left ECT patients showed more field-
dependence on the second test* all twelve right ECT patients
showed less field dependence* the controls showed little or no
change (Cohen, Berent & Silverman, 1973).
Witkin (1965) relates the field-dependence-independence
perceptual dimension to a cognitive differentiation dimension:
the person who is field-dependent also does less well at "solving
problems which require isolating essential elements from the con
text in which they are presented and using them in different
contexts" (1965, p. 319). The cognitive differentiation dimension
is manifest in a global/diffuse versus an articulated cognitive style.
The field-dependent/global style has been associated with
hysterical neurosis (Zukmann, 1957), character disorders, somatiti-
zation, alcoholism, and patients whose primary symptom is affective
discharge (see the review by Witkin, 1965). A field independent/
articulated personality style has been associated with paranoia
(Janucci, 1964; Powell, 1964) and obsessive-compulsive disorders
(Zukmann, 1957).
Sociopathy and Hysteria
The absence of anxiety and the inability to learn from experience
are pathonomic in sociopathy. In contrast to anxiety neurotics,

148
"the most outstanding single feature of the sociopath's WAIS test
profile is his systematic high scores on the performance as compared
to the verbal part of the scale" (Wechsler, 1958, p. 176). The
same pattern was found to be characteristic of hysterics (Schafer,
1948). These relationships indicate a relative predominance of
right hemisphere processes in these disorders. The correlation
between low levels of left hemisphere functioning and low anxiety
is also striking. The PIQ greater than VIQ WAIS pattern has been
associated with "acting out" tendencies (Ogden, 1967). Silverman,
Buchsbaum, and Stierlin (1973) found that acting out adolescents
showed evoked potential augmenting. Augmenting has also been
associated with high scores on a sensation seeking scale (Zuckerman,
1974) and with alcoholism (Knorring, 1979; Coger, 1976), traits
which are common in sociopathy and hysteria (see Ball's, 1978).
Smokier and Shevrin (1979) used lateral eye movements as an
index of relative hemispheric activation in obsessive-compulsive
and hysterical personality style subjects (designated by modified
standard tests)] They found significantly more left-looking among
the hysterical.subjects indicating increased right hemisphere acti
vation in this disorder. They suggested that these personality
styles may be related to "predominant use of one or the other
hemisphere" (p. 952).
The dissociative reactions seen in hysterical neurosis (amnesia,
depersonalization, fugue, multiple personality) are indicative of a
failure to encode or access memories in the verbal system (cf.,
Gazzaniga, 1977). The finding that hysterical conversion symptoms

149
tend to be lateralized to the left half of the body (Galin,
Diamond & Braff", 1977) suggests a right hemisphere involvement
in these phenomena. The hysteric's characteristic indifference to
these manifestations is similar to that seen with right cortical
lesions, which has been interoreted here as the failue of the right
cortex to trigger subcortical motivation and mobilization processes
In sociopathic patients the CNV is absent or develos only
very low voltage (Cohen, 1974). The tendency for sociopaths and
hysterics to be field-dependent with a global/diffuse cognitive
style was noted earlier. This is consistent with the poorly articu
lated self-conceDts and value systems generally seen in these
personality types. The egocentricity and impulsiveness which
characterize these people tynifies the reflexive, context-bound
functioning of the right hemisphere. All of these trends indicate
that the left hemisphere is less involved in the organization of
behavior in these individuals.
Neurotic symptoms were found to be common following right
temporal lobectomy for epilepsy, while psychopathic disorders were
more frequent following this operation on the left side (Taylor,
1972). These findings provide significant support for the present
functional meta-system model which postulates that normal emotional
responsiveness depends on a motivation system pathway which extends
from the right amygdala (which is activated by the monitoring
system) to the left amygdala and on to the left prefrontal lobe

150
where the motivational signals are experienced as subjective emotion.
When this pathway is disrupted (by excision of the left temporal
components) emotional responsiveness is lost. When the right
temporal components are removed, motivational signals are generated
only on the left side and neurotic symptoms result. Failure of the
right hemisphere's monitoring system to activate the right limbic
system results in the failure to mobilize the left hemisphere
(so right processes have increased influence on behavior) and lack of
motivation signals to guide left hemisphere processes.

CHAPTER III
METHOD
This study was designed to affirm or disaffirm the consequents of
the proposed theoretical model of personality function and psychopathol
ogy and to demonstrate a confluence of psychological and neurological
observations. Specifically, the investigation attempted to ascertain
whether groups of adult psychiatric patients classified according to
the constructs of the proposed model as suffering from hypo- or hyper
dominance spectrum disorders differed significantly from each other on
a ratio of test scores that have been demonstrated to be sensitive to
damage to the left and right cerebral hemispheres, respectively,
without differing significantly in overall performance on the
instruments.
Both spectrum disorders are personality disturbances defined by
chronically diminished social performance and/or dysfunctional behavior
patterns without a clear precipitating factor. Both exclude gross
impairments of perception, orientation or memory. Hyper-dominance
spectrum disorders are defined as disorders in which the primary
symptoms are manifested intrapsychically and/or are characterized by a
central tendency to maladaptive overutilization of formal thought
processes. Hypo-dominance spectrum disorders are defined as disorders
in which the primary symptoms are manifested extrapsychically and by
a central tendency to maladaptive underutilization of formal thought
processes. DSM III diagnostic categories which meet these criteria
are listed in Appendix A.
151

152
Subjects
A total of 42 adult psychiatric patients served as subjects in
this study. Twenty outpatients were taken from an urban community
mental health center and three rural satellite clinics. Twenty-two
inpatients were taken from a state forensic hospital. Subjects were
assigned to the hyper- or hypo-dominant groups by DSM III Axis I and
II diagnoses (see Appendix A).
The hypo-dominant group (N=22) consisted of 13 males and nine
females. Eighteen were right-handed, one left-handed and three
ambidextrous by self-report. The mean age was 30.45 years with a
standard deviation of 12.14 years. The diagnoses within this
group included 14 antisocial personality disorder (eleven of whom
were classified as mentally disordered sex offenders undergoing
inpatient treatment at the state forensic hospital); three histrionic
personality disorder; two dependent personality disorder; one
avoidant personality disorder; one narcissistic personality disorder;
one psychogenic Dain disorder.
The hyper-dominant groun (N=20) consisted of 14 males and six
females. Ten were right-handed, one left-handed and three ambidextrous
by self-report. The mean age was 42.65 years with a standard
deviation of 13.73 years. The diagnoses within this grouo included
11 schizophrenia, paranoid types (all of whom were adjudicated
incompetent to stand trial and undergoing inpatient treatment at
the state forensic hospital); two generalized anxiety disorder;
two compulsive personality disorder; three dysthymic disorder;
one simple phobia; one agoraphobia with panic attacks.

153
All subjects were selected by diagnosis and were asked to
participate by their primary therapist. Each subject read an
informed consent statement (Appendix B) before being tested.
Instruments
The dependent measures in this study were scores on the Street
(1931) Gestalt Completion Test; the WAIS-R Object Assembly, Similari
ties and Information Subtests (Wechsler, 1981); and the Mini-Mult, a
71 item abbreviated form of the MMPI (Kincannon, 1968). The Street
Test and the Object Assembly Subtest have both been shown to be
differentially sensitive to right hemisphere cognitive processes and
right hemisphere damage impairs performance on these tests (Bogan,
Dezure, Ten Houten & March, 1972; Direnzi & Sninnler, 1966; Ogden,
1967; Matarazzo, 1972; Rapaport, 1951; Black, 1974). The Similarities
and Information Subtests of the WAIS-R have been shown to be differ
entially sensitive to the left hemisphere cognitive processes and
left hemisphere damage impairs performance on these tests (Bogan
et al., 1972; Ogden, 1967; Matarazzo, 1972; Rapaport, 1951). Bogan
et al. (1972) demonstrated that patients with partial sectioning
to the cerebral commissure show higher scores on the Street Test
and higher Street/Similarity ratios than patients with complete
sectioning of the commissures. This was interpreted by the authors
as indicating that the Street Test normally required right hemi
sphere processing.
Scales 3 and 4 from the MMPI were selected as representing
symptoms that are characteristic of the hypothesized hypo-dominance
spectrum disorder. High scorers on Scale 3 (hysteria) show a

154
"general denial of physical health . denial of psychological or
emotional problems and of discomfort in social situations. . .
They react to stress by developing physical symptoms . [and]
are not likely to report anxiety, tension or depression" (Graham,
1977, pp. 38-39). High scorers on Scale 4 (psychopathic deviate)
show "absence of satisfaction with life, family problems, delinquency,
sexual problems, and difficulties with authorities" (Graham, 1977,
p. 41). Scales 5 and 7 were selected as representing symptoms that
are characteristic of hypothesized hyDer-dominance spectrum
disorders. The high scorers on Scale 6 (paranoia) show paranoid
symptoms. Scale 7 (psychasthenia) is a good index of psychological
turmoil, including excessive doubts, anxiety, compulsions and
obsessions, unreasonable fears, and tension (Graham, 1977). Graham
concluded that the Mini-Mult is "useful for comoaring groups, particu
larly if they are psychiatric patients ..." (1977, p. 219).
Procedure
Demographic information and diagnostic data were taken from the
subjects' clinical records. No identifying information was recorded.
The Mini-Mult was administered by the subjects' primary therapist.
Subjects were tested by the author before or after their regularly
scheduled therapy session. Street Test, Similarities, Object Assembly,
and Information Subtests were administered in that order. Diagnos
tic data, demographic information and test responses were entered on
a subject data form. The WAIS-R subtests and the Mini-Mult were
scored by the author using standard scoring criteria (Wechsler, 1981;
Kincannon, 1968). The Street Test as originally standardized
(Street, 1931) includes a number of antiquated items and the

155
scoring criteria for two items were altered for this study. (The
complete socring criteria used are presented in Appendix C.)
Hypotheses
The following hypotheses were generated:
1. The ratio of the sum of the raw scores on the Street Gestalt
Completion Test and the WAIS-R Object Assembly Subtest to the
QAIS-R Similarities and Information Subtests (Street + Object
Assembly/Similarities + Information ) will differ between groups
with the hyper-dominant grouD showing relatively better performance
on the Similarities and Information Tests and the hypo-dominant
group showing relatively better performance on the Street and Object
Assembly Tests.
2. The sum of the standardized scores (z-scores) on all the
tests (Street + Object Assembly + Similarities + Information) will
not differ between the groups.
3. The ratio of the sum of the MMPI T-scores for scales
three and four to the sum of the T-scores for scales six and
seven (3T + 4T/6T + 7T) will differ significantly between groups
with the hyper-dominant group showing relatively higher scores on
scales six and seven and the hypo-dominant groups showing relatively
higher scores on scales three and four.

CHAPTER IV
RESULTS
Left versus Right Hemipshere Cognitive Functioning
Between Groups
Hypothesis 1 stated that the hyper- and hypo-dominant spectrum
disorders would differ on the ratio of scores on tests that have
been demonstrated to be sensitive to left and right hemisphere
cognitive functioning. Specifically, it was hyoothesized that the
hyper-dominant group would show relatively better performance on
the Similarities and Information WAIS-R Subtests and the hypo-
dominant group would show relatively better performance on the
Street Test and the WAIS-R Object Assembly Subtest. The mean
ratio (Street + Object Assembly/Si Hilarities + Information) for the
hyper-dominant group was x = 1.15 (SD = 0.42) while the mean ratio
for the hypo-dominant group was x = 1.91 (SD = 0.84), indicating
that the groups differed in the predicted directions. A student's
t-test for independent samples (Robson, 1973) performed on these means
revealed that the differences between groups was significant
(t = 3.4640, p < 0.001, one-tailed). It was concluded that
psychiatric inpatients and outpatients assigned to the hyper
dominant spectrum disorder group performed better on tests sensitive
to left hemisphere cognitive functioning relative to tests
sensitive to right cognitive functioning; their ratio scores
156

157
differed significantly (p < 0.001) from patients assigned to the
hypo-dominant group, who scored relatively better on tests sensitive
sensitive to right, relative to left, ehmisphere cognitive functioning,
as predicted by the proposed model.
Overall Performance on the Tests Sensitive to
Right versus Left Hemisphere Cognitive Functioning
Hypothesis 2 suggested that any differences discovered in the
test of Hypothesis 1 would be due only to differences in the ratios
between groups and not to any other pattern of overall performance
on the tests (e.g., intelligence). To test this hypothesis,
standardized scores (z-scores based on the total n) were derived for
each subject's individual test scores (following Robson, 1973) and
added togehter for each group (Street + Object Assembly +
Similarities + Information). The mean z-score was x = 0.09 for
the hypo-dominant group and x = -0.04 for the hyper-dominant group.
A student's t-test for independent samples (Robson, 1973) performing
on these means revealed no significant differences between groups
(t = 0.87^gg, p<0.4, two-tailed, N.S.). It was concluded that
there were no significant differences on overall test performance
between groups and that the differences in ratios found in the
test of Hypothesis 1 were not due to any factor (e.g., intelligence)
other than the one being tested.
Differences in MMPI Scores
Hypothesis 3 amounted to a verification of the subjects' diagnoses
and provided an additional test of the ability of the proposed
model to predict membership in diagnostic categories based on an
assessment of personality traits. Specifically, it was hypothesized

158
on an assessment of personality traits. Specifically, it was
hypothesized that the hyper-dominant group would score relatively
higher on MMPI clinical scales (6 and 7) which indicate symptoms
that are manifested intrapsychically and suggest a central tendency
to chronic and maladaptive overutilization of formal thought processes
while the hypo-dominant group would score relatively higher on MMPI
clinical scales (3 and 4) which indicate symptoms that are magnified
extrapsychically and suggest a chronic and maladaptive underutiliza
tion of formal thought processes. The mean ratio (3T + 4T/6T = 7T)
for the hyper-dominant group was x = 1.01 (SO = 0.12), while the
mean ratio for the hypo-dominant group was x = 1.12 (SD = 0.14),
indicating that the groups differed in the predicted directions.
A student's t-test for independent samples (Robson, 1973) performed
on these means revealed that the difference between groups was
highly significant (t = 2.75^ p < 0.005, one-tailed). It was
concluded that psychiatric inpatients and outpatients assigned to
the hyper-dominant spectrum disorder group scored relatively higher
on MMPI clinical scales which indicate symptoms that are manifested
intrapsychically and suggest a chronic and maladaptive overutiliza
tion of formal thought processes; their ratios differed significantly
from those of patients assigned to the hypo-dominant group, who
scored relatively higher on MMPI clinical scales which indicate
symptoms that are manifested extrapsychically and suggest a chronic
underutilization of formal thought processes, as predicted by the
proposed model.

159
Age
The mean age in the hyper-dominant group was x = 42.65 years
(SD = 13.73 years). The mean age in the hypo-dominant group was
x = 30.45 years (SD = 12.14 years). A student's t-test performed
on these means revealed significant differences between groups
(t = 2.12^g, p < 0.05, two-tailed).

CHAPTER V
DISCUSSION
This study was an attempt to affirm or disaffirm certain central
consequents of the proposed model of personality function and psycho
pathology. With this type of design a failure to support the hypotheses
is unequivocal: the theory cannot be true. If the consequences
are clearly verified there is some indication that the theory may be
true. All of the hypotheses in this experiment were suDported at
high levels of statistical significance. It was concluded that the
proposed theoretical model of personality function and psychopathology
is tenable and may provide useful operational definitions for the
applied psychologist.
The value of an experimental test of a theory is determined,
in large part, by the strength of the chain of reasoning between the
hypotheses and the data. In this study that connection is relatively
straightforward. Based on the functional meta-system model it was
postulated that the manifestations of personaltiy dysfunction (i.e.,
psychopathology) will be determined by the properties of the
cerebral hemisphere with greater access to the limbic system. It was
proposed, therefore, that there are two major subtypes of psycho
pathology. Since the functional impact of disturbances within the
meta-system is thought to be experienced in the left cerebral hemi
sphere (which alone possesses the requisites for self-awareness
160

161
and considered response generation), it is convenient to express
these two forms of pathology in terms of hyper- or hyDo-dominance.
Any given pathological syndrome will reflect both the point of
A
disturbance within the meta-system and the attempts of the rest of
the system to compensate, but most may be classified as hyper-
or hypo-dominance spectrum disorders with the primary symptoms
manifesting themselves either intra- or extrapsychically, resoectively.
A major consequence of hyper- or hypo-dominance is the avail
ability of right hemisphere cognitive products to the left for
utilization in response generation. The validity of the instruments
used as dependent measures in this study is well established and
their sensitivity to lateralized cognitive processes has been
adequately documented. The relationship between these indices of
lateralized brain functions and personality traits (as measured by
the relevant clinical scales of the MMPI) was clearly demonstrated
and congruent with the predictions of the theory. The agreement
between the MMPI data and the diagnoses used to assign subjects to
the experimental groups also enhances the value of the results.
A number of other factors contribute to the validity and general -
izability of the results in this study. Considerable design risks
were undertaken in that variables were introduced which would have
invalidated the results had they not conformed to predictions.
The assignment of mentally disordered sex offenders exclusively to
the hypo-dominant group and general psychiatric forensic inpatients
to only the hyper-dominant groups could not have been justified if
the two experimental groups had differed in overall performance on the

162
cognitive function measures. Given the outcome, however, the
inclusion of these two inpatient populations greatly enhances general-
izability. The absence of overall between-group differences on these
dependent measures also effectively rules out extraneous factors
(e.g., I.Q.) in the production of the results and dramatically
emphasizes the importance of the predicted left-right ratio variable.
Since the hyper-dominant group was significantly older (p < .05)
than the hypo-dominant group, it could be argued that age accounted
for the observed differences between groups. It is well established,
however, that hypo-dominant type symptoms tend to diminish with aging
(e. g., See Balis, ch. 4) and the observed difference is believed to
reflect the normal distribution of symptoms in the population.
An important limitation in this study is the fact that only cognitive
factors and personality traits were observed. One could reject the
functional-meta-system theory and interpret the results in terms of
Bogan's (1969c) "hemisphericity"/"dual mind" notion. In this case
the demonstrated correlation between lateralized cognitive functions
and personality traits would be seen as manifestations of "mind
left" versus "mind right." This alternative is discredited, however,
by the documented differences between split-brain patients and those
right hemispherectomy patients for whom relevant data are available.
Subjects deprived of their right hemisphere should tend toward
"hyper-dominance" type symptoms in this case, but the opposite is
true: as noted earlier, these patients show all the symptoms of
hypo-dominance, including egocentricity, affective lability, and
apparent lack of concern about the social consequences of their

163
behavior. The functional meta-system model accounts for this,
and the fact that split-brain subjects do not tend to develop
pathological symptoms, by hypothesizing that the right hemisphere
normally monitors the environment for significant (e.g., social)
stimuli, relates them to personal experience, and signals the left
hemisphere appropriately via the (subcortical) limbic system.
Perhaps the most important asset of the proposed model is the
fact that a rigorous and comprehensive test of the full range of
its predictions can be accomplished with existing technology. The
essential cortical-limbic system hyoerconnection postulate could
be evaluated by observing galvanic skin response (GSR) on the left
side in the hyper-dominant group. Augmentation of cortical averaged
evoked potentials (AEP) to increasing stimulus intensity would be
predicted in the hypo-dominant group and reduction in the other,
thus verifying the predicted cognitive mode. Differences in the
contingent negative variation (CNV) between obsessives, anxiety
neurotics, hysterics, and psychopaths were noted earlier and are
congruent with the tenets of the model.
An ideal test of the theory would include replication of the
cognitive and personality results of the present study, peripheral
physiological observations of indices of limbic system function (bi
lateral GSR), perceptual reactance (augmenting/reducing), and
information processing efficiency (CNV). This could be done with
cell sizes large enough to permit meaningful comparisons of
diagnostic entities to be made within and between grouDS. Such a
study might result in complete operational definitions of

164
psychopathological phenomena with important implications for the
practice of psychotherpay. Such a fully validated model would
permit an integrated approach to diagnosis and treatment planning.
An analysis of the ordering of events in a pathological psychological
process might allow specification of the relationships betv/een
significant experiences (recorded in the right hemisphere system),
the conditions and manner in which these (and associated generalized
expectations) are elicited by stimuli, and the way in which those
are interpreted and/or dealt with by the left hemisphere system. This
information would allow the goals of the treatment process to be
operationalized and indicate intervention points and methods. Such
a plan might specify, for example, that a specific corrective emotional
experience or metaphorical association of specified generalized
expectations to an identified stimulus, followed by a well-defined
cognitive behavior modication and a particular social skill enhance
ment would be the most efficient intervention package. Finally,
biofeedback techniques might find new significance in both the
processes of diagnosis and treatment.

APPENDIX A
HYPO- AND HYPER-DOMINANCE SPECTRUM DISORDERS
BY DSM III DIAGNOSTIC CLASSIFICATION
Hyper-dominance Spectrum
Disorders
295.30 Schizophrenia, paranoid
type
300.40 Dysthymic disorder
300.21 Agoraphobia with panic
attacks
300.22 Agoraphobia without
panic attacks
300.23 Social phobia
300.49 Simple phobia
300.01 Panic disorder
300.30 Obsessive-compulsive
disorder
300.02 Generalized anxiety
disorder
300.00 Atypical anxiety
disorder
301.00 Paranoid personality
disorder
301.40 Compulsive personality
disorder
Hypo-dominance Spectrum
Disorders
300.81 Somatization disorder
300.11 Conversion disorder
307.80 Psychogenic pain disorder
300.70 Hypochondriasis
300.71 Atypical somatoform
disorder
300.12 Psychogenic amnesia
300.13 Psychogenic fugure
300.14 Multiple personality
300.60 Depersonalization
disorder
300.15 Atypical dissociative
disorder
312.31 Pathological gambling
312.32 Kleptomania
312.33 Pyromania
312.34 Intermittent explosive
disorder
312.35 Isolated explosive
disorder
312.39 Atypical impulse control
disorder
301.50 Histrionic personality
disorder
301.70 Antisocial personality
disorder
301.60 Dependent personality
disorder
301.81 Narcissistic personality
disorder
165

APPENDIX B
INFORMED CONSENT STATEMENT
Project Title: Functional Brain Systems and Personality Dynamics
You are being asked to participate in a research project
which was designed to find out whether certain patterns of per
ceiving and processing information are related to specific emo
tional or nervous problems. You were selected because the types
of problems you have reported are similar to the ones being
investigated. If successful, the study could lead to more effec
tive ways of helping people with those problems.
Participation in this research project will take about 15 minutes
of your time. No monetary compensation will be awarded. No identi
fying information will be recorded so it will be impossible to
report your results to anyone and you will not be recontacted.
There are no risks or discomforts involved. You will be given a
true-false questionnaire by your therapist or case manager; all
the researcher will ask you to do is to identify different types
of visual patterns and answer a series of nonpersonal questions.
You may withdraw your consent and discontinue at any time without
prejudice. If you have further questions, you may contact
David Lindquist at Mental Health Services, Inc., 374-5690.
166

APPENDIX C
REVISED SCORING CRITERIA FOR THE STREET (1931) GESTALT COMPLETION TEST.
Credit any response that indicates that the following has been seen:
Item 1: a dog
Item 2: a boat or ship
Item 3: a cat
Item 4: a stove
Item 5: a baby
Item 6: a table
Item 7: a man in uniform with a rifle or a fisherman
with a fishing pole*
Item 8: a horse
Item 9: a rabbit
Item 10: a locomotive
Item 11: a boy on a tricycle
Item 12: a man's face
Item 13: a man kneeling*
*Altered from original in Street (1931)
167

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BIOGRAPHICAL SKETCH
David Lindquist was born on November 22, 1944, at Mitchell Air
Force Base on Long Island, New York. He grew up in a half dozen
cities around the country during the forties and fifties and
attended as many educational institutions during that period.
During the sixties he dropped out of high school, served in the
Air Force, was married and eventually divorced, obtained a high school
equivalent certificate, and worked at a variety of jobs. He took
courses at community colleges on a part-time basis beginning in 1968,
and earned an A.A. degree in 1973.
He came to the University of Florida in 1974, majored in
psychology, and received a B.A. degree with high honors in 1976. He
worked in several inpatient and outpatient settings while continuing
his graduate study toward the doctoral degree in counseling psychology
and has developed a specialization in forensic psychology.
188

I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is
fully adequate, in scope and quality, as a dissertation for the degree
Doctor of Philosophy.
irry Grater/ Chairman
Professor of Psychology
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is
fully adequate, in scope and quality, as a dissertation for the degree
Doctor of Philosophy.
James I. Morgan
Associate Professor of
Counselor Education
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is
fully adequate, in scope and quality, as a dissertation for the degree
Doctor of Philosophy.
Paul Schauble
Professor of Psychology
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is
fully adequate, in scope and quality, as a dissertation for the degree
Doctor of Philosophy.
David Suchman
Professor of Psychology
I certify that I have read this study and that in tpy opinion it
conforms to acceptable standards of scholarly presentation and is
fully adequate, in scope and quality, as a dissertation for the degree
Doctor of Philosophy.
Robert- Ziller
Professor of Psychology
- N

This dissertation was submitted to the Graduate Faculty of the
Department of Psychology in the College of Liberal Arts and Sciences
and to the Graduate School, and was accepted as partial fulfillment
of the requirements for the degree Doctor of Philosophy.
May 1935
Dean for Graduate Studies and
Research



41
a lateral-frontal-anrygdala lateral-hypothalamus fcilitory circuit
and an inhibitory orbitofrontal--amygdalamedial-hypothalamus
circuit. "Activation"is an attention focusing process involved in
perceptual expectancies and motor readiness to respond. This system
is based on the locus ceruleus, in the periaqueductal gray, which
supplies norepinephrine to the forebrain. Activation was thought
to be modulated by the ancient motor control system in the basal
ganglia. Together, these systems provide for appropriate attending
to novel or significant stimuli and prepare the organism to respond
cognitively and behaviorally. The hippocampus was seen to integrate
the functioning of these systems and to exert ultimate control over
cortical mobilization through a mutually inhibitory relationship with
the reticular formation.
Phasic Control Systems: The Frontal Lobes and Thalamus
A novel (possibly significant) stimulus elicits an orienting
response (OR) from the organism. The psychological phenomena
associated with the OR are familiar to all who have had experience
with "things that go bump in the night." The complete orienting
reaction includes
The suppression of ongoing behavior, the orienting of
the body and receptor towards the new stimulus, changes
in the peripheral autonomic nervous system, and, perhaps
less obvious, preparations for associating the new stimulus
with memories from the past and expectancies of the
future. (Issacson, 1974, p. 110)
Pribram (1973) noted that the stimulus sampling aspects of the
orienting reSDonse differed from the processes necessary to register
a stimulus in awareness and memory (which must be accomplished


91
the septal area (which receives the bulk of these terminals), the
anterior-ventral thalamus, the cingulate gyrus and the neocortex.
Stein and Wise (1971) suggest that this dorsal NE system is involved
in regulating cognitive activities.
The ascending serotonin (5-HT) system originates in the median
raphe' nucleus which is situated in the core of the reticular
formation and receives collateral branches from the sensory nerves.
5-HT cells in the raph^; have terminations throughout the central
gray of the midbrain. Axons from the raphe' also rise in the MFB and
distribute to the same areas as the dorsal NE system, with the
addition of terminals in the basal ganglia and more widespread
distribution in the neocortex. This system seems to be most
directly involved in regulating tonic arousal.
Brodie and Shore (1957) suggested that NE and 5-HT exert
opposing effects that modulate a variety of CNS functions (e.g.,
sleep, appetite, sexual drive). It appears that these parallel
systems control behavior by reciprocal action in balanced systems.
While functional specificity is determined by the neural circuitry
involved at a given anatomical level, the cumulative effects of
specific functional outcomes produced a preponderance of one or
the other transmitter seem to be consistent and to result in
coordination across levels. It appears that a preponderance of
the indolamine 5-HT, for example, results in the release of tonic
mobilizing energy at the brainstem level, the suppression of
ongoing behavior at the hypothalamic level, the identification of
"negative reinforcement" at the limbic system level, and


162
cognitive function measures. Given the outcome, however, the
inclusion of these two inpatient populations greatly enhances general-
izability. The absence of overall between-group differences on these
dependent measures also effectively rules out extraneous factors
(e.g., I.Q.) in the production of the results and dramatically
emphasizes the importance of the predicted left-right ratio variable.
Since the hyper-dominant group was significantly older (p < .05)
than the hypo-dominant group, it could be argued that age accounted
for the observed differences between groups. It is well established,
however, that hypo-dominant type symptoms tend to diminish with aging
(e. g., See Balis, ch. 4) and the observed difference is believed to
reflect the normal distribution of symptoms in the population.
An important limitation in this study is the fact that only cognitive
factors and personality traits were observed. One could reject the
functional-meta-system theory and interpret the results in terms of
Bogan's (1969c) "hemisphericity"/"dual mind" notion. In this case
the demonstrated correlation between lateralized cognitive functions
and personality traits would be seen as manifestations of "mind
left" versus "mind right." This alternative is discredited, however,
by the documented differences between split-brain patients and those
right hemispherectomy patients for whom relevant data are available.
Subjects deprived of their right hemisphere should tend toward
"hyper-dominance" type symptoms in this case, but the opposite is
true: as noted earlier, these patients show all the symptoms of
hypo-dominance, including egocentricity, affective lability, and
apparent lack of concern about the social consequences of their


108
The characteristics of the contingent negative variation
(CNV) were reviewed by Cohen (1974). Briefly, the CNV is a special
form of cortical evoked response consisting of a gradually developing
wave of negative potential which originates in the frontal lobes and
sweeps posteriorally over the cortex whenever there is a stimulus-
response contingency or expectancy built into a situation. This
wave becomes abruptly positive when the required perceptual or
motor response is executed. The amplitude of the signal is directly
related to the reinforcement ratio, with peak amplitude at 100%
reinforcement. Reaction time is shortest following high amplitude
CNVs. Perhaps the most interesting characteristic of the CNV is the
fact that it responds to verbal instructions: the waves disappear
when the instruction which produced the expectancy is countermanded,
and fail to appear when the subject is told in advance that the
contingent signal will not be forthcoming. The association between
perceptual reduction processes, the CNV, and dopamine is strengthened
by the finding that evoked potential reducers have increased levels
of homovanillic acid (a dopamine metabolite) in the cerebral spinal
fluid (Gottfries et al., 1974).
The Functional Elements of the Personality Structure
It has been assumed here that the phylogenetic transition from
instinct-based responding to self-determined behavior required the
evolution of new mechanisms to insure the survival of the individual
and the species. Specifically, it was hypothesized that automatic
neural systems were needed to (1) Monitor the environment for
significant stimuli (b) Motivate the organism to respond in the
presence of such stimuli-, and (c) Mobi 1 ize the appropriate


71
input and end with afferents which are capable of modulating the
activity of the brainstem RAS.
A determination of novelty or significance might be made at
the level of the secondary sensory association cortices where raw
receptor impulses are converted into functional (i.e., "meaningful")
information, although such duplication of effort in all the modalities
would be cumbersome and inefficient. Further, the relative signifi
cance of a stimulus may depend on the internal state of the organism
(e.g., satiation, the presence or absence of certain hormones, etc.).
The fact that information related to this added consideration is
most readily available in subcortical structures is another argument
favoring a centralized location for the mechanisms involved with the
decision to orient or habituate. The hippocampus meets all of the
criteria specified above.
Luria concluded that "many nuerons in the hippocampus and
connected nuclei do not respond to modality-specific stimuli of
any sort, but serve to compare present stimuli with traces of
past experience; they react to every change in the stimulus and
thus play to some extent the role both of 'attention neurons' and
of 'memory neurons'" (1973a, p. 289). According to Luria the
hippocampus provides for the "elimination of responses to irrelevant
stimuli and enables the organism to behave in a strictly selective
manner" (1973a, pp. 271-272). It appears that the hippocampus
accomplishes this complex task by coordinating the activities of
the cortical and subcortical mechanisms which are directly in
volved in the processes of attention, memory and learning.


97
such stimulation demonstrated excellent memory for the learning experi
ence. Kessner interpreted this result as the failure to encode the
negative affect associated with the experience.
In order to discriminate the relative contribution of the
amygdala and the hippocampus to these phenomena Baker, Kessner,
and Mi chai (1981) took advantage of the fact that suitable reminder
cues often serve to facilitate existing memories and thus enhance
recall. Rats with electrodes implanted in the amygdala or hippo-
capmus and unoperated animals received a single training trial with
footshock (FS) while licking a water tube in a goal box. Retention
was evaluated 24 hours later as an increase in latency to enter the
goal box and lick the tube at least ten times. Following this first
retention test four of the groups received a reminder cue (non
contingent FS) in a different enviroment, followed immediately by
hippocampus or amygdala stimulation or no stimulation. Twenty-three
hours later all groups were tested a second time for retention of
the passive-avoidance learning in the original apparatus. Operated
and nonoperated controls that received a reminder cue but no brain
stimulation exhibited a marked increase in latency on the second
(relative to the first) retention test, demonstrating that the
reminder cue effectively enhanced retention of the aversive experience.
Conversely, operated and nonoperated controls which did not receive a
reminder cue showed a decrease in latency on the second test,
indicating that some extinction had taken place due to re-exposure to
the original apparatus during test number one. Animals whose amygdala
function was disrupted by electrical stimulation following the reminder


173
Gardner, W. J. Removal of the Right Cerebral Hemisphere for Infil
trating Glioma. Journal of the American Medical Association,
1933, 101, 822-826.
Gardner, W. J., Karnosh, L. J., McClure, C. C., & Gardner, A. K.
Residual Function Following Hemispherectomy for Tumour, and for
Infantile Hemiplegia. Brain, 1955, 78, 487-502.
Gazzaniga, M. S. The Bisected Brain. Mew York: Appleton-Century-
Crofts, 1970.
Gazzaniga, M. S. Consistency and Diversity in Brain Organization.
Annals of the New York Academy of Sciences, 1977, 299, 415-423.
Gazzaniga, M. S., & Ledoux, J. The Integrated Mind. New York:
Plenum Press, 1977.
Gazzaniga, M. S., Risse, G. L., Springer, S. P., Clark, E., &
Wilson, D. H. Psychologic and Neurologic Consequences of
Partial and Complete Cerebral Commissurotomy. Neurology,
1975, 25, 10-15.
Geshwind, N. Disconnection Syndrome in Animals and Man. Brain,
1965, 88, 237-294.
Geshwind, N. Selected Papers on Language and the Brain. New York:
D. Reidel Publishing Co., 1974.
Geshwind, N. Specializations of the Human Brain. Scientific
American, 1979, 241 (3), 180-199.
Geshwind, N., & Levitsky, W. Human Brain: Left-Right Asymmetries
in Temporal Speech Region. Science, 1968, 161, 186-187.
Gionotti, G. Emotional Behavior and Hemispheric Side of the Lesion.
Cortex, 1972a, 8, 41-55.
Gionotti, G. Studies on the Functional Organization of the Minor
Hemisphere. International Journal of Mental Health, 1972b
1, 78-82.
Gionotti, G., & Lemmo, M. Comprehension of Symbolic Gestures in
Aphasia. Brain and Language, 1976, 3, 451-460.
Glass, D.H., Ison, J. R., & Thomas, G. J. Anterior Limbic Cortex and
Partial Reinforcement Effects on Acquisition and Extinction of
a Runway Response in Rats. Journal of Comparative Physiological
Psychology, 1969, 69, 17-24.
Glees, P. Experimental Neurology. London: Oxford University
Press, 1961.


124
guarantees that the species would continue to survive. (The
psychological correlates of emotion and arousal activated by the
left hemisphere will be discussed in the section on psychopathology,
below!)
The normal chain of events within the meta-system might be
traced as follows (see Fig. 4): neuronal signals, originating
in the sensory receptors, which reflect novel or potentially
significant environmental stimuli excite the brainstem raphe'
nuclei by way of sensory nerve collaterals. The raohe' nuclei activate
the reticular formation resulting in generalized cortical arousal
(orienting). (1) The right hippocampal system responds to environ
mental stimuli by causing the (cortical) memory traces which
represent the organism's previous experience with similar stimuli
to be activated; (2) if any of these memory traces contain coded
emotional attributes the right amygdaloid como!ex is activated by
them; (3) the amygdala, (a) acting via the septal area, focuses
the hippocampal memory search on the significant memory trace with
the secondary effect of releasing hippocampal inhibition of the
reticular formation and (b) influences the reticular formation
directly to release mobilizing energy and (c) relays its activation
to the left amygdala via the anterior commissure; (4) left amygdala
activation is then forwarded (via the dorsomedial thalamic nucleus
to the left orbitofrontal cortex where it achieves conscious
awareness as subjective emotional experience; (5) the dorsolateral
area of the left prefrontal lobe utilizes this motivational informa
tion to formulate intentions and will attempt to resolve the


78
Data on the fornix is relatively scarce, and there have been
reports of negative findings. In his influential review, Brierly
(1977) took a very conservative stance on this subject:
The most discrete link between the hippocampal and
diencepahlic regions is the fornix. It is surprising,
therefore, that with the exception of the case reported
by Sweet, Tall and and Ervin (1959), bilateral division
of the fornix (usually in the region of the intra
ventricular foramen) has not resulted in a disorder
of memorizing (Dott, 1938; Cairnes & Mosberg, 1951;
Garcia-Bengochea and his colleagues, 1954). This
finding suggests that the two groups of structures
linked by the fornix cannot be regarded as a unitary
system subserving the process of memorizing, at least
until major interconnections other than the fornix
have been identified, (pp. 221-222)
Other interconnections are available. Smithies (1966) describes
a "massive direct hippocampal-hypothalamic pathway" that runs
diffusely through the subthalamus and which he suggests might be
"quite able to carry on hippocamoal and limbic circuit function in
the absence of [the fornix]" (p. 122). A seond look at the reports
cited by Brierley, however, allows the possibility that his
conclusions are premature.
Sweet et al. (1959) emphasized that their patient's "conversa
tions, social amenities, and general demeanor gave little evidence
of [her] severe deficits unless they were specifically looked for"
(p. 76). They also noted that she lacked spontaniety, made little
effort to converse, and was apparently indifferent to her deficits
(cf. Korsakoff's syndrome). The very brief report of Bengochea,
De la Torre, Esquival, Vieta and Fernandez (1954), after a "short
follow-up" of their patients (whose fornices were severed in an
experimental operation to relieve intractable epilepsy) did not


99
Mishkin and Aggleton (1981) reviewed data which demonstrate
that the amygdala receives highly processed sensory information
from all of the secondary and polymodal association areas of the
posterior neocortex in addition to its reciprocal connections with
the hypothalamus. They suggested that these connections provide a
mechanism for complex stimuli to become intergrated with, and
later, to evoke emotional responses which are organized at the
level of the hypothalamus.
Smythes (1966) suggested that the hippocampal system may lay
down memories and the amygdala system may determine what memories are
to be laid down. While this may overstate the case somewhat, it
is clear that these two functional systems work synergistically to
provide the organism with pertinent information from its past
experience which is caoable of motivating and facilitating appropriate
responding based on the requirements of the situation. The septal
area appears to play an essential role in the interaction of the
amygdalar and hippocampal systems and may integrates their activity
with that of the brain's arousal systems.
The septal area is interposed anatomically and functionally
between the amygdala and hippocampus (via the stria terminal is
and fornix, respectively) and has a reciprocal relationship with
both (Swanson & Cohen, 1976). The septum is connected with the
brainstem reticular formation (via the MFB) and is in a position
to modulate the activity of the ARAS through its connection with
the habenula (via the stria medularis), which is a station between the
RF and ILTN. (All of these anatomical relationships are illustrated


126
simultaneously. They found that a nonverbal WS reduced RTs of
both hands equally, no RT asymmetries occurred when the verbal WS
merely forewarned the subject (simple RT condition), right hand
RTs were faster than left following a verbal WS only when a response-
linked decision process was required (the go/no go condition in
which the WS both forewarned and dictated whether or not a response
should be made). The results of these two studies suggested to the
authors that
The right hemisphere may activate the left hemisphere
via interhemispheric pathways, wherease the extent to
which the left hemisphere can activate the right hemi
sphere is considerably less. Essentially, a 'one-way
street' is proposed in which the right hemisphere
activates the left more than the left hemisphere acti
vates the right (i.e., asymmetric interhemispheric
activation). (Bowers & Hielman, 1976, p. 7)
Altered GSR following unilateral brain injury. Hielman,
Schwartz and Watson (1978) studied arousal responses in patients
with left hemisphere damage (an aphasia syndrome), right hemisphere
damage (with the neglect/indifference syndrome), and no brain
damage, by electrically stimulating the forearm ipsilateral to
the brain injury and measuring GSR from the fingers on that side.
They found that the right hemisphere group showed singificantly
less GSR than either the left hemisphere or control groups (five
of the seven right hemisphere pateints had no measurable GSR at
all). The left hemisphere group had an exaggerated GSR relative
to the controls. These results were not attributable to differences
in sensory input or lesion size. The authors concluded that patients
with right hemisphere injuries and neglect have defective arousal.


80
It should be noted that a colloid cyst of the third ventricle
tends to produce confusion, dulling of attention and memory, and
sometimes a progressive dementia prior to its surgical removal.
These factors would make a pre-post evaluation of memory function
very difficult. Still, the scantiness of the reported date in the
studies reviewed above is unfortunate. It is evident that injuries
to different parts of this system result in different expressions
of the disorder. It is reasonable to conclude, however, that
damage to the fornix has adverse effects on memory function which
vary as to the quality and degree, and may leave a greater possibility
of recovery of function.
Mammillary bodies and mammillothalamic tract. The mammillary
bodies are a collection of nuclei at the posterior boundary of the
hypothalamus. They form a major relay station for hippocampal
output on its way to the thalamus (via the mammillothalamic tract)
and to the midbrain reticular formation (by way of the marnmillo-
tegmental tract).
In humans, damage which is apparently limited to the mammillary
bodies has resulted in the full Korsakoff amnesic syndrome (Remy,
1942; Delay & Brion, 1951; Gruner, 1956; Symonds, 1966), although
Victor (1964) suggested that additional damage to the thalamus
was necessary to produce the disorder. In the rat, lesions of the
mammillary bodies or of the mammillothalamic tract impaired the
ability to perform a spatial discrimination in a T-maze in order to
avoid footshocks (Thompson, Langer & Rich, 1964). Krieckhaus
(1962, 1964) found that complete or partial destruction of the


131
Penfield, W. The Permanent Record of the Stream of Consciousness,
Proceedings XIV Of the International Congress of Psychology,
Montreal. Acta Psychologica, 1954, 47-69.
Penfield, W. The Mystery of the Mind. Princeton: Princeton
University Press, 1975.
Penfield, W., & Evans, J. The Frontal Lobe in Man: A Clinical Study
of Maximum Removals. Brain, 1940, 58, 115-138.
Peretz, E. The Effects of Lesions of the Anterior Cingulate Cortex
on the Behavior of the Rat. Journal of Comparative and
Physiological Psychology, 1960, 53, 540-548.
Perris, L., Rosadini, G., & Rossi, G. F. Determination of Side of
Cerebral Dominance with Amobarbital. Archives of Neurology,
1961, 4, 173-181.
Peterson, L. R. Search and Judgement in Memory. In B. Kleinmantz
(Ed.), Concepts and the Structures of Memory. New York:
John Wiley & Sons, 1967.
Petrie, A. Personality and the Frontal Lobes. New York: Blakeston,
1952.
Petrie, A. Some Psychological Aspects of Pain and Suffering. Annals
of the New York Academy of Science, 1960, 86^, 13-27.
Petrie, A. Individuality in Pain and Suffering. Chicago: University
of Chicago Press, 1967.
Powell, B. J. A Study of the Perceptual Field Approach of Normal
Subjects and Schizophrenic Patients Under Conditions of an
Oversize Stimulus. Unpublished Doctoral Dissertation,
Washington University, 1964.
Pribram, K. H. Memory and the Organization of Attention. In D. B.
Lindsley & A. A. Lumsdaine (Eds.), Brain Function (Vol. 4).
Berkeley: University of California Press, 1967.
Pribram, K. H. The Primate Frontal Cortex Executive of the Brain.
In K. H. Pribram & A. R. Luria (Eds.), Psychophysiology of the
Frontal Lobes. New York: Academic Press, 1973.
Pribram, K. H., & McGuinness, 0. Arousal, Activation and Efforts in
the Control of Attention. Psychological Review, 1975, 82, 116-
149.
Price, J. L. The Efferent Projections of the Amygdaloid Complex in
the Rat, Cat and Monkey. In Y. Ben-Ari (Ed.), Inserm Symposium
No. 20, The Amygdaloid Complex. New York: Elsevier/North-Holland
Biomedical Press, 1981.


23
In spontaneous speech the key substantives are often missing
and the remaining parts of speech lack their organizing influence.
Grammatical parts and forms are used abundantly but incorrectly.
Adjectives,, adverbs, conjunctions and prepositional phrases are
strung together haphazardly and the result may resemble a schizo
phrenic's "word salad" (Luria, 1973). Nonsense syllables and neo
logisms are frequent. The guiding thought behind a verbal produc
tion may be evident, but it becomes obscured by tangential associa
tions and incomprehensible babbling.
The Wernicke's aphasic understands little of what is said and
seems to rely on nonverbal cues in order to respond to a situation.
They are also unable to comprehend nonverbal symbolism and so their
communication is vague and concrete at all levels.
Perhaps the most striking feature of Wernicke's syndrome is
the patients.' complete lack of awareness of and indifference to their
deficits. They appear unconcerned and will vehemently deny any
problems.
The same deficits are evident in reading and writing tasks.
The patient can read single words but does not seem to grasp their
meaning or relate them to him or herself. They may correctly
repeat a simple command written on a card but will make no attempt
to comply. Unable to formulate an acceptable sentence on their
own, they are able to arrange cards with words printed on them to
form a syntactically correct sentence. However, the key substan
tives are likely to be misplaced (e.g., "The man bit the dog.")
confirming again that the patient's facility is with granmar as
opposed to meaning.


117
punishment by withholding that response (Isaacson, Douglas, Lubar
& Schmaltz, 1973, note that the inhibition of a response is actually
an active process). In the one-way active avoidance problem the
animal must learn to actively avoid punishment (e.g., footshock
[FS]) by making a specified response (e.g., jump over a hurdle in
response to a conditioned stimulus [CS1 which signals an impending
FS). In the two-way active avoidance task the animal must jump
between the two compartments of a shuttle-box on alternate trials
in response to a CS in order to avoid punishment. Isaacson et al.
(1971), point out that this situation produces a conflict between an
active- and a passive-avoidance response: on any given trial the
animal must leave the compartment it is in to avoid the signaled
FS (active-avoidance) and must enter the compartment in which it
was shocked on the previous trial (passive-avoidance). In the
differential reinforcement of low rates of responding (DRL)
situation the animal must learn to withhold responses for a pre
determined amount of time after the last response before a response
will be rewarded. On the fixed interval (FI) reinforcement
schedule normal animals learn to stop responding in the first half
of the interval following a reinforced response and to slowly
increase their rate of responding in the second half of that
interval. Continued responding in the first half of the interval
is considered to be a perseveration error; higher than normal rates
of responding in the second half of the interval are considered
anticipatory errors.


113
reciprocal connections with the hypothalamus (via the stria terminal is
and the ventral amygdalofugal fiber system) and with the septal
nuclei (via the stria terminal is).
The amygdala is involved in the integration of information
from the internal and external enviroments which relates specifically
to the survival of the individual and its ability to reproduce.
Based on such an integration, the amygdala attaches motivational
significance to previously neutral stimuli by associating those
stimuli with their internal consequences. The amygdala produces
the signals which permit the positive and negative attributes of
an experience to be encoded in memory through an interaction with
the hippocampus (via the septal area). When that memory trace is
reactivated on a later occasion the amygdala responds to its encoded
emotional and reinforcing attributes, computes the reinforcement
value of the present stimulus in light of the immediate internal
status of the organism, and participates in focusing the hippocampal
systems on significant memory traces. Qualitative and quantitative
motivational information is relayed from the amygdala to the orbital-
frontal cortex where it is translated into a subjective experience
(.e.g, fear, anger, anxiety, etc.) of appropriate intensity. The
tertiary association area in the Tdorsolateral prefrontal cortex
responds to this experience and will attempt to resolve it by
programming the post-central problem-solving mechanisms to formulate
a behavioral response to the stimulus. The amygdala may also
initiate activity in the sympathetic and parasympathetic control
mechanisms in the hypothalamus to prepare the organism physically


56
3. "Illusory" emotional experiences.
Based on his analysis of the data, Penfield (1975) postulated the
existence of two related brain systems: a "mechanism of recall,"
and a "mechanism of interpretation." The latter involved the temporal
cortex (exclusive of the speech areas) and was referred to by Penfield
as the "nonverbal concept mechanism." Penfield comoared its func
tion with nonverbal concepts to the operation of the speech cortex
with verbal concepts.
Somehow [this mechanism] seems to analyze the components
of sensation, compares them with previous experience,
and by that analysis and comparison, transmits into
consciousness their present and immediate significance
... [an emotional response] is a signal that rises
into consciousness as a result of an interpretation of
what the present situation may bring the subject in
the immediate future.. (Mullan & Penfield, 1959, p. 283)
It appears that the cognitive products of these mechanisms fit
the criteria for generalized expectations. Penfield's evidence
assists in understanding the different types of memory referred
to by Rapaport and by Breuer and Freud and indicates the neural
substrate of these processes. (Penfield's data and conclusions
will be reviewed and evaluated in detail presently.)
It has been assumed here that at the base of personality there
are mechanisms whereby cognitive operations and affective processes
are appropriately activated by significant memories. The present
task is to describe neurological evidence which accounts for the
memory phenomena reviewed above in terms that fit the criteria
for "functional systems" as defined by Luria, and satisfy the
evolutionary imperatives outlined at the beginning of this chapter.


147
(for the treatment of depression) within 48 hours. A second rod-
and-frame test was administered five hours after the treatment.
Twelve subjects whose first treatment was rescheduled for non-
clini cal reasons served as controls and were retested five hours
after the missed first appointment. The results were highly
significant; all twelve left ECT patients showed more field-
dependence on the second test* all twelve right ECT patients
showed less field dependence* the controls showed little or no
change (Cohen, Berent & Silverman, 1973).
Witkin (1965) relates the field-dependence-independence
perceptual dimension to a cognitive differentiation dimension:
the person who is field-dependent also does less well at "solving
problems which require isolating essential elements from the con
text in which they are presented and using them in different
contexts" (1965, p. 319). The cognitive differentiation dimension
is manifest in a global/diffuse versus an articulated cognitive style.
The field-dependent/global style has been associated with
hysterical neurosis (Zukmann, 1957), character disorders, somatiti-
zation, alcoholism, and patients whose primary symptom is affective
discharge (see the review by Witkin, 1965). A field independent/
articulated personality style has been associated with paranoia
(Janucci, 1964; Powell, 1964) and obsessive-compulsive disorders
(Zukmann, 1957).
Sociopathy and Hysteria
The absence of anxiety and the inability to learn from experience
are pathonomic in sociopathy. In contrast to anxiety neurotics,


187
Whitty, C. W. M. Some Early and Transient Changes in Psychological
Function Following Anterior Cingulectomy in Man. International
Journal of Neurology, 1966, 3-4, 403-409.
Wigan, A. L. The Duality of the Mind. London: Longman, 1844.
Winocur, A., & Mills, J. A. Transfer Between Related and Unrelated
Problems Following Hippocampal Lesions in Rats. Journal
of Comparative and Physiological Psychology, 1970, 73_, 162-169.
Wise, C. D., Berger, B. D., & Stein, L. Serotonin: A Possible Mediator
of Behavioral Suppression Induced by Anxiety. Disorders of the
Nervous System, 1970, 21 34-37.
Wishart, T. B., & Mogunson, G. J. Effects of Lesions of the Hippo
campus and Septum Before and After Passive Avoidance Learning.
Physiology and Behavior, 1970, 5, 31-34.
Witkin, H. A. Psychological Differentiation and Forms of Pathology.
Journal of Abnormal Psychology, 1965, 70 (5), 317-336.
Zaidel, E. The Elusive Right Hemisphere of the Brain. Engineering
and Science, 1978, 2 10-19.
Zangwill, D. L. Psychological Deficits Associated with Frontal Lobe
Lesions. International Journal of Neurology, 1966, 3-4, 395-409.
Zuckerman, M. Sensation Seeking and Cortical Augmentinq-Reducing.
Psychophysiology, 1974, 11, 535-542.
Zukmann, L. Hysteric Compulsive Factors in Perceptual Organization.
Unpublished Doctoral Dissertation, New School for Social
Research, 1957.


CHAPTER IV
RESULTS
Left versus Right Hemipshere Cognitive Functioning
Between Groups
Hypothesis 1 stated that the hyper- and hypo-dominant spectrum
disorders would differ on the ratio of scores on tests that have
been demonstrated to be sensitive to left and right hemisphere
cognitive functioning. Specifically, it was hyoothesized that the
hyper-dominant group would show relatively better performance on
the Similarities and Information WAIS-R Subtests and the hypo-
dominant group would show relatively better performance on the
Street Test and the WAIS-R Object Assembly Subtest. The mean
ratio (Street + Object Assembly/Si Hilarities + Information) for the
hyper-dominant group was x = 1.15 (SD = 0.42) while the mean ratio
for the hypo-dominant group was x = 1.91 (SD = 0.84), indicating
that the groups differed in the predicted directions. A student's
t-test for independent samples (Robson, 1973) performed on these means
revealed that the differences between groups was significant
(t = 3.4640, p < 0.001, one-tailed). It was concluded that
psychiatric inpatients and outpatients assigned to the hyper
dominant spectrum disorder group performed better on tests sensitive
to left hemisphere cognitive functioning relative to tests
sensitive to right cognitive functioning; their ratio scores
156


16
on new sources of information about the internal and external
environments and to utilize new and more efficient forms of learning
in organizing its responding. It is important to appreciate that
the final product of limbic system operations in the paleomammalian
brain is the inhibition of lower centers. However, these same
mechanisms later came to exert important influences on the neo-
cortical systems.
The cerebral cortex of the neomammalian brain developed in close
association with the limbic system and basal ganglia. New nuclei were
added to existing sensory and motor systems, culminating finally
in the arrangement now found in primates. More efficient information
processing in the neocortical additions permits more precise sensory
discriminations and rapid, fine-grain movements of the extremities
(Isaacson, 1974) but these new systems still operate in close con
junction with the older subcortical mechanisms (Schade1 & Ford,
1973).
The importance of the limbic system in human experience can
hardly be overstated. These structures are implicated in most,
if not all, forms of psychopathology. They are the probable sub
strate for the therapeutic effect of most psychotropic medications
(Broekkamp & Lloyd, 1981) and the targets for all forms of "psychia
tric surgery." Unfortunately, existing theory regarding the
functional significance of the limbic system is primarily descriptive
in nature or consists of generalizations so broad as to be of
little use to the applied psychologist.


t
TEMP CTX
HPC MB
Ant. Thai. CING
>IPL
00
CT>
Papez's circuit
Figure 3. The position of Papez's circuit within a larger cortical circuit. (TEMP CTX -
temporal cortex; HPC hippocampus; MB mammillary body; Ant. Thai anterior nuclei of the
thalamus; CING cingulate gyrus; IPL interior parietal lobule.


30
occurrence of signs revealing the capacity of the subject to keep in
contact with the external world" (p. 103). They acknowledged that
"the suppression of expressive and receptive speech functions make
such a task quite difficult with the patients receiving barbiturate
in the dominant hemisphere" (p. 109). It seems that it is only
the appearance of aphasiz (after dominant hemisphere anesthetization)
that is specific to human consciousness in this study, and these
findings are consistent with those of Terzian and Serafetini des
et al.
Rosadini and Rossi did not report the results of their test of
subjects' ability to recall what had occurred during the Wada
procedure but this question was addressed directly in an experiment
reported by Gazzaniga (1977). This author found that "information
encoded while the left hemisphere was anesthetized was uninterpretable
by the verbal system when the left hemisphere returned to normal
functioning . when information is encoded by other than the
verbal system the person is not consciously aware of the information"
(p. 150).
Another approach to the localization of conscious awareness
involved an analysis of the temporal discrimination for simultaneity
when two visual stimuli were presented separately to the left and
right visual half-fields, separated by a very brief interval.
Efron (1963a; 1963b) found that normal right-handed subjects reported
that the two flashes occurred simu"!taneously only when the
light flashed in the left visual half-field was presented several
milliseconds earlier than the light flashed in the right visual field.


3
structure and describe the ways in which these interact to produce
psychological health and psychopathology. Such a model might lead to
new operational definitions of psychological phenomena which, in
turn, may suggest new intervention points and methods.
The purpose of a theory is to integrate known facts within a
single framework and account for them in terms of a small number of
interrelated concepts. Existing theories suffer from a lack of
integration. The discipline of psychology has hindered such integra
tion by institutionalizing a tradition of subspecialization: theories
of attention, perception, cognition, motivation, emotion, behavior,
etc. are developed in relative isolation. The failure of individual
theorists to understand and to appreciate the interrelatedness of
these various personality operations may account for the inadequacy
of psychological theory in general. It is assumed here that the
component parts of the personality can only be properly understood
in the context of their relationship to the whole; that more will
be gained from a gross (but comprehensive and testable) description
of the personality infrastructure than from a detailed examination
of a single personality operation.
Rather than subdivide the personality structure on the basis of
notions of what it should do; it might be more productive to approach
the problem on the more basic level of what it must do. The formula
tion of the model will be guided by a set of assumptions about the
evolutionary pressures which shaped the personality structure.
These "evolutionary imperatives" are as follows:


132
they have difficulty sustaining mental activity; they become ego
centric, shallow, and seemingly unaware of the social consequences
of their behavior (Gardner, 1933; Rowe, 1937; Bell & Karnosh,
1949; Mensh, Schwartz, Matarazzo & Matarazzo, 1952; Gardner et al.,
1955; Austin & Grant, 1962; Bruell & Albee, 1962). It may be deduced
from the above that normal, adaptive, responsiveness to the environ
ment depends on the products of right hemisphere processes. The fact
that the emotional impact of a situation is available to the left
hemisphere in split-brain patients but not in right hemispherectomy
patients indicates that the right cerebral hemisphere in humans
is responsible for making determinations regarding the significance
of environmental stimuli. The contrast between the cerebral dis
connection and right hemispherectomy syndromes further suggests
that relatively normal emotional resDonsiveness, judgement, and
adaptability can be maintained as long as the motivating and mobilizing
products of the right hemisphere's cognitive operations can be
transmitted to the left (i.e., can activate the left amygdala and
ARAS). In the absence of the right hemisphere's modulating input
the left hemisphere is subject to erratic and inappropriate emotional
experience because the neocortex on that side is not adequately
prepared to mediate these processes.
Patients with lesions in their left cerebral hemispheres
typically have a catastrophic/depressive reaction while those with
right hemisphere damage show a characteristic indifferent/euphoric
response (e.g., Gionotti, 1972a). Transient, but similar emotional
reactions are commonly seen during recovery from unilateral


133
hemisphere anesthetization using the Wada technique (e.g., Pern's,
Rosadini & Rossi, 1961) and have been reDorted after unilateral
ECT (Deglin & Nikolaenko, 1975). Such findings have prompted
speculation that each hemisphere tends toward a different emotional
state.
The fundamental problem with the lateralized emotional valance
theory is the fact that the actual direction of the valance cannot
be specified. Tucker (1981) notes that the phenomena observed
following the unilateral loss of cortical function might result
from the disinhibition of the emotional tendency of the opposite
hemisphere or from the release of subcortical processes on the
same side.
Dimond, Farrington, and Johnson (1976) reported experimental
evidence suggesting that the right hemisphere has a negative
valance. They presented three short films to normal subjects'
left or right visual half-fields (using a special arrangement of
spectacles and contact lenses) and to free-viewing controls. The
right hemisphere group rated the films as significantly more
unpleasant and horrific than did the left hemisphere and control
groups (who did not differ from each other). Dimond et al.,
concluded that the right hemisphere tends toward a negative
emotional appraisal of incoming stimuli similar to the "characteristic
perception of the depressed patient" (p. 691). This finding would
seem to support the contralateral disinhibition interpretation of
the emotional phenomena seen after unilateral brain injury.
However, the contralateral disinhibition notion is discredited by


120
noted that rats with FNX cuts or septal lesions were hyperactive and
avoided mainly by shuttling spontaneously between compartments,
expending a great deal of energy in the process. The fact that
animals with MFB cuts were not hyperactive and learned to respond
approriately to the CS suggested to Grossman that the altered
responding by HPC and septal animals in this paradigm was due to
"an interruption of reticulo-septo-hippocampal interconnections"
(1076, p. 395).
Grossman (1976) reported that sectioning the FNX, ST, or MFB
failed to mimic the deficit in acquiring the one-way active avoidance
task seen in animals with amygdalar, septal, and hippocampal damage.
As noted earlier, an animal should be able to encode and retrieve
one strategy in a particular situation (as long as no competing
responses were available) if the cortex has access to amygdala
generated emotional information during training. An alternate
pathway for this amygdala information is available via a branch from
the ST to the SM (see Fig. 2). The one-way deficit was reproduced
by sections :of the SM (which appear to be downstream from the ST
branch) performed by Ross in Grossman's laboratory (Grossman,
1976).
A Functional Meta-system
Human beings differ from lower forms in that a large part of
human brain development and organization occurs postnatally.
The special and evolved integrative functions which distinguish homo
sapiens are related to specialization of the secondary and specifically
human tertiary association areas which do not reach functional


14
lower animals depend on the sense of smell for such survival-
related activites as food-getting, the detection of enemies, and
mating (MacLean, 1949), the role of these subcortical structures
in motivational and emotional processes was not appreciated until
the 1930s. Since Kluver and Buey (1938) published their description
of altered emotional behavior following the bilateral removal of
the temporal lobes, including the amygdalae and hippocampi, limbic
system components have been the subject of intense scrutiny. The
massive literature which has accumulated is filled with confusing
inconsistencies and conflicting findings. This may be due in part
to the fact that the function of these structures appears to vary
according to enviromental circumstances (Olds, 1958). Although
a number of tentative models of limbic system function have been
offered (e.g., Papez, 1937; Gloor, 1956; Olds, 1958) none has
found general acceptance.
MacLean (1970) distinguishes three major types of systems in
the mammalian brain which correspond to stages in its evolutionary
development. He describes a protoreptilian core-brain, a paleo-
mammalian brain (the limbic system) and a neomammalian brain (the
neocortical areas). This triune brain conceptualization provides
a useful perspective on the hierarchical arrangement of anatomical
and functional systems and their relationship to behavior (Isaacson,
1974).
The protoreptilian brain represents the fundamental core of
the nervous system consisting of areas homologous to parts of the
upper brainstem and midbrain, hypothalamus, and basal ganglia.


I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is
fully adequate, in scope and quality, as a dissertation for the degree
Doctor of Philosophy.
irry Grater/ Chairman
Professor of Psychology
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is
fully adequate, in scope and quality, as a dissertation for the degree
Doctor of Philosophy.
James I. Morgan
Associate Professor of
Counselor Education
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is
fully adequate, in scope and quality, as a dissertation for the degree
Doctor of Philosophy.
Paul Schauble
Professor of Psychology
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is
fully adequate, in scope and quality, as a dissertation for the degree
Doctor of Philosophy.
David Suchman
Professor of Psychology
I certify that I have read this study and that in tpy opinion it
conforms to acceptable standards of scholarly presentation and is
fully adequate, in scope and quality, as a dissertation for the degree
Doctor of Philosophy.
Robert- Ziller
Professor of Psychology
- N


22
of language is generally intact. The ability to understand and
utilize nonverbal symbolism (e.g., gesture, pantomime) is also
spared. Intellectual functioning is relatively unimpaired. Patients
retain the ability to reason logically, to abstract and to generalize,
and to respond to context appropriately. Their associational
processes are not loosened, tangential or pressured. They do not
produce paraphasias or confabulate. Finally, these patients are
acutely aware of their deficits; they may be appropriately depressed
and are prone to sudden, transient emotional outbursts.
Exactly the opposite clinical picture is evident when Wernicke's
area, on the lateral, convex surfaces of the left temporal lobe, is
damaged. Wernicke's aphasia is characterized by impaired language
comprehension with fluently articulated but nonsensical speech.
Unlike the Broca's aphasic, words are spoken clearly with normal
sounding cadence, intonation and melody (prosody). However, the
speech of the Wernicke's aphasic is lacking in content and may
consist almost entirely of semantic jargon which has little communi
cation value. These patients appear to have lost control of the
language mechanisms at all levels. It is as if the selection
thresholds for phonemes, words, and ideas were all lowered.
Repetition is poor and marked by paraphasic errors in which the
correct sounds may be present but emerge in the wrong order.
Patients are able to name only the most familiar objects accurately,
although the word produced may come from the same category as the
target. Prompting with the initial wordsounds seldom helps.


49
Watson and Hielman (1976) who showed also that right hemisphere
patients were impaired in the vocal expression of emotion. The
efforts of patients with right temporoparietal damage to impart a
sad, happy or angry tone to their voices were rated as incorrect
significantly more often than controls. Ross and Mesulam (1979)
presented case studies of two well-educated patients who had com
parable damage in the right supra-sylvian area, which is homologous
to Broca's area on the left side. Both patients showed flattened
affect and had completely lost the ability to laugh, cry, or
otherwise express any emotion in their speech. Their ability to
experience and comprehend emotions was unchanged. The authors
noted that the organization of emotion in the right hemisphere
seems to mirror that of language in the left: the area homologous
to Wernicke's area being essential for comprehension, and to Broca's
area, for expression.
Gazzaniga and Ledoux (1977) suggested that right hemisphere
functioning in humans is distinguished only by contrast to the
left; it continues to perform its functions in the same manner
as elsewhere in the phylum. Although it should be noted that the
human right hemisphere is in possession of tertiary association
areas and so would perform those tasks more efficiently, the
data reviewed above are not inconsistent with Gazzaniga's interpre
tation. The brains of lower forms (and, apparently, the right
brain in humans) are primarily concerned with neuronal signals
which represent the survival needs of the organism within the
immediate environment. Interaction with the social environment


46
Amygdala Circuits and the Prefrontal Lobes
The role of the amygdala in emotional processes established by
K1uver and Buey in 1938, has been assumed to be affected through this
structure's close relationship with the hypothalamus. The amygdala
seems to direct behavior toward biological goals (Halgren, 1981)
and is implicated in the control of species-specific behaviors related
to survival needs, including defensive and aggressive behaviors,
sexual activity, and feeding (Isaacson, 1974). In lower forms these
processes might depend on a simplified (instinctive) form of
memory in which stimulus and response are yoked (Pribram & McGuiness,
1975). In addition to mediating emotional states the amygdala is
involved in the analysis of reinforcement contingencies. Amygdala
lesions have been shown to produce impaired recognition of stimuli
associated with rewards (Weiskrantz, 1956; Schwartzbaum, Thompson
& Kellicut, 1965; Jones & Mishkin, 1972) and inability to resDond
appropriately to changes in the magnitude of rewards (Schwartzbaum,
1960).
Strong interconnections with the hypothalamus (via the stria
terminalis and ventral amygdalofugal fibers) give the amygdala
immediate access to information concerning the internal status of
the organism (Price, 1981). The amygdala also receives processed
sensory information from all of the secondary sensory association
areas (Van Hoesen, 1981). Mishkin and Aggleton (1981) noted that
this arrangement places the amygdalae in a position to integrate
external events with their internal consequences, which would
permit the attachment of emotional and motivational significance


112
coding/decoding operations through a two-way septal-hippocampal inter
action. The memory indexing process is further modulated or pro-
grammed by processed motivational input from the prefrontal lobe
to the cingulate cortex. This system accomplishes three principal
tasks in the course of its operation: (a) it monitors incoming
stimuli and matches them against cortical representations of the
organism's previous experience with similar stimulus configurations;
(b) when a potentially significant stimulus configuration is
encountered the system activates the organism by switching on the
Mobilization System; and finally, (c) it causes pertinent informa
tion related to those stimuli to be presented to conscious awareness.
It appears that there are two such systems in the human brain,
each specialized to deal with a different type of information. The
first, lateralized to the right hemisphere, performs the functions
listed above with experiential data. The second, operating in the
left hemisphere, deals with language and verbal concepts. It is
probable that environmental stimuli trigger an initial search in
the experiential (right hemisphere) system. (It may be noted
that these right hemisphere processes would be unconscious and
that their products might be excluded from consciousness.)
The Motivating System
The Motivating System is centered on the amygdala. This structure
receives processed sensory information from all of the sensory
modalities and is interconnected with the entire prefrontal lobe
both directly (via the uncinate fasciculus) and indirectly (by
way of the dorsomedial thalamic nuclei). It also has strong


169
Bell, E., & Karnosh, L. J. Cerebral Hemispherectomy, Report of a
Case Ten Years After Operation. Journal of Neurosurgery,
1949, 6, 285-293.
Bengochea, F., De la Torre, E., Esquival, 0., Vieta, R., &
Fernandez, C. The Section of the Fornix in the Surgical
Treatment of Certain Epilepsies. Trans. Am. Neurol. Assoc.,
1954, 79, 176-178.
Benton, A. Differential Behavioral Effects of Frontal Lobe Disease.
Neuropsychologia, 1968, 6, 53.
Bergin, A. E., & Lambert, M. J. The Evaluation of Terapeutic
Outcomes. In S. L. Carfield & A. E. Bergin (Eds.), Handbook
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& Sons, 1978.
Black, F. W. Cognitive Effects of Unilateral Brain Lesions Secondary
to Penetratinq Missle Wounds. Perceptual and Motor Skills,
1974, 38, 387-391.
Bogan, J. E. The Other Side of the Brain I: Dysgraphia and
Dyscopia Following Cerebral Commissurotomy. Bulletin of
the Los Angeles Neurological Societies, 1969a, 34 (2), 73-105.
Bogan, J. E. The Other Side of the Brain II: An Appositional Mind.
Bulletin of the Los Angeles Neurological Societies, 1969b,
34 (~3), 135-162.
Bogan, J. E., & Bogan, G. M. The Other Side of the Brain III: The
Corpus Callosum and Creativity. Bulletin of the Los Angeles
Neurological Societies, 1969, 34 (4), 191-220.
Bogan, J. E., DeZure, R., Ten Houten, W. D., & Marsh, J. F. The
Other Side of the Brain IV: The A/P Ratio. Bulletin of the
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Neuropsychologia, 1976, 14, 123-129.
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on Hippocampal EEG. Psychonomic Science, 1970, 18, 181-183.
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1977.


134
the fact that the emotional responses seen with unilateral hemi
sphere anesthetization occur only as the effects of the anesthetic
are clearing: they should be most pronounced when the contralateral
hemisphere was completedly incapacitated (see Tucker, 1981).
The indifference/euphoric reaction seen after right hemisphere
injury is often accompanied by neglect for the left half of the
body and for the left extrapersonal space (Gionotti, 1972b).
This neglect syndrome is usually associated with damage to the
right IPL, although it has been reported after lesions of the right
frontal lobe (Hielman and Valenstein, 1972) which presumably programs
the operations of the right post-central areas. Hielman and
Valenstein note that the mechanism of neglect remains unknown but
observe that
The inability of sensory stimuli to excite or alert an
organism (neglect) cannot be completely explained by
defect in sensory synthesis. The lesions that pro
duce neglect must also interfere with centrifugal
or descending pathways that normally would permit
integrated sensory stimuli to excite or alert the
organism. (1972, p. 663)
These authors suggest that neglect following parietal lobe lesions
may result from disconnection between sensory association areas and
the limbic system. What is most puzzling, and perhaps most
significant, is the fact that neglect is rarely, if ever, seen
following left hemisphere lesions.
The puzzles of lateralized emotional valance and unilateral
neglect resolve themselves if the basic premises of the proposed
functional meta-system hypothesis are accepted. This model
suggests that the primary role of the amygdala is to generate


9
central fissure (Rolando). Incoming somato-sensory, visual and
auditory nerve impulses are relayed, via the thalamus, from contra
lateral receptor surfaces to primary projection areas located in
the parietal, occipital and temporal lobes (see Fig. 1, areas 1, 2,
3; 41; 17), respectively (Noback & Demerest, 1972). In each case the
modally specific, somatotopic organization of nerve impulses in
the projection areas is transformed into functional information
(i.e., acquires meaning) in an adjacent secondary association
area (areas 5, 7, 18, 19; 42). With each new level of processing
there is increasingly complex synthesis of information and decreased
modal specificity (Luria, 1973a). Damage to a primary Drojection
area results in a loss of sensation (e.g., blindness) while lesions
of a secondary association area are likely to produce the inability
to recognize a stimulus in that modality (agnosia). Conversely,
artificial stimulation of a projection area produces a discrete
sensory experience while stimulation of a secondary association
area elicits a more elaborate sensory hallucination whose complexity
is related to the level within the hierarchy that is activated
(see Mullan & Penfield, 1959).
Progression within the hierarchy is reversed in the motor
systems. Specificity of control increases as the secondary (pre
motor) areas (areas 6 and 8) coordinate and fine tune their influence
on the pyramidal cells of the primary motor cortex (area 4) with
the assistance of continuous feedback from the sensory modalities.
Lesions of the primary motor cortex produce contralateral paresis
while stimulation elicits flexion of individual muscle groups.


47
to sensory stimuli. Kessner (1981) reported experimental evidence
that demonstrated the essential role of the amygdala in encoding
and retrieving the positive and negative attributes of a specific
memory. In lower forms the identification of a motivationally
significant stimulus might result in the release of species-specific
behaviors, but in humans behavior is self-determined. Halgren
(1981) concluded from his amygdala stimultion studies with humans
that "the amygdala helps organize the discharge of emotional tension
into consciousness" (p. 404) and noted that this would allow the
directing of consciousness toward biological goals. The amygdala's
input to the neocortex is directed to the entire prefrontal lobe
both directly, via the uncinate fasciculus, and indirectly, by
way of the dorosomedial thalamus (Noback & Demerest, 1972; Price,
1981).
While damage to the dorsolateral area of the prefrontal lobes
has been associated with intellectual disturbances, lesions of the
orbito-frontal cortex (and orbital undercutting, which disconnects
this area from the amygdala) result in emotional changes (Lewin,
1961). Eli thorn, et al. (1958) concluded that this type of damage
produced a "generalized impairment of the ability to form appropriate
emotional responses" (p. 250), including the ability to elaborate
on the affect appropriate to the concepts present in consciousness.
In contrast to the planning deficits, loss of energy and interest,
and affective dullness seen after dorsolateral frontal damage, orbito-
frontal injuries often lead to euphoria, impulsive (disinhibited)
behavior, and the appearance of "greediness, selfishness, and


138
experientially-based alterations in the disposition of the organism
relative to a stimulus object, situation, or event. The cognitive
information contained in the GE can only pass to the left hemisphere
over the neocortical commissures. This exchange permits the verbal
system to symbolize its reinforcement history in a given situation.
It is possible that the more facile mechanisms in the left hemisphere
might be conditioned to somehow prevent this exchange of information
to avoid the evocation of psychological pain. In this case, the
individual would experience the arousal and emotion appropriate
to the stimulus but remain ignorant of its nature, and consequently,
impotent in his or her efforts to resolve the subjective experience.
At least two separate mobilizing systems co-exist within each
half of the brain: a serotonergic system produces indiscriminant
cortical arousal which facilitates the functioning of the monitoring
system in the right hemisphere; a dopaminergic activation and attention
focusing system is essential to the cognitive operations of the left
hemisphere's problem-solving/response-generating system. Any
process which altered the balance between these two systems (within
or between hemispheres) would disrupt normal functioning. The
biochemical mechanisms which form the neural substrata of the
motivating and mobilizing systems are modulated by descending
influences from the prefrontal lobes (via the amygdalae) and from
the temporal lobes (via the hippocampi). The normal functioning of
these modulating pathways might thus be subverted by conditioning
processes (in the interest of "defending the ego") at the expense
of normal, integrated functioning. In the absence of integrated


134
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1974.
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Visual Evoked Responses to Sine Wave Light. Psychophysioloqy,
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57
The neuropathological correlates of human amnesia syndromes and data
from related animal studies will be reviewed. It will be hypothe
sized that human beings possess two autonomous, lateralized memory
systems, centered on the hippocampi, each of which is intimately
associated with its own emotional and cortical activating system.
Human Amnesia Syndromes
A number of terms are used to describe different aspects of
memory function and dysfunction. Immediate memory, usually measured
by digit span, probably reflects the ability to hold information
in the primary or secondary sensory cortex as long as voluntary
attention (directed by the nonspecific thalamic nuclei) is focused
upon it (Smithies, 1966). The terms short-term ("recent") and
long-term ("remote") memory indicate recollection over increasingly
greater periods of time, but both are almost certainly subserved
by the same physiological processes (Brierley, 1977). The term
retrograde amnesia refers to a period of time before an accident or
illness for which the patient's ability to recall is diminished
or lost. Anterograde amnesia is an inability to retain in memory
events that occur after such an injury or illness.
The combination of a severe retrograde amnesia and a debilitating
anterograde amnesia is the hallmark of the Wernicke-Korskoff
syndrome, the most common form of memory disease. This illness is
most frequently seen as a result of brain lesions brought on by
dietary (thiamine) deficiencies in chronic alcoholics, although the
lesions and illness may be produced by a number of toxic or disease
processes (see the excellent review by Brierley, 1977). This illness


43
lesions of the temporal, parietal, or occipital lobes may have
sensory, orientation, or intellectual deficits, but their attention
and concentration remain sustained and directed by intentions.
Luria and his co-workers suggested that the frontal syndrome reflected
the loss of this selectivity (Luria, Homskaya, Blinkov, & Critchley,
1967). An understanding of the functioning of thalamic systems
suggests a mechanism by which the frontal lobes might select or
"recruit" psychological operations in the post-central cortex.
The thalami are a pair of egg-shaped masses located beneath the
cortex in the center of the cerebral hemispheres. The thalamus
is the final processing point for cortical input and the central
integration station of the nervous system. The brief review of
thalamic anatomy and functioning presented below is based on reviews
by Noback and Demerest (1972) and Chusid (1976).
The ventral half of the thalamus contains the specific relay
of nuclei of the sensory-motor systems. The nuclei of the dorsal
tier are association nuclei which have reciprocal connections with
the association areas of the post-central cortex and no sub
cortical connections; the dorsolateral and posterolateral nuclei
are interconneected with the parietal lobe, and the pulvinar with
the temporal and parietal lobe.
The dorsomedial and anterior thalamic nuclei are association
nuclei involved in emotion and memory, respectively. The dorso
medial nucleus receives input from the amygdala and lateral hypo
thalamus and has reciprocal connections with the association areas
of the prefrontal lobe. The anterior nuclei of the thalamus receive


176
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Rat Limbic System and Midbrain Reticular Formation Upon Short-
and Long-term Memory. Physiology and Behavior, 1974, 12_, 5.
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Megaugh (Ed.), The Chemistry of Mood, Motivation, and Memory.
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and Emotional Behavior in the Rat. Ph.D. Dissertation,
University of Michigan, 1961.
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273-283.
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Bulletin, 1968, 70, 285-295.
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8
constitute the basic elements of the personality structure will be
proposed. The eighth section will describe the way in which the
basic elements are organized into a functional meta-system which
forms the infrastructure of personality and functions to assure the
emission of adaptive behavior. This model will be supported with
evidence concerning the psychological correlates of neurological
syndromes. In the final section the psychological phenomena
associated with various forms of psychopathology will be related
to neurological indices which reflect the operation of the lateralized
subsystems and their interaction within the functional meta-system.
It will be concluded that many psychodiagnostic entities may be
classified as hypo- or hyper-dominance spectrum disorders which
are functionally related to chronic, and maladaptive, under- or
over-utilization of the mechanisms in the left hemisphere of the
brain.
Review of Basic Brain Anatomy and Organization
Cortical Mechanisms
Although certain phylogenetically new areas of the cerebral
cortex are of special interest when discussing "higher mental
functions," these areas must be considered in their physiological
and evolutionary context. A brief review of basic brain systems
and anatomy will establish this perspective.
In man, as in all mammals, large portions of the cerebral
cortex are devoted to the more elementary functions of processing
sensory stimuli and the initiation and control of movements. The
brain structures subserving these basic functions are divided at the


44
the output from the hippocampus and have reciprocal connections
with the cingulate cortex.
Lying between and separating the major thalamic association
nuclei, the nonspecific (intralaminar, midline and reticulate)
nuclei have only intrathalamic and subcortical connections. They
receive their main input from the brainstem reticular formation and
from the rostral end of the ascending reticular activating system
(ARAS). The intralaminar nuclei have the "remarkable property of
being able to exert a controlling influence upon the rhythmic
electrical activity of the entire cortex" (Jasper, 1949, p. 406).
Jasper noted that this system is in a position to provide a central
coordinating mechanism for cerebral activities:
A central integrative mechanism with ready access to all
afferent and elaborative systems of both hemispheres,
and closely related to autonomic spring of action; is
necessary to explain consciously directed thought and
behavior. It seems that the thalamic reticular system,
with its diffuse cortical projections, relations to
afferent and efferent systems, relations to mesencephalic
hypothalamic and striatal systems, is a good candidate
for this office. (Jasper, 1949, p. 419)
Discussion
The brain's mobilization systems with their brainstem, thalamic,
and forebrain components, regulate consciousness, unconsciousness,
and the differential consciousness of attention. Sensory signals
representing possibly significant stimuli cause the reticular
formation to initiate diffuse cortical arousal. When a stimulus
has been identified, cortical activation systems facilitate the
organization of cerebral activity to deal with the situation
appropriately. It appears that the frontal lobes direct this process


17
The Interpretation of Neurology Literature
The neurological data which form the basis of this review were
gathered over many decades, from a variety of populations, by
scientists with widely divergent theoretical beliefs. The psycholgist
who is unfamiliar with this literature must become aware of its
inherent problems before venturing interpretations based upon it.
Those who come under the neurologist's care have, almost
invariably, suffered a catastrophic change in their lives and
personalities. Much of the data is drawn from individuals who have
incurred brain damage as a result of a head wound, cerebrovascular
accident (stroke), or cerebral tumor (neoplasm). While offering an
otherwise unavailable opportunity to study higher brain functions,
these devastating "natural experiments" invariably preclude the
rigorous application of scientific method to that study.
Although techniques continue to improve, it is unusually
impossible to define the parameters of a lesion accurately. Thus,
crucial independent variables cannot be precisely isolated and
controlled. Cause and effect interpretations are further
hampered by the fact that a patient's pre-morbid level of functioning
may be impossible to ascertain. Generalizability is also compromised
when the subject has a chronic brain disease, such as epilepsy,
or has undergone a surgical intervention to control that condition,
such as temporal lobectomy or cerebral commissurotomy. In these
cases, as with patients who have been subjected to "psychiatric
surgery," it is reasonable to suggest that the "pre-treatment"
psychological organization may have been grossly abnormal.


28
efforts in this area have been hampered by the lack of adequate
operational definitions (for such phenomena as consciousness, cogni
tion, and thought) which distinguish the hither mental processes
of humans from the brain functions of lower forms.
Cerebral dominance for consciousness has been investigated
using the Wada carotid amobarbital test, a procedure developed to
localize language functions prior to neurosurgery. The Wada test
involves injecting sodium amobarbital into one common carotid
artery and results in the anesthetization of only the cerebral hemi
sphere on the side of the injection. Terzian (1964) reported an
absolute and immediate arrest of any communication, both verbal and
nonverbal, in the first thirty to sixty seconds after the injec
tion of the drug into the carotid artery of the dominant side
which he interpreted as a transient loss of consciousness.
Serafetinides and his co-workers reported similar results and
noted that the phenomenon rarely occurred following barbiturization
of the non-dominant hemisphere (Serafetinides, Driver & Hoare,
1964, 1965a, 1965b). They concluded that unconsciousness, and by
implication consciousness, is in general linked with the function
of the hemisphere dominant for speech (Serafetinides et al., 1965a).
Rosadini and Rossi (1967) attempted to replicate these findings
using more strictly operationalized definitions of consciousness.
In one group (48 cases) the criteria consisted of an "analysis of
the capacity of the patient to keep in contact with the examiner
through verbalizations or movements, to react to noxious stimuli


90
involve an interaction between NE and the indolamine serotonin
(5-HT). Histological researchers have identified six major mono
amine systems in the rat brain (Fxe, 1965; Fuxe, Hamberger &
Hokfelt, 1968).
The niqro-striatal DA system originates in cell bodies in the
substancia nigra whose axons extend through the lateral hypothalamus
to terminate on cells in the caudate nucleus. This tract is known
to regulate the activity of the ancient extrapyramidal motor
control system.
The meso-limbic DA system originates in the ventral tegmental
area of the pons and projects mainly to the septal area and related
nuclei in the limbic forebrain. Dysfunction in this system is
assumed by many to be the source of schizophrenic disorders.
The meso-cortical DA system is less well defined but appears
to include projections from the ventrotegmental to the frontal
cortex and from the substancia nigra to the anterior cingulate
cortex (see Meltzer, 1979).
The ventral NE system originates in the reticular formation
(medulla and pons) and ascends through the median forebrain bundle
(MFB) to terminate on cells throughout the hypothalamus and amygdala.
Stein, Wise and Berger (1972) suggested that this system primarily
regulates motivational activities.
The dorsal NE system arises in the locus ceruleus in the pons
and may supply up to 70% of the NE in primate brains (Redmond,
1979). The axons in this system ascend through the MFB and give
off branches to the hypothalamus, the hippocampus and amygdala,


87
level of the temporal cortex. This match would take the form of a
finished "gestalt" in the experiential system and a formal "concept"
in the verbal system. With the appearance of such a match the
temporal component would terminate the search process and relay the
product to the cdntralateral system. Thus, the recognition of
a gestalt might initiate the search for a related verbal concept or,
conversely, a verbal formulation may trigger a scan for pertinent
experiential associations. The description of these two complementary
systems seems to provide adequate explanation for the qualitatively
different types of memory which were described by Breuer:.and Freud
and by Rapaport, respectively as noted at the beginning of this
section.
Substantial support for the model described above may be found
in a situation where hippocampal activity is induced from within
the limbic system (rather than from the neocortex, as was the case
in Penfield's experiments). Such is the case when amygdala stimula
tion produces after-discharges in the hippocampus. Halgren (1981)
reviewed the effects of amygdala stimulation in conscious humans and
noted that such stimulations sometimes produces "complex formed
hallucinations, sometimes complete scenes as in a dream or vivid
recollection and sometimes more vague, apparently similar to an
intruding thought . and illusions of familiarity (deja vu)"
(p. 345). Halgren suggested that it is the activation of distant
normal tissue subsequent to amygdala stimulation which produces the
resulting mental phenomena. He cites evidence that


6
"unit for receiving, analyzing and storing information," operating
in the post-central (sensory) areas of the brain; and the "unit
for programming, regulation and verification of activity," operating
in the frontal lobes (Luria, 1973a, ch. 2).
Luria's concepts represent a major advance in the understanding
of the fundamental operating characteristics of brain systems.
Although his formulations are too basic to be of much use to
the applied psychologist, he has established a format and a methodol
ogy which will be followed here. The present investigation will
focus on identifying and describing the interactions of the functional
brain systems which satisfy the requirements of the evolutionary
imperatives outlined above. Evidence suggests that the substrate
for these systems will be found in those anatomical areas for which
Luria acknowledged he had inadequate data for his own analyses: the
medio-basal zones of the cortex and the right hemisphere of the brain.


45
by recruiting psychological operations in the post-central cortex.
The frontal lobe may accomplish this through its influence on the
nonspecific thalamic nuclei which, in turn, control the phasic
activation of specific cortical systems.
It is important to note that the two biochemically mediated
subsystems which control the mobilization processes are duplicated
in both halves of the brain. Unilateral prefrontal lobe lesions
have been found to produce deficits which resemble those seen after
damage to post-central lesions on the same side: right frontal
lesions have been associated with disturbances of emotion and spatial
abilities while left-sided injuries lead to disorders of speech and
thought (Zangwill, 1966; Benton, 1968; Luria, 1973a).. Processes
which disrupt the biochemical balance between the two control
mechanisms have far-reaching psychological consequences which will
be reviewed in a later section.
Motivation: Emotion and Affect
The greatest risk involved in giving up instinct-based responding
in favor of self-determined behavior is the possibility that the
individual might fail to respond appropriately in survival-related
situations. The evolution of the species could not have occurred
if this problem had not been solved. The forces which motivate
adaptive behavior must, by definition, be the single most powerful
influence on the personality structure. The evidence indicates that
the source of these forces lies in the limbic system. It appears
that separate cortical mechanisms mediate their internal experience
and external expression.


Mammillary bodies and mammillothalamic tract..80
Cingulate cortex 81
Discussion 83
Interfaces and Interactions of the Monitoring,
Motivating, and Mobilization Systems 89
The Biochemistry of Emotion, Motivation,
and Learning 92
Emotion, Amygdala Circuits and Memory 95
Discussion 103
Biochemical and Electrophysiological Aspects
of Cortical Mobilization Processes 104
The Functional Elements of the Personality
Structure 108
The Problem-Solving/Response-Generatinq
System 109
The Memory System 110
The Motivating System 112
The Mobilization System 114
Learning and Memory: Animal Studies 115
A Functional Meta-system 120
Lateralized Mobilization Processes 125
Asymmetrical reaction time to laterally
presented stimuli 125
Altered GSR following unilateral brain
injury 126
Asyrmietrical biochemical and electro-
physiological processes 127
Bilateral Interaction in Emotion and Cognition..128
Mental Health and Psychopatholgy 136
The Psychopathological Correlates of
Unilateral Temporal Lobe Epilepsy 139
Discussion: Hyper- and Hypo-dominance
Spectrum Disorders 141
Schizophrenia and the Affective Disorders 143
Anxiety Disorders, Obsessive-Compulsive
Illness, and Paranoia 145
Sociopathy and Hysteria 147
III METHOD 151
Subjects 152
Instruments 153
Procedure 154
Hypotheses 155
IV RESULTS 156
Left versus Right Hemisphere Cognitive
Functioning Between Groups 156
Overall Performance on the Tests Sensitive to
Right versus Left Hemisphere Cognitive
Functioning 157
v


34
It appears, then, that the left hemisphere extracts meaning from
the relationship of individual parts to each other, while the
right gathers meaning from the pattern of the whole.
Bogan (1969b) proposed that the right "mind" utilized a
different mode of thought which he characterized as aopositional to
denote the ability to appose, or compare, information. He contrasted
this with the propositional mode utilized by the left hemisphere. A
number of investigators have distinguished similar dichotomies of
information processing style. Luria (1973a) spoke of narrative
versus relational processes. Gal in (1974) suggested that the right
hemisphere solved problems through a process of multiple converging
determinants as opposed to a left hemispheric style which utilized
a single causal chain. Sechenov (quoted by Luria, 1973a) postulated
that the human brain utilizes two forms of integrative activity:
organization into simultaneous and primarily spatial groups, and
into temporally organized successive series. This is consistent with
Spearman's conclusion that intelligence comprises two components: the
eduction of correlates used in analogical reasoning and the eduction
of relations, the basis of abstract reasoning (see McFie & Piercy,
1952). Campbell (1974) noted that "abstraction means escaDe from
the present . what distinguishes man from animals is the length of
time through which his consciousness extends (p. 335). Finally,
Bogan (1969b) observed that the most important distinction between
the left and right hemispheric modes might be "the extent to which
the linear concept of time participates in the ordering of
thought" (p. 160).


APPENDIX B
INFORMED CONSENT STATEMENT
Project Title: Functional Brain Systems and Personality Dynamics
You are being asked to participate in a research project
which was designed to find out whether certain patterns of per
ceiving and processing information are related to specific emo
tional or nervous problems. You were selected because the types
of problems you have reported are similar to the ones being
investigated. If successful, the study could lead to more effec
tive ways of helping people with those problems.
Participation in this research project will take about 15 minutes
of your time. No monetary compensation will be awarded. No identi
fying information will be recorded so it will be impossible to
report your results to anyone and you will not be recontacted.
There are no risks or discomforts involved. You will be given a
true-false questionnaire by your therapist or case manager; all
the researcher will ask you to do is to identify different types
of visual patterns and answer a series of nonpersonal questions.
You may withdraw your consent and discontinue at any time without
prejudice. If you have further questions, you may contact
David Lindquist at Mental Health Services, Inc., 374-5690.
166


36
construct such a consistent frame of reference the right hemisphere
is bound to the immediate context with only the influences of the
physiological status of the organism (and the left hemisphere) to
guide its processes. Complex motivations, therefore, cannot exist
in the right hemisphere. Likewise, so-called "pictorial thinking,"
if temporally ordered and goal directed, must be organized by the
left hemisphere. Gal in (1974) suggested that the context bound,
egocentric and impulsive nature of right hemisphere congition
resembled Freud's notion of primary process thinking. Higher mental
processes in the right hemisphere almost certainly qualify as
cognitions (i.e., a way of knowing) and may account for the phenomenon
of intuition (knowledge without awareness of the process by which
it was gained). However, the term "thought" seems misleading and
"information processing" might be preferable. As noted above, there
is no direct evidence that mental events occurring in the right
hemisphere are directly experienced in the conscious left hemisphere;
one is left to ponder the question of whether a tree falling in the
right brain would make a sound if the left wasn't listening.
The restrictions outlined above are in no way inconsistent
with the demonstrated role of the human right hemisphere in the
analysis of emotional communications and the modulation of affective
expression (e.g., Ross & Mesulam, 1979). The brains of lower
forms (and, apparently, the right brain in humans) are primarily
concerned with neuronal signals which represent the survival needs
of the organism within the immediate environment. Interaction with


24
There is less agreement about the third major language
disorder, known as anomic or amnesic aphasia, which results from
more posterior lesions located in the angular gyrus in the parieto
occipital area. Part of the confusion may stem from the fact that
its main symptom, the loss of the ability to name objects, is
common to all aphasic disorders. However, the anomic aphasia
syndrome is distinguished by the fact that the naming disorder is
accompanied by relatively intact comprehension of written and spoken
language and normal spontaneous speech. The ability to read and
to repeat are also spared in anomic aphasia.
The anomic aphasic has no difficulty using words in their
appropriate context but cannot find the word in isolation of
context; he is unable to divorce himself from the immediate
situation. The anomic aphasic cannot produce the name of objects
on demand even though he knows what they are. When an object is
designated the patient is unable to produce its name, and conversely,
given a name, the patient is not certain what it refers to.
Other language difficulties are evident. The patient's
spontaneous speech seems to be either too detailed or too general.
Thinking is very concrete; the patient will interpret proverbs
literally. The patient is aware of the diabilities and will often
develop strategies to compensate for them.
Lanugage, Symbolism, and Meaning
The theory of the functional organization of language develooed
by Wernicke in 1885 is still generally accepted today. According
to this model the underlying structure of an utterance arises in


79
mention any attempt at quantification of behavior. They simply
stated that "so far, in none of the 12 surviving cases there has
been [sic] any unfavorable neurological or psychiatric sequela"
(p. 177). It seems possible that these authors may have missed
subtle symptoms in their apparently superficial evaluation.
Wilder Penfield (quoted in Sweet et al, 1959) underscored the
fact that "patients who have [bilateral hippocampal lesions] do
not forget their skills. Two of them were able to carry out most
complicated skills learner previous--glove cutting and engineering
drawing" (p. 81). Unfortunately, the only behavioral measure
reported by Cairnes and Mosberg (1951) involved a return to work.
These authors noted that some of their pateints (who had incurred
fornix damage in the course of surgery to remove colloid cysts
of the third ventricle) showed initial confusion, loss of memorizing,
and amnesia for the period surrounding the operation, but: "after
operation all [but one of their nine surving cases] returned to
work, and . showed no disturbance of emotion or intelligence"
(p. 564). Thus
Four of the five young women . are doing normal
housework; three have borne children. The other
young woman is in regular work as a clerk, and is
free from complaints. . Two older women . .
are also doing their housework [although one has a
'slight impairment of memory']. ... Of the two men,
one is working regularly as a doIicemen, (p. 568)
The extent of the lesions in these patients is unclear. The authors
report only that "each had . partial or complete division of
the anterior columns of the fornix" (p. 564). (It seems possible
that these surgical fornicotomies spared the precommissural fornix.)


29
and to describe at the end of the examination what happened during
the examination itself" (p. 103). In a second group the criteria
for consciousness consisted of "a simple stimulus-resoonse test"
in which "the patients were instructed to work a switch held in
the hand ipsilateral to the intracarotid injection [i.e., the
hand controlled by the unanesthetized hemisphere] any time they
heard a given sound or saw a flash of light" (p. 103). Behavioral
and clinical events indicating unconsciousness were required to
last more than one minute in order to "permit their safe detection."
They found that 47 of their 69 cases did not meet their criteria
for unconsciousness and the 22 cases which did occurred in roughly
the same percentage following left and right injections; aphasia
occurred in 16 cases (15 left, one right); only five of the 21
cases (three left, two right) evaluated with the stimulus-response
test failed to operate the switch held in the hand controlled by
the unanesthetized hemisphere in response to the signals used.
Although their results were complicated by existing neuropathological
and cerebrovascular abnormalities, these authors concluded that
"the existence of a cerebral dominance for consciousness is not
supported" (p. 111). The usefulness of this study appears to
be severely limited by the criteria used to evaluate consciousness
in the unanesthetized hemisphere: a large number of animals of
various species have demonstrated the ability to "work a switch"
in response to visual and auditory stimuli and to respond to noxious
stimuli, but these lower forms are not considered conscious in the
same sense that humans are. The authors were looking for "the


175
Heath, R. G., & Galbraith, G. C. Analysis of Sodium Amytal Effects
on Frontal Lobe-Subcortical Interrelations. Int. J, Neurol,,
1965, 3-4, 348-357.
Henschen, S. E. On the Function of the Right Hemisphere of the Brain
in Relation to the Left in Speech, Music, and Calculation.
Brain, 1926, 49, 110-123.
Herzog, A. G., & Van Hoesen, G. W. Brain Research, 1976, 115, 57.
Hielman, K. M., Scholes, R., & Watson, R. T. Auditory Affective
Agnosia: Disturbed Comprehension of Affective Speech. Journal
of Neurology, Neurosurgery, and Psychiatry, 1975, 38^, 69-72.
Hielman, K. M., Schwartz, H., & Watson, R. T. Hypoarousal in
Patients with the Neglect Syndrome. Neurology, 1978, 28,
229-232.
Hielman, K. M., & Valenstein, E. Frontal Lobe Neglect. Neurology,
1972, 22, 660-664.
Hielman, K. M., & Van Den Able, T. Right Hemisphere Dominance for
Mediating Cerebral Activation. Neuropsychologia, 1977, 17,
315-321.
Hillix, W. A., & Marx, M. H. (Eds.). Systems and Theories in
Psychology: A Reader. New York: West Publishing Co.,
1974.
Isaacson, R. L. The Limbic System. New York: Plenum Press, 1974.
Isaacson, R. L., Douglas, R. J., Lubar, J. F., & Schmaltz, L. W.
A Primer of Physiological Psychology. New York: Harper &
Row, 1973.
Isaacson, R. L., Douglas, R. J., & Moore, R. Y, The Effect of
Radical Hippocampal Ablation on Acquisition of Avoidance
Response. Journal of Comparative and Physiological Psychology,
1961, 54, 625-628.
Isaacson, R. L., & Wickelgren, W. 0. Hippocampal Ablation and
Passive Avoidance. Science. 1962, 138. 1104-1106.
Ison, J. R., & Thomas, G. J. Anterior Limbic Cortex and Partial
Reinforcement Effects on Acquisition and Extinction of a
Runway Response in Rats. Journal of Comparative and Physiological
Psychology, 1969, 69, 17-24.
Iverson, S. D. Brain Dopamine Systems and Behavior. In L. I. Iverson,
S. D. Iverson, & S. H. Snyder (Eds.), Handbook of Psychopharmacology
Vol. 8: Drugs, Neurotransmitters and Behavior. New York:
Plenum Press, 1977.


65
activation of a distant, secondary ganglionic station (Penfield,
1975, ch. 7). In his later formulations then, Penfield referred
to those temporal structures as the "interpretive cortex" and postu
lated that his electrode had activated a final common pathway to
a secondary center which in turn produced the illusions of
comparative interpretation. Since the temporal lobe forms the
principal source of input into the hippocampus (which was known to
be related to memory), Penfield assumed that this was the secondary
center in question. He suggested that
The hippocampi seem to store keys-of-access to the
record of the stream of consciousness. With the
interpretive cortex, they make possible the scanning
and the recall of experiential memory. (Penfield,
1975, p. 36)
Penfield's finding that illusions of familiarity were associated
with activity of the temporal lobe in the right hemisphere is
complemented by Kimura's (1963) evidence that the right temporal
lobe appears to be more involved in the analysis of unfamiliar stimu
li. Kimura presented familiar and unfamiliar visual stimuli to the
right and left visual fields of patients with lesions of the right
or the left temporal lobe. The right temporal group was impaired in
the perception of the unfamiliar stimuli but not the familiar.
Kimura interpreted her results in terms of the verbal i dentifiability
of a stimulus:
It seems clear that a frequent (though not a necessary)
concomitant of familiarity in a perceptual sense is the
possibility of verbal identification. Where increased
familiarity with a stimulus object, or class of objects,
is associated with the repeated naming of the object,
the ability instantly to attach a name to it represents
an important step in the development of a concept. It


182
Pucetti, R. Bilateral Organization of Consciousness in Man. Annals
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Rapaport, D. Emotions and Memory. New York: Science Editions,
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Redmond, D. W. New and Old Evidence for the Involvement of a Brain
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Consciousness. Brain, 1967, 90, 101-112.
Ross, E. D., & Mesulam, M. M. Dominant Language Functions of the
Right Hemisphere: Prosody and Emotional Gesturing. Archives
of Neurology, 1979, 315, 144-148.
Rotter, J. Generalized Expectancies for Internal versus External
Control of Reinforcement. Psychological Monographs, General and
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Rotter, J. Some Problems and Misconceptions Related to the Construct
of Internal versus External Control of Reinforcement. Journal
of Consulting and Clinical Psychology, 1975, 43 (1), 56-67.
Rowe, S. N. Mental Changes Following the Removal of the Right
Cerebral Hemisphere for Brain Tumor. American Journal of
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Schafer, R. The Clinical Application of Psychological Tests.
New York! International University Press, 1948.


131
considered here would suggest that this is because the normal inter
actions between the hemispheres which involve motivation and arousal
take place at a subcortical level. Sperry pointed out that the
right hemisphere of these patients demonstrated "appropriate emotional
reactions" but that "apparently, only the emotional effect gets
across [to the left], as if the cognitive comDonents of the
process cannot be articulate through the brainstem . the
affective component gets across to the speaking hemisphere, but
not the more specific information" (1968, p. 732).
The proposed model specifies further that the right hemisphere
in humans specializes in analyzing and recording experiential data
and thus becomes the repository of the reinforcement history of the
individual. The emotional responsiveness and behavior of adults
whose right hemispheres were removed (because they had been invaded
by life-threatening, fast growing tumors) is markedly different
from the split-brain patient. Although normal intelligence
is generally retained, such people show a lessened capacity for
adaptability (Glees, 1961) and "suffered a loss in terms of
personality values . defects in judgement and . impairment
in insight, in emotional control, in initiative, and in perseverance"
(Gardner, Karnosh, McClure & Gardner, 1955, pp. 500-501). These
individuals, whose left hemisphere was deprived not only of sensory
and cognitive information from the right (as was the case with
cerebral commissurotomy patients), but also of that hemisphere's
motivational and mobilizing inputs, show a clear pattern of deficits:
their affect tends to be labile, inappropriate, and poorly modulated;


105
of a person before and after prefrontal lobotomy exemplify the
extremes on a continuum of "perceptual reactance" which can be
seen in the normal population. Petrie (1960; 1967) administered her
kinesthetic figura! aftereffects test to a variety of normal and
abnormal populations and demonstrated that, after repeated tactile
stimulation of the fingertios, some people consistently overestimate
(augment) the size of a standard stimulus while others consistently
underestimate (reduce) its size. Reducers tolerate pain well and
augmenters do not. Petrie argues that augmenters and reducers are
displaying basic differences in their methods of perceiving the
environment.
Several investigators studying visual evoked potentials to
light flashes have noted that some subjects showed a reduction in
the amplitude of these cortical responses as stimulus intensity
increased, a phenomenon which Kamphuisen and van Leeuwen (1968)
called "paradoxical diminution." Buchsbaum and Silverman (1968)
reported a significant correlation between flash visual evoked
responses and perfromance on Petrie's kinesthetic figura! after
effects test: augmenters showed an increasing cortical response as
stimulus intensity increased while reducers showed a leveling off
or decrease in the amplitude of their cortical responses. This
finding was replicated by Spilker and Callaway (1969) using
different stimuli and different electrode placement, von Knorring,
Espvall and Perris (1974) demonstrated empirically that reducers
classified by visual averaged evoked responses had significantly
higher pain thresholds and pain tolerance levels than augmenters.


39
Cortical Mobilization: Attention,
Arousal, and Activation
Consciousness and cognition become possible only when minimum
levels of cortical tone are attained. These tonic levels, reflected
in the desynchronized EEG pattern, permit sensory discriminations,
motor acts and other cognitive operations to take place. Once
activated, the cortex has the ability to make phasic modifications
of its level of activity and to voluntarily direct its attention.
The fundamental systems which govern cortical tone and attention,
however, are automatic and capable of overriding voluntary controls.
It is clear that these systems, with their ability to control the
level and content of consciousness, will exert a significant
influence on the personality structure.
The Reticular Activating System and Tonic Arousal
The mobilization of the cortex is accomplished by the brainstem
reticular formation (RF). This structure is a network of highly
interconnected neurons which adjusts its level of activation by
integrating input from the sensory pathways, limbic system struc
tures, and the neocortex. Impulses from the reticular formation
lower the activation thresholds of the neurons it projects to.
When the reticular formation is relaxed, cortical tone is lowered
and the organism sleeps (Moruzzi & Magoun, 1949).
The reticular activating system (RAS) regulates the state of
activation of the brain in two ways: the ascending reticular
activating system (ARAS) affects the brain diffusely and sets the
generalized (tonic) level of arousal; descending influences direct


58
was described independently in the 1880s by Karl Wernicke (whose
work with aphasia was reviewed earlier) and S. S. Korsakoff, a
Russian psychiatrist. Wernicke focused on the acute stage of the
illness during which the patient is usually depressed, fearful and
anxious; often paranoid; and always severely confused and disoriented
(Wernicke's encephalopathy). Patients who survive this acute stage
become stabilized in the phase known as Korsakoff's psychosis.
This chronic state is characterized by severe memory disorders and
profound changes in the patients personality, motivation, and affect.
Against a background of retained intellectual skills and intact
remote memory, the victim of Korsakoff's psychosis suffers a retro
grade amnesia for periods of up to several years before the onset
of the illness, and an almost total inability to recall any new
information once his or her attention is distracted from it. Conse
quently, these patients live virtually in the immediate present and
are always disoriented as to time, place and situation. These
patients are, at best, only vaguely aware of their inability to learn
new material. They often produce confabulations to cover gaps in
their recollection and fuse or combine ("reduplicate") experiences
from different periods in their lives (Gardner, 1975). In both
cases they believe that their statements reflect reality. In most
cases the patient's affect is blunted, although some patients have
exhibited chronic euphoria (e.g., Remy, 1942). They show reduced
spontaneity and initiative and a "lack of desire for alcohol, sex,
and other traditional reinforcers"'(Gardner, 1975, p. 11188).


BIOGRAPHICAL SKETCH
David Lindquist was born on November 22, 1944, at Mitchell Air
Force Base on Long Island, New York. He grew up in a half dozen
cities around the country during the forties and fifties and
attended as many educational institutions during that period.
During the sixties he dropped out of high school, served in the
Air Force, was married and eventually divorced, obtained a high school
equivalent certificate, and worked at a variety of jobs. He took
courses at community colleges on a part-time basis beginning in 1968,
and earned an A.A. degree in 1973.
He came to the University of Florida in 1974, majored in
psychology, and received a B.A. degree with high honors in 1976. He
worked in several inpatient and outpatient settings while continuing
his graduate study toward the doctoral degree in counseling psychology
and has developed a specialization in forensic psychology.
188


84
and finally, causes pertinent information concerning that stimulus
to be presented to conscious awareness. It appears that there are
two such systems in the human brain, each specialized to deal with a
different type of information. The first, lateralized to the right
hemisphere, performs the functions listed above with experiential
data. The second, operating in the left hemisphere, deals with
language and verbal concepts which may be based on gesta!ten that
are assembled and stored in the contralateral system. All of the
functions noted above are impaired, to a greater or lesser extent,
in various manifestations of the amnesic syndrome.
The evidence suggests that some learning does not take place in
amnesia victims, that is, memory traces are stored. However, the
amnesic subject has difficulty gaining access to those memory
traces when they are needed. More specifically, they are unable to
recognize and select the appropriate memory trace in a given situation
from the set of available traces. Access to the proper trace is
facilitated with adequate cueing. It appears that the function of
the hippocampal system is to assure the activation of appropriate
memory traces based on the requirements of the situation. In the
present formulation, significant (experientially based) memory traces
have been designated by the superordinate term "generalized expecta
tions." It appears that the hippocampal system is responsible
for the generalizing of these expectations from one situation to
another, similar, situation.
In lower forms, where survival is dependent on instincts, the
identification of a significant stimulus may culminate in the release


158
on an assessment of personality traits. Specifically, it was
hypothesized that the hyper-dominant group would score relatively
higher on MMPI clinical scales (6 and 7) which indicate symptoms
that are manifested intrapsychically and suggest a central tendency
to chronic and maladaptive overutilization of formal thought processes
while the hypo-dominant group would score relatively higher on MMPI
clinical scales (3 and 4) which indicate symptoms that are magnified
extrapsychically and suggest a chronic and maladaptive underutiliza
tion of formal thought processes. The mean ratio (3T + 4T/6T = 7T)
for the hyper-dominant group was x = 1.01 (SO = 0.12), while the
mean ratio for the hypo-dominant group was x = 1.12 (SD = 0.14),
indicating that the groups differed in the predicted directions.
A student's t-test for independent samples (Robson, 1973) performed
on these means revealed that the difference between groups was
highly significant (t = 2.75^ p < 0.005, one-tailed). It was
concluded that psychiatric inpatients and outpatients assigned to
the hyper-dominant spectrum disorder group scored relatively higher
on MMPI clinical scales which indicate symptoms that are manifested
intrapsychically and suggest a chronic and maladaptive overutiliza
tion of formal thought processes; their ratios differed significantly
from those of patients assigned to the hypo-dominant group, who
scored relatively higher on MMPI clinical scales which indicate
symptoms that are manifested extrapsychically and suggest a chronic
underutilization of formal thought processes, as predicted by the
proposed model.


154
"general denial of physical health . denial of psychological or
emotional problems and of discomfort in social situations. . .
They react to stress by developing physical symptoms . [and]
are not likely to report anxiety, tension or depression" (Graham,
1977, pp. 38-39). High scorers on Scale 4 (psychopathic deviate)
show "absence of satisfaction with life, family problems, delinquency,
sexual problems, and difficulties with authorities" (Graham, 1977,
p. 41). Scales 5 and 7 were selected as representing symptoms that
are characteristic of hypothesized hyDer-dominance spectrum
disorders. The high scorers on Scale 6 (paranoia) show paranoid
symptoms. Scale 7 (psychasthenia) is a good index of psychological
turmoil, including excessive doubts, anxiety, compulsions and
obsessions, unreasonable fears, and tension (Graham, 1977). Graham
concluded that the Mini-Mult is "useful for comoaring groups, particu
larly if they are psychiatric patients ..." (1977, p. 219).
Procedure
Demographic information and diagnostic data were taken from the
subjects' clinical records. No identifying information was recorded.
The Mini-Mult was administered by the subjects' primary therapist.
Subjects were tested by the author before or after their regularly
scheduled therapy session. Street Test, Similarities, Object Assembly,
and Information Subtests were administered in that order. Diagnos
tic data, demographic information and test responses were entered on
a subject data form. The WAIS-R subtests and the Mini-Mult were
scored by the author using standard scoring criteria (Wechsler, 1981;
Kincannon, 1968). The Street Test as originally standardized
(Street, 1931) includes a number of antiquated items and the


130
half-brain knew what the word was. In both instances,
the command kiss elicited an emotional reaction that
was detected by the verbal system of the left hemi
sphere, and the overt verbal response of the left
hemisphere was basically the same, regardless of
whether the word was presented to the right or left
half-brain. In other words, the verbal system of the
left hemisphere seemed able to accurately read the
emotional tone and direction of a word seen by the
right hemisphere alone. (Gazzaniga & Ledoux, 1977,
p. 151)
Thus, P.S.'s left hemisphere appears to have experienced a direction
ally specific emotion in the absence of a cognition, and the
required information must have been obtained from the right
hemisphere via the AC. In this case it is not possible to differen
tiate whether the response was made possible by information passed
between his amygdalae, or by cognitive information passed from the
right temporal lobe to the left, or both. The interpretation of
this phenomenon is further complicated by the fact that P.S.
possessed language in his right hemisphere. The finding demonstrates,
however, that the interaction of the hemispheres in emotion as
proposed in the present model is tenable. The likelihood of such
an interaction is supported by evidence concerning differences
in the emotional responsiveness of split-brain patients and adults
who have had their right hemispheres removed because of fast
growing tumors.
The most surprising initial observation concerning split-
brain subjects is still the most significant: there were no
readily observable changes in the intelligence, behavior, or
personalities of these patients following the disconnection of
their cerebral hemispheres (Sperry, 1968). The model being


64
unchanging ganglionic record of subjective experience" (Penfield,
1954, p. 67). Niesser suggested that "most of the cases described
by Penfield seem more like generic and repeated categories of events
rather than specific instances" (p. 168). (It will be noted that
Niesser's formulation is congruent with the notion of a "generalized
expectation," as defined earlier.)
Penfield's second mechanism was suggested by another category
of electrically induced phenomena which consisted of the "misrepre
sentation or altered interpretation of present experience" (Mullan &
Penfield, 1959, p. 269). Prominent among these were "illusions of
recognition" during which "present experience seemed familiar,
strange, altered, or unreal" (p. 270). These "illusions of compara
tive interpretation" were associated with stimulation of the temporal
cortex in the hemisphere that was minor for handedness and speech.
The authors believed that "in normal life, these are signals that
rise into consciousness, signals that depend on subconscious
comparison of past experience with the present" (p. 283).
In 1951 Penfield proposed that portions of the temporal lobes
be called "memory cortex" in the belief that his electrode had
activated a neuronal record which was stored there. He was obliged
to revise this theory in 1958 because of a new understanding of the
physiology of electrical brain stimulation. When an electrode passes
a current into the cerebral cortex, the current completely disrupts
the patient's normal use of that gray matter (e.g., stimulation
of the speech areas produces momentary aphasia). Therefore, any
positive responses are produced by axon-conduction and the functional


This dissertation was submitted to the Graduate Faculty of the
Department of Psychology in the College of Liberal Arts and Sciences
and to the Graduate School, and was accepted as partial fulfillment
of the requirements for the degree Doctor of Philosophy.
May 1935
Dean for Graduate Studies and
Research


109
psychological operations to determine the form of that response.
It was postulated that these mechanisms would be arranged in
functional systems (as defined by Luria, 1973a) and that these
central organizing factors would form the infrastructure of
personality. The brain mechanisms which appear to satisfy these
requirements will be outlined below. It will be proposed that
these systems form the basic functional elements of the personality
structure. Supporting evidence from animal studies of learning
and memory will be presented at the end of this section.
The Problem-Solving /Response-rGeneratinq System
It is evident that the psychological operations which generate
considered responses in a given situation take place in the neural
tissue of the association areas in the post-central cortex of the
left cerebral hemisphere. These mechanisms normally have access
to, and employ the products of, mechanisms lateralized in homologous
areas of the right hemisphere. Both of these areas solve problems
through a process of categorization, but they arrive at their
solutions in different ways. The right hemisphere is specialized
to perceive overall patterns in input; it solves problems through a
convergence of information, synthesizing basic units into meaningful
gestalts. The left hemisphere shares these spatial abilities to some
extent, but they have been superseded by more complex skills. The
dominant hemisphere has been genetically prepared to analyze
information along a temporal dimension and this ability allows
mental activity to escape from the bounds of the inmediate spatial
context. This freedom, in turn, permits the mental manipulation


5
Neurological theories of mental function center on clinical
observations of patients with localized brain lesions. The history
of neurological thought has revolved around the question of how these
data are to be interpreted. For many years higher mental functions
were treated as discrete "faculties." The results were inconsistent
and of little value. The failures of these "narrow localizers" led
to theories which attempted to account for mental functions on the
basis of the "mass action" of the brain. Where earlier models were
too specific, these theories proved too general to be useful.
In the 1920s Goldstein broke with the tradition of attempting to
infer functions directly from deficits and proposed instead an
"analysis of basic disturbances." This approach led finally to
Luria's conceptualization of mental activities as the product of the
interaction of complex functional systems. In Luria's formulation
a mental function is the result of contributions from a number of
concertedly working zones. Therefore, that function may be destroyed,
or disturbed differently, by lesions in different locations. Luria
(1973a) described the characteristics of a functional system:
The presence of a constant (invariant) task, performed
by variable (variative) mechanisms, bringing the process
to a constant (invariant) result, is one of the basic
features distinguishing the work of every "functional
system." The second distinguishing feature is the
complex composition of the "functional system," which
always includes a series of afferent (adjusting) and
efferent (effector) impulses. (Luria, 1973a, p. 28)
Luria outlined three principal functional units in the brain:
the "units for regulating tone and waking and mental states,"
v t
centered on the reticular activating system in the brainstem; the


96
1981) and related to survival needs, including defensive and aggressive
behaviors, sexual activity, and feeding (isaacson, 1974). In lower
forms these processes might depend on a simplified (instinctive)
form of memory in which stimulus and response are yoked (Pribram &
McGuinness, 1975). In addition to mediating emotional states the
arnygdala is involved in the analysis of reinforcement contingencies.
Amygdala lesions have been shown to produce impaired recognition of
stimuli associated with rewards (Weiskrantz, 1956; Schwartzbaum,
Thompson & Kelli cut, 1964; Jones & Mishkin, 1972) and inability to
respond appropriately to changes in the magnitude of rewards
(Schwartzbaum. .1960).
The more recent discovery of strong anatomical interconnections
between the amygdala and the neocrotex (Aggleton, Burtin & Passingham,
1980; Herzog & Van Hoesen, 1976; Turner, Mishkin & Knapp, 1980)
allows for the involvement of this structure in higher levels of
mental processing and memory. Kessner (1981) suggested that long
term memory "consists of a set of bundle of traces, each representing
some attribute or feature of a learning experience." He postulated
that "the amygdala mediates the encoding, storage, and retrieval
of specific emotional attributes often associated with reinforcement
contingencies in specific situations" (p. 332). In a test of this
hypothesis Kessner and Conner (1974) showed that five seconds of
bilateral subseizures level electrical stimulation of the amygdala
(which disrupts the normal functioning of that structure) following
footshock in a passive-avoidance paradigm resulted in amnesia for
that training experience. Implanted and unoperated controls without


25
Wernicke's area. It is then transferred via large fibre bundles
(the arcuate fasciculus) to Broca's area where it evokes a
detailed program for vocalization which is governed by the rules
of grammar and syntax. The program developed in Broca's area, the
linear scheme of the sentence, is supplied to the adjacent face area
of the motor cortex which in turn drives the muscles which produce
the vocalization. Thus, the content of speech originates in
Wernicke's area and finds its form in Broca's area.
Wernicke's area is also essential for the comprehension of
language. Auditory stimuli are relayed from the Organ of Corti
to the primary auditory projection areas in Heschell's gyrus in the
left temporal lobe. At this point, as with the other sensory projec
tion areas, the information is somatotopically organized and retains
its modal specificity; to be understood it must be transferred
to the secondary auditory association area (Wernicke's area)
where the somatotopical organization is converted into a functional
organization (Luria, 1973a). Here, the fundamental phonemic
characteristics of language are isolated and identified. Processing
by Wernicke's area is essential for both the encoding and decoding
of meaning. An intact Wernicke's area is also essential for the
expression and comprehension of meaning through symbolic gesture
and pantomime (Goodglass & Kaplan, 1973; Gionotti & Lemmo, 1976).
It is interesting to note in this regard that "illusions of
interpretation" emerge in consciousness after electrical stimulation
of the temporal lobe in the right hemisphere but not after stimula
tion of any other brain area (Mullan & Penfield, 1959).


83
(in runway problems and bar-press alternation: Barker & Thomas, 1965,
1966; Barker, 1967), and the failure to exhibit behaviors which were
indicative of opium addiction (Marques, 1971). The last, especially,
is supportive of the hypothesis voiced by some authors that cingulate-
lesioned animals are unable to anticipate the emotional consequences
of their behavior for both rewards and punishments (Glass, Ison,
& Thomas, 1969; Isaacson, 1974).
Anterior cingulectomy has been termed the psychosurgical
"operation of choice" for the treatment of severe obsessional and
anxiety disorders (Lewin, 1961; Whitty, 1966). Whitty noted that
one of the long-term effects of cingulectomy was "relative neglect
of the impact of external events." Finally, Pechtel, McAvoy,
Levitt, Kling & Massermann, (1958) concluded that lesions of the
cingulate gyrus in humans resulted in, among other things, "amnesia
for previous learning" and "impairment of new learning skills."
Discussion
The failure to transfer learning between hemispheres seen in
cerebral commissurotomy preparations (animal and human) indicates that
the storage of memories is a neocortical function (Geshwind, 1965).
Generalized memory disorders, on the other hand, are only produced
by bilateral damage to the diencephalic structures of the hippocampal
system. It is evident that these structures in the circuit of
Papez form part of a functional system which monitors the environ
ment, matches incoming stimuli with representations of the organisms
previous experience with similar stimulus configurations, activates
the organism in the presence of potentially significant stimuli,


LIST OF FIGURES
FIGURES
1 Cytoarchitectural map of the lateral and medial
surfaces of the human cerebral cortex, with numbers
representing the areas of Brodman 10
2 Partially schematized representation of the
limbic system 13
3 The position of Papez's circuit within a
larger cortical circuit 86
4 Schematic representation of the proposed model of
the physiological substrate of personality Ill


170
Brodie, B. B., & Shore, P. A. A Concept for a Role of Serotonin and
Norepinephrine as Chemical Mediators of the Brain. Annals of the
New York Academy of Science, 1957, 66^, 631-642.
Broekkamp, C. L., & Lloyd, K. G. The Role of the Amygdala on the
Action of Psychotropic Drugs. In Y. Ben-Ari (Ed.), Inserm
Symposium No. 20, The Amygdaloid Complex. New York: Elsevier/
North-Holland Biomedical Press, 1981.
Bruder, G. E., & Yozawitz, A. Central Auditory Processing and
Laterality in Psychiatric Patients. In J. Fruzelier & P.
Flor-Henry (Eds.), Hemisphere Asymmetries of Function and
Psychopathology. Amsterdam: Elsevier, 1979.
Bruell, J. H., & Albee, G. W. Higher Intellectual Functions in a
Patient with Hemispherectony for Tumors. Journal of Consulting
Psychology, 1962, 26 (1), 90-98.
Buchsbaum, M. W., & Silverman, J. Stimulus Intensity Control and
the Cortical Evoked Response. Psychosomatic Medicine, 1968,
30, 12-22.
Butters, N., & Cermack, L. S. The Role of Cognitive Factors in the
Memory Disorders of Alcoholic Patients with the Korsakoff
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Cairnes, H., & Mosberg, W. H. Colloid Cysts of the Third Ventricle.
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Campbell, B. G. Human Evolution. Chicaqo: Aldine Publishing Co.,
1974.
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Coger, R. W. Alcoholism: Averaged Visual Evoked Response Amplitude-
Intensity Slope and Symmetry in Withdrawal. Biological
Psychiatry, 1976, U, 435-443.
Cohen, G. Hemispheric Differences in Serial Versus Parallel
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Cohen, J. Cerebral Psychophysiology: The Contingent Negative
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1974.


137
According to the model normal, adaptive functioning requires that
the problem-solving/response-generating system receive adequate
information from the monitoring, mobilizing and motivating systems.
Mental health would require, and result from, the integrated
functioning of these basic personality elements. Conscious left
hemisphere processes must be mobilized appropriately by emotional
experience and "channelized" by generalized expectations which are
congruent with reality (cf., Kelly, 1963). Such integrated
functioning was implicit in Rogers (1963) description of the
"fully functioning person":
This person would be open to his experience . .
every stimulus, whether originating within the
organism or in the environment, would be freely
relayed through the nervous system without being
distorted by a defensive mechanism . the
self and personality would emerge from experience,
rather than experience being translated or twisted
to fit a preconceived self-structure . since he
would be open to his experience . this person
would find his organism a trustworthy means of
arriving at the most satisfying behavior in each
existential situation. (1963, pp. 18-20)
Any process that interfered with the operation of an individual
element, or prevented adequate interaction among them, would diminish
the adaptability of the organism. The system is vulnerable to
interference at several levels.
The model suggests that, at a cognitive level, generalized
expectations (GEs) are the most potent influence on behavior.
These significant experiential memories are assembled and stored
in the right half of the neocortex and are accessed by the hippocampal
system lateralized in that hemisphere. Such GEs are here defined as


TABLE OF CONTENTS
ACKNOWLEDGEMENTS iii
LIST OF FIGURES vii
ABSTRACT viii
CHAPTER
I INTRODUCTION 1
II THE PHYSIOLOGICAL SUBSTRATE OF PERSONALITY:
A SELECTIVE REVIEW OF THE LITERATURE 7
Review of Basic Brain Anatoniy and Organization 8
Cortical Mechanisms 8
Subcortical Systems 12
The Interpretation of Neurology Literature 17
Human Consciousness 18
Right versus Left 18
Language and the Left Hemisphere 20
Aphasia: Anatoniy and Syndromes 21
Language, Symbolism, and Meaning 24
Human Consciousness, Self-Awareness,
and Thought 26
Discussion 35
Cortical Mobilization: Attention, Arousal,
and Activation 39
The Reticular Activating System and
Tonic Arousal 39
Phasic Control Systems: The Frontal Lobes
and Thalamus 41
Discussion 44
Motivation: Emotion and Affect 45
Amygdala Circuits and the Prefrontal Lobes 46
Affective Expression 48
Discussion 50
Memory Functions 51
Human Amnesia Syndromes 57
Memory and the Neocortex 63
Discussion 68
The Limbic System, RAS, and Memory 70
Papez Circuit and Memory 74
Fornix 75
iv


48
tactlessness" (see Lewin, 1961). Faust (1966) noted that such
patients resemble psychopaths in that they are unable to profit
from experience and are in constant conflict with their environ
ment and the law. Zangwill (1966) pointed out that the tactlessness
conrnon in frontal lobe patients does not result from a loss of
knowledge of social conventions, but from the failure to regulate
behavior in accordance with those standards.
Disconnecting the orbito-frontal cortex from the aniygdala
(orbital undercutting) has been reported to be the most effective
psychosurgical operation for relieving the symptoms of anxiety
and depression (Elithorn et al., 1958; Lewin, 1961; Levinson &
Meyer, 1965). Elithorn et al. (1958) noted that this procedure
increased reactions of "a hysterical type" and is contraindicated
for those conditions. It is interesting to note that drugs that
reduce anxiety and produce euphoria (e.g., the barbiturates) have
been found to exert an uncoupling effect between the frontal lobes
and the limbic system (Heath & Galbraith, 1965).
Affective Expression
The right hemisphere plays an essential role in both the
comprehension of emotional Communications and the expression of
affect. Patients with right hemisphere lesions showed impaired
recall of stories with emotional content versus neutral stories
(Wechsler, 1973). Hielman, Scholes and Watson (1975) demonstrated
that judgements of the emotional mood of a speaker (sad, happy,
angry, indifferent) made by patients with right temporoparietal
lesions were significantly impaired relative to patients with left
sided lesions and controls. This finding was replicated by Tucker,


APPENDIX C
REVISED SCORING CRITERIA FOR THE STREET (1931) GESTALT COMPLETION TEST.
Credit any response that indicates that the following has been seen:
Item 1: a dog
Item 2: a boat or ship
Item 3: a cat
Item 4: a stove
Item 5: a baby
Item 6: a table
Item 7: a man in uniform with a rifle or a fisherman
with a fishing pole*
Item 8: a horse
Item 9: a rabbit
Item 10: a locomotive
Item 11: a boy on a tricycle
Item 12: a man's face
Item 13: a man kneeling*
*Altered from original in Street (1931)
167


CHAPTER II
THE PHYSIOLOGICAL SUBSTRATE OF PERSONALITY:
A SELECTIVE REVIEW OF THE LITERATURE
In the following sections the neurological and physiological
psychology literature pertaining to the functional/anatomical
organization of personality will be reviewed. The data will be
related to psychological factors and any conclusions will be noted
in discussions at the end of each section. The first section is a
review of basic brain anatomy and organization. The second section
will examine the physiological substrata of consciousness and conclude
that human consciousness, characterized by self-awareness, is a
manifestation of processes which occur in the left hemisphere of
the brain. The third section will trace the systems involved in
cortical activation and note the existence of two mechanisms within
each hemisphere which have opposite effects on the form of cognitive
processes. The fourth section will examine the role of the aniygdala
and frontal lobes in the subjective experience of emotion and of
the right hemisphere in the expression of affect. The fifth section
will develop the basis for a theory of memory function and the role
of memory systems in organizing affective, arousal, and cognitive
processes. The sixth section will focus on the interactions of the
emotion, arousal, and memory systems and examine the role of bio
chemically mediated systems in coordinating these processes. In
the seventh section a model of four functional systems which
7


2
based on the former have been criticized as untestable and therefore
unscientific (Eysenck, 1970). The latter movement lost impetus with
the discovery by Olds and Milner (1954) of "pleasure" centers in the
brain. This revelation undermined the basic assumption of the learning
theorists that behavior could be explained simply by defining the
rules governing stimulus-response relationships.
Successful scientific theories are built on paradigms that
describe the fundamental properties and mechanics of their subject.
The fundamental units of personality are networks of neurons in the
brain. Sigmund Freud (1948) attempted to relate mental structures
to anatomical locations but was forced to abandon his effort because
the neurology of the time was not adequate. Instead he and subsequent
theorists were forced to base their models on suppositions about
the products of the personality processes. As noted above, the
results have been less than satisfactory. The science of neurology
has made significant progress in the interim and a large amount of
useful information has accumulated. These data have been virtually
ignored by the discipline of psychology. The integration of neuro
logical data and psychological theory may provide a basis for a useful
paradigm for the psychotherapist. The present work is intended as
a step toward such an integration.
The purpose of this study is to develop and test a heuristic
model of personality function, based on an understanding of its
physiological substrate, with the ultimate goal of imoroving the
effectiveness of psychotherapeutic interventions. Such a model
should identify the basic elements and processes of the personality


174
Gloor, P. Telencephalic Influences Upon the Hypothalamus. In W. S.
Fields (Ed.), Hypothalamic-Hypophysical Interrelationships.
Springfield, Illinois: C. C. Thomas, 1956.
Gloor, P. Etudes Electrographiques de Certaine Connexions RhinenceDh-
aliques. In Ph. Alajouanine (Ed.), Physioloqie et Pathologie
du Rhinencephale. Paris, France: Masson, 1961.
Goodglass, H., & Kaplan, E. Disturbance of Gesture and Pantomime in
Aphasia. Brain, 1973, 86, 703-720.
Gottfries, C. G., Perris, C., & Roos, B. E. Visual Averaged Evoked
Responses (AER) and Mono Amine Metabolites in Cerebrospinal
Fluid (CSF). Acta Psychiatric Scandinavica, 1974, 255, 135-142.
Graham, J. R. The MMPT: A Practical Guide. New York: Oxford
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Gray, H. Anatomy, Descriptive and Surgical. New York: Bounty Books,
1977.
Green J. F., & Arduini, A. Hippocampal Electrical Activity in Arousal.
Journal of Neurophysiology, 1954, T7, 533-557.
Griffith, H., & Davidson, M. Long Term Changes in Intellect and
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surgery, and Psychiatry, 1966, 29, 571-576.
Grossman, S. P. Behavioral Functions of the Septum: A Re-analysis.
In J. F. DeFrance (Ed.), The Septal Nuclei. New York: Plenum
Press, 1976.
Gruner, J. E. Sur la Pathologie des Encephalopathies Alcoliques.
Revue Neurol., 1956, £4, 682-689.
Gruzelier, J. H., & Hammond, N. Schizophrenia: A Dominant Temporal-
Limbic Disorder. Research Communications in Psychology,
Psychiatry, and Behavior, 1976, 1, 33-72.
Halgren, E. The Amygdala Contribution to Emotion and Memory: Current
Studies in Humans. In Y. Ben-Ari (Ed.), Inserm Symposium No. 20,
The Amygdaloid Complex. New York: Elsevier/North-Holland Bio
medical Press, 1981.
Haliday, A. M., Davidson, K., & Brown, M. W. Comparison of Effects on
Depression and Memory of Bilateral ECT and Unilateral ECT to
the Dominant and Nondominant Hemisphere. British Journal of
Psychiatry, 1968, U4, 997-1012.
Hanson, N. R. Patterns of Discovery. New York: Cambridge University
Press, 1965.


10
Figure 1. Cytoarchitectural map of the lateral and medial
surfaces of the human cerebral cortex, with numbers repre
senting the areas of Brodman. (Redrawn from Noback & Demerest,
1972).


88
Amygdala stimulation seldom evokes a mental phenomenon
unless it also evokes an after-discharge . most
amygdala stimulations, even if they evoke an after
discharge (AD), do not evoke any reported mental phenome
non . thus, there is no direct connection between
amygdala activity and hallucinatory experiences. . .
Simultaneous recordings from multiple brain areas
indicate that amygdala ADs seldom remain localized.
Initial spread is to the ipsilaterla hippocampus and
hippocampal gyrus. AD may remain confined to these
structures, in which case no mental phenomenon is
necessarily evoked. Further spread is to the ipsi-
lateral limbic cortex (orbital, insular and cingular)
and diencephalon (especially the anterior nucleus of
the thalamus, but also to the centre median, pul vinar
and dorsomedial nuclei). ADs seldom spread to the
neocortex, which may however desynchronize." (pp. 396-
397, emphasis added)
These findings are congruent with the present formulation, which
would interpret the appearance of "complex formed sensory hallucina
tions" following the stimulation described above as the result of
hippocampal output evoking experiential memory traces in the posterior
association cortex by way of the anterior thalamus and cingulate
cortex. The appearance of identical "experiential hallucinations"
following temporal lobe stimulation and limbic system ADs supports
Penfield's contention that final common pathways in the temporal
lobe may produce activity in the hippocampal system which utlimately
results in the appearance of mental phenomena.
The hippocampal system is directly involved in the mechanics
of learning. Hippocampal activity seems to assure that the organism
attends to novel stimuli and sets the conditions which permit changes
to be encoded into the neural models which form the centeral represen
tations of those stimuli. Pribram and McGuinness (1975) suggested
that such changes in neural represenations "may be conceived as


63
that his patients were deficient in making associations among new
ideas and in connecting past and present experience.
The suggestion of bilevel processes in memory noted above are
particularly interesting given that language and conscious awareness
have been associated with temporal lobe structures of the left
hemisphere, and Penfield's (1975) report that electrically induced
"illusions of recognition" were elicited only by stimulation of
temporal structures of the right hemisphere.
Memory and the Neocortex
As noted earlier, Penfield's experiments with electrical stimula
tion of the cortex in conscious patients led him to postulate the
existence of two separate memory systems: a "mechanism of recall"
and a "mechanism of interpretation." The existence of the former
was suggested by the fact that, following stimulations of the exposed
cortex, some of his patients reported vivid auditory and/or visual
experiences ("flashbacks"); it seemed to the patients as if they
were reliving prior experiences in their lives, although they retained
their awareness of the operating room environment. Since many of
these sensory sequences were trivial, yet perceived as familiar,
Penfield concluded that his electrode was tapping a "continuous
record of conscious experience." Although widespread areas of the
cortex were exposed and explored, these "experiential responses
come only from the temporal lobe, never from any other part of the
brain" (Penfield, 1975, p. 31).
Niesser (1967) presented persuasive arguments refuting Penfield's
claim that "nothing is lost . the brain of every man contains an


74
around the peak frequency narrowed during discrimination; and the
location of the peak shifted in discrimination reversal procedures.
In summary, the hippocampus appears to be able to monitor
incoming stimuli and match them against (cortical) neuronal models
which represent the past experience of the organism with related
stimuli. In addition to facilitating appropriate access to these
memory traces hippocampal activities have been directly associated
with the triggering of cortical processes which permit further
analysis of a stimulus, emotional and behavioral responses as
warranted, and/or alterations in the neuronal model itself (i.e.,
learning).
Papez Circuit and Memory
The hippocampus forms part of a continuous pathway within the
limbic system which Papez (1937) believed to be the substrate of
emotion. Papez was aware that destruction or stimulation of limbic
structures produced major alterations in emotional behavior and
believed that emotional expression depended on the integrative action
of the hypothalamus. He was also convinced that subjective emotional
experience required the participation of the cerebral cortex.
Papez outlined an anatomical circuit through which he thought
emotion might arise in either of those two centers. Thus
Incitations of cortical origin would first pass to
the hippocampal formation and then down by way of
the fornix to the mammillary body. From this they
would pass upward through the mammillothalamic tract
... to the anterior nuclei of the thalamus and thence
by the medial thalamocortical radiation (in the
cingulum) to the cortex of the gyrus cinguli . .
Radiations of the emotive process from the gyrus
cinguli to other regions in the cerebral cortex


141
may lead to action in which the patient "takes the law
into his own hands." Sensing emotional importance in
even the smallest acts, he performs these ritualisti-
cally and repetitively. Since details bear the
imprimatur of affective significance, they may be
mentioned in lengthy, circumstantial speech or
writing, (o. 465)
Discussion: Hyper- and Hypo-dominance Spectrum Disorders
The proposed functional meta-system model specifies that, in
survival situations, emotion and arousal are normally triggered
by the right ("monitoring") system and are resolved by the left
("problem-solving") system. The right memory system also provides
appropriate context information and generalized expectations (via
the cerebral commissures) to assist in this process. When the
verbal left hemisphere initiates limbic system processes directly
(e.g., by triggering its own amygdala) there would be a tendency
toward a vicious circle: the increased attentional/emotional
activity would exacerbate the focus on the initiating cognitive
theme (rather than eliciting an objective response to environmental
requirements). In normal circumstances this reverberating unilateral
process might account for mood (the maintenance of affective tone
in the absence of external stimuli). If abnormally exacerbated,
the process might produce the rigidified cognitive phenomena
associated with various pathological syndromes (e.g., anxiety,
depression, paranoia, obsessions). In addition, the overactive
verbal system would systematically initiate searches in the contra
lateral memory system for experiential data which were congruent
with its abnormal cognitive state. Finally, memories encoded with
"artifically" induced emotional significance would, by definition,
tend to deviate from reality.


32
His right hemisphere has a sense of self, for it knows
the name it collectively shares with the left. It has
feelings, for it can describe its mood. It has a sense
of who it likes and what it likes, for it can name its
favorite people and its favorite hobby. The right
hemisphere in P.S. also has a sense of the future, for
it knows what day tomorrow is. Furthermore, it has
goals and aspirations for the future, for it can name
its occupational choice. . The fact that this mute
half-brain could generate personal answers to ambiguous
and subjective questions demonstrates that in P.S. the
right hemisphere has its own independent response-
priority determining mechanisms, which is to say, its
own volitional control system, (pp. 143-145)
P.S. is the only split-brain patient with advanced language skills
in his right hemisphere and the only patient to demonstrate double
consciousness. Ledoux, Wilson and Gazzaniga (1977) stressed the
fact that "in all other patients, where linguistic sophistication
is lacking in the right hemisphere, so too is the evidence for
consciousness" (p. 420).
The capacity for speech and conceptual thought is clearly innate
in homo sapiens; only the symbols themselves must be learned
(Campbell, 1974). Recent evidence has clarified the anatomical
substrate of this genetically transmitted specialization. The
development of language capabilities in the human species is
correlated with the anatomical expansion and interconnection of
the association areas in the left hemisphere (Campbell, 1974). The
posterior area of the planum temporale, which forms a part of the
secondary auditory cortex (Wernicke's area) is significantly larger
on the left side (Geshwind & Levitsky, 1968. The enlargement of
this area can be explained in terms of its distinctive cellular
organization (Galaburda, LeMay, Kemper & Geshwind, 1978) and the


4
1. The physiological substrate of personality evolved to
assure the emission of adaptive behavior by the organism. Survival
pressures shaped an organization that was able to respond effectively
to significant stimuli and produce behavior that enhanced the
organism's well-being.
2. The evolution of this substrate proceeded from an organiza
tion based on "hard-wired" stimulus-response instincts to an organiza
tion which allowed increased latitude in behavior in order to take
advantage of the organism's developing problem-solving abilities.
3. In order to permit self-determination of behavior and,
at the same time, to insure survival, new mechanisms were required
to assure that the organism would (a) attend to significant
stimuli; (b) be motivated to respond effectively to those stimuli;
(c) accomplish the mobilization of the necessary psychological
resources to determine the form of that response (referred to
hereafter as the Monitoring, Motivating, and Mobilization systems).
4. Although these systems interact, they must be functionally
separated to the extent that they do not interfere with each other's
normal operation. Similar functions might be carried out in other
brain areas, but the automatic activation of these systems will
assure that they dominate responding to stimuli which relate to the
survival and well-being of the organism.
5. As these mechanisms are essential for the survival of
the individual and species they will form the central organizing
processes of personality; personality dynamics will center on their
operation.


CHAPTER V
DISCUSSION
This study was an attempt to affirm or disaffirm certain central
consequents of the proposed model of personality function and psycho
pathology. With this type of design a failure to support the hypotheses
is unequivocal: the theory cannot be true. If the consequences
are clearly verified there is some indication that the theory may be
true. All of the hypotheses in this experiment were suDported at
high levels of statistical significance. It was concluded that the
proposed theoretical model of personality function and psychopathology
is tenable and may provide useful operational definitions for the
applied psychologist.
The value of an experimental test of a theory is determined,
in large part, by the strength of the chain of reasoning between the
hypotheses and the data. In this study that connection is relatively
straightforward. Based on the functional meta-system model it was
postulated that the manifestations of personaltiy dysfunction (i.e.,
psychopathology) will be determined by the properties of the
cerebral hemisphere with greater access to the limbic system. It was
proposed, therefore, that there are two major subtypes of psycho
pathology. Since the functional impact of disturbances within the
meta-system is thought to be experienced in the left cerebral hemi
sphere (which alone possesses the requisites for self-awareness
160


129
nuzzle the man. Prior to the completion of the neocortical
disconnection the hemisphere that was blind but capable of feeling
fear received adequate information from the sighted side to activate
the intact amygdala and motivate the organism appropriately. However,
when this blind but fearful hemisphere was isolated the animal's
behavior was determined by the sighted but fearless side. (The
animal responded with normal fear and aggression when touched on
the trunk or limb.) The result was, in effect, a functional
bilateral amygdalectomy with regard to visual stimuli (Puccetti,
1977), demonstrating that without adequate cortical input to the
amygdala the organism cannot behave in an adaptive manner.
When a visual stimulus is presented tachistoscopically to
one hemisphere (by way of the left or right visual half-fields)
in a split-brain patient, the opposite hemisphere is "blind"
with respect to that stimulus. Case P.S. differs from earlier
split-brain patients in that his anterior commissure (which connects
the amygdalae and temporal lobes) was left intact. The combination
of these factors led to an important discovery in the course of
a standard experiment with P.S.:
On the verbal command test . where a word was
lateralized to the right hemisphere and P.S. was
instructed to perform the action described by the
word, his reaction to the word kiss proved revealing.
Although the left hemisphere of this adolescent boy
did not see the word, immediately after kiss was
exposed to the mute right hemisphere, the left
blurted out, "Hey, no way, no way. You've got to
be kidding." When asked what it was that he was
not going to do, he was unable to tell us. Later,
we presented kiss to the left hemisphere and a
similar response occurred: "No way. I'm not going
to kiss you guys." However this time the speaking


72
Early investigators were puzzled by the fact that cortical
activation (EEG desynchronization) was accompanied by synchronous
slow-wave activity (4-8 Hz theta rhythms) in the hippocampus
(e.g., Green & Arduini, 1954). The most constant behavioral corre
lation of hippocampal theta activity in animals of different species
is orienting towards, and attending to, stimuli in the environment
(Isaacson, 1974). Cortical activation, such as that seen in the
orienting response, is accomplished by the brainstem RAS in conjunction
with the nonspecific thalamic nuclei. The hippocampus appears to
regulate the process of involuntary attention by performing switching
functions through a mutually inhibitory relationship with the reticu
lar formation (Smithies, 1966). The Soviet neuropsycholgist Vinogradova
provided important insights into the mechanics of this process.
By observing unit activity with microelectrodes Vinogradova
(1970) determined that all of the neurons in the hippocampus monitor
incoming stimuli, habituating to repetition and dishabituating to
any change in the stimulus configuration. Such responsiveness
requires constant matching of the stimulus with a related neuronal
model (Sokolov, 1963). That these models exist in the cortex, and
not in the hippocampus itself, is established by evidence that the
quality of sensory information is almost completely erased in
hippocampal neurons (Gloor, 1961).
Vinogradova distinguished two types of neurons in the hippo
campus: A-neurons (30-40%) which are activated by a stimulus and
I-neurons (60%) which are inhibited. She went on to proDOse a
mechanism whereby the hippocampus is able to modulate the processes
of attention and learning:


33
incomplete development of this cellular architecture has been
related to language dysfunction (see Geshwind, 1979).
In her exhaustive study of hundreds of brain injured war
veterans, Semmes (1967) discovered that elementary sensory and
motor capacities were focally represented in the left hemisphere and
diffusely represented in the right. She proposed that this difference
indicates the mechanism of hemispheric specialization: focal
organization favoring fine control and the integration of similar
units (e.g., manual skills and speech) and diffuse organization
favoring multimodal coordination (e.g., the various spatial
abilities).
Gazzaniga and Ledoux (1977) observed that nearly every demonstra
tion of a right hemisphere advantage in split-brain patients has
involved manipulo-spatial activities and concluded that
[This advantage] exists so long as manipulative activities
are involved in either the stimulus perceptions or the
response production. . The probable neural substrate of
these manipulo-spatial acts involves the inferior parietal
lobule of the right hemisphere in humans. In the left
hemisphere, however, linguistic functions occupy the
inferior parietal lobule. . The superior performance
of the right hemisphere of split-brain patients on such
tasks does not reflect the evolutionary specialization
of the right hemisphere, but instead represents the
price paid by the left hemisphere in acquiring language.
. . Our view is not that the right hemisphere is
specialized in some unique way in man. Rather, it
continues to do what it does elsewhere in the phyla.
(pp. 420-421, emphasis added)
Campbell (1974) noted that "spatial relationships involving
depth and distance may appear to be predominately spatial concepts,
but they are not of space but about space; of themselves they are
spaceless and concerned with pattern rather than place" (p. 337).


172
Eysenck, H. J. Personality Organization. In W. A. Hillix &
M. H. Marx (Eds.), Systems Theories in Psychology: A Reader.
New York: West Publishing Co., 1974.
Faust, C. Different Psychological Consequences Due to Superior
Frontal and Orbitofrontal Lesions. Int., J. Neurol., 1966,
3-4, 410-419.
Fawcett, J. Biochemical and Neuropharmacological Research in the
Affective Disorders. In E. J. Anthony & T. Benedek (Eds.),
Depression and Human Existence. Boston: Little, Brown & Co.,
1975.
Flor-Henry, P. Psychosis, Neurosis and Epilepsy. British Journal
of Psychiatry, 1974, _124, 144.
Flor-Henry, P. On Certain Aspects of the Localization of the Cerebral
Systems Regulating and Determining Emotion. Biological
Psychiatry, 1979, 14 (4), 677-698.
Freud, S. The Unconscious: Collected Papers. Vols. 4 and 5.
London: Hogarth Press, 1948.
Fuxe, K. Evidence for the Existence of Mono Amine Neurons in the
Central Nervous System: IV. Distribution of Mono Amine
Nerve Terminals in the Central Nervous System. Acta Physiologica
Scandinavica, 1965, 64, 37-85.
Fuxe, K., Hamberger, B., & Hokfelt, T. Distribution of Noradrenaline
Nerve Terminals in Cortical Areas of the Rat. Brain Research,
1968, 8, 125-131.
Gaffen, D. Loss of Recognition Memory in Rats with Lesion of the
Fornix. Neuropsychologia, 1972, 10, 327-341.
Galaburda, A. M., LeMay, M., Kemper, T. L., & Geshwind, N. Right-
Left Asymmetries in the Brain. Science, 1978, 199, 852-856.
Gal in, D. Implications for Psychiatry of Left and Right Cerebral
Specialization. Arch. Gen. Psychiatry, 1974, 31, 572-583.
Galin, D. Lateral Specialization and Psychiatric Issues: Speculations
and Development and the Evolution of Consciousness. Annals of
the New York Academy of Science, 1977, 299, 397-411.
Galin, D. Diamond, R., & Braff, D. Lateralization of Conversion
Symptoms: More Frequent on the Left. American Journal of
Psychiatry, 1977, 134, 578-530.
Gardner, H. The Shattered Mind. New York: Knopf, 1975.


92
cognitive patterns consistent with the goals of orienting at the
neocortical level. An opposite pattern is evident when the catechola
mines predominate. The specific systems involved in these functions
will be described in the following sections.
The Biochemistry of Emotion, Motivation, and Learning
The fundamental prerequisite for adaptive behavior is a system
to sort environmental inputs and pair them with appropriate responses.
The more primitive life forms must rely on the experience of their
species, accrued over evolutionary history and trasmitted genetically,
to accomplish these tasks. The adaptability of such forms is
strictly limited by the repertoire of sensory discriminations and
motor response sequences they are programmed to perform. Kety (1972)
noted that adaptability would be greatly enhanced if the organism
could learn to determine and preserve those sensory, evaluation, and
response patterns which, in the individual's own experience, pro
vided the ultimate survival advantage. He suggested that neuronal
and neurochemical mechanisms which utilized constant reference to
a small number of inborn values or states would allow the nervous
system to become increasingly elaborate and effective in performing
these functions. Kety postulated that only three such inborn states
should be required: arousal, pain, and pleasure. He proposed
that these mechanisms would interact to produce an emotional state,
consisting of increased arousal and the activation of particular
circuits, with a resulting drive to approach or avoid the confronting
stimulus. Arousal would accompany both pleasure and pain but might
occur in the absence of either in response to novel stimuli or to


73
The hippocampus exerts a tonic inhibitory influence upon
the reticular formation, blocking activatory processes
through the tonic discharge of its I-neurons when
novelty is absent and registration [a change in the
neuronal model] is not needed. But when a stimulus which
is not registered in the memory system appears, this
inhibitory control is blocked (I-neurons become silent],
arousal occurs, and the process of registration starts.
(1970, p. 114)
Vinogradova's hypothesis is supported by observations of electrical
activity in the brains of animals in classical conditioning paradigms.
Theta rhythms (associated with activity of Vinogradova's A-neurons)
are found in the early stages of learning but disappear when the
response has become well established (Isaacson, 1974). During
conditioning the time course of the theta rhythm and the orienting
response are matched. As the latter is replaced by the stabilized
conditioned response, theta dies out (I-neurons become active again)
and the hippocampus resumes its inhibitory controlover the RAS,
thus ending the orienting response and thereby allowing the fully
developed conditioned response to materialize (Smithies, 1966, p. 90).
The hippocampal-cortical interaction was apparent in an
analysis of the characteristics of hippocampal theta rhythms in
situations requiring different types of cortical information
processing. Bremner (1970) investigated the effects of orienting,
simple conditioning, discrimination, and discrimination reversal
tasks on various parameters of the theta rhythm using the habituated
organism (rat and man) as a baseline. He found that theta power
(amount of energy) increased in the presence of stimuli which elicited
orienting and arousal and decreased in the interval preceding a
response in the conditioning situation; the range of energy distribution


Copyright 1985
by
David Lindquist


148
"the most outstanding single feature of the sociopath's WAIS test
profile is his systematic high scores on the performance as compared
to the verbal part of the scale" (Wechsler, 1958, p. 176). The
same pattern was found to be characteristic of hysterics (Schafer,
1948). These relationships indicate a relative predominance of
right hemisphere processes in these disorders. The correlation
between low levels of left hemisphere functioning and low anxiety
is also striking. The PIQ greater than VIQ WAIS pattern has been
associated with "acting out" tendencies (Ogden, 1967). Silverman,
Buchsbaum, and Stierlin (1973) found that acting out adolescents
showed evoked potential augmenting. Augmenting has also been
associated with high scores on a sensation seeking scale (Zuckerman,
1974) and with alcoholism (Knorring, 1979; Coger, 1976), traits
which are common in sociopathy and hysteria (see Ball's, 1978).
Smokier and Shevrin (1979) used lateral eye movements as an
index of relative hemispheric activation in obsessive-compulsive
and hysterical personality style subjects (designated by modified
standard tests)] They found significantly more left-looking among
the hysterical.subjects indicating increased right hemisphere acti
vation in this disorder. They suggested that these personality
styles may be related to "predominant use of one or the other
hemisphere" (p. 952).
The dissociative reactions seen in hysterical neurosis (amnesia,
depersonalization, fugue, multiple personality) are indicative of a
failure to encode or access memories in the verbal system (cf.,
Gazzaniga, 1977). The finding that hysterical conversion symptoms


102
that monkeys with cingulate damage treat their companions like
inanimate objects, which he interpreted as a "loss of social con
sciousness."
4. fiiesser (1967) noted that the content of the complex
hallucinatory experiences reported by Penfield's subjects a'fter
electrical stimulation of the right temporal cortex was not
surprising: "What is surprising is only the vividness of the
imagery" (Ntesser, 1967, p. 169). Whitty (1966) reported that,
for one to three days postoperatively, patints who had undergone
anterior cingulectomy experienced "increased vividness of thoughts
and images so that a subjective difficulty is experienced in
distinguishing between thoughts and exterior happenings" (p. 404).
5. The cingulate cortex receives projections from the brain
stem ascending 5-HT and dorsal NE systems and has the highest
density of NE terminals in the cortex (Fuxe et al., 1968).
6. Redmond and his colleagues (see Redmond, 1979) reported
that monkeys with bilateral locus ceruleus lesions showed an
"absence of emotional responses to threat" (p. 158). Encephalitis
lethargica is a disease which affects the brainstem and the peri
aqueductal gray area (which contains the locus ceruleus) in
particular. Penfield (1975) noted that postencephalic patients
often developed an obsessive-compulsive syndrome: such patients
were "preoccupied with compulsive thoughts, compulsive utterances,
and compulsive behaviors to an extent that often caused severe
disability" (p. 98).


101
The identification of emotionally significant stimuli in
phylogenetically lower forms may result in the release of species-
specific responses at the level of the hypothalamus, but in higher
forms the neocortex is more involved in motivating behavior. It is
assumed here that the more recently developed systems which provide
the substrate for higher mental processes must be programmed in
some way to insure survival. It appears that biochemically mediated
mechanisms may have evolved in parallel over the more ancient control
circuits to perform this function. Stein and Wise (1971) suggested
that the dorsal NE system was involved in the regulation of cognitive
processes. Several lines of evidence, taken together, strongly
suggest that the cingulate cortex and NE participate in the integra
tion (encoding and decoding) of emotional experiences with memory
to facilitate adaptive functioning at the highest levels:
1. Based on the review in the previous section it was hypothe
sized that the cingulate cortex is the final station in a hippocampal
memory circuit which performs memory indexing functions.
2. The cingulate cortex is interposed anatomically between the
prefrontal lobe and the inferior parietal lobule (I PL) and is
thus in a position to coordinate the actions of these two highest
centers of information processing.
3. Animals with cingulate lesions are unable to acguire
avoidance learning (see previous section). Ison and Thomas (1969)
concluded from their studies that cingulate lesions "dissociated
conditioned emotional goal responses from instrumental performance"
(p. 17). Ward (quoted in Isaacson et al., 1971, p. 232) reported


Key to the schematic representation of the proposed mode) of the physiological substrate of personality. Most connections are reciprocal; arrows
indicate pathways emphasized in the text. Heavy black lines indicate memory indexing information from the hippocampal Monitoring System. Dashed
lines indicate emotion/reinforcement information from the amygdalar Motivating System. Dotted lines indicate activating information from the
reticular formation Mobilizing System. AMYG--amygdala; aniyg--contralateral amygdala; DM--dorsomedial thalamic nucleus; Pre-FR CTX--prefrontal
cortex; MOTmotor cortex; BG basal ganglia; TEMP CTX--temporal lobe cortex; AIS--autonomic nervous system; HT--hypothalamus; SEPT--septal area;
RF--reticular formation; ILTN--intralaminar thalamic nuclei (includes all nonspecific thalamic nuclei; Post-ASSH CTX--postcentral association cortex
(includes secondary association areas and inferior parietal lobule); SENS--primary projection sensory cortex; HPC--hippocampus; hpc--contralateral
hippocampus; MB--manmi 11 ary body; ANT--anterior thalamic nuclei; ClNG--cingulate gyrus; CC--eorpus callosum; AC--anterior comnissure; ufuncinate
fasciculus; vaf--ventral aipygdalofugal fibers; st--stria terminalis; sm--stria medullaris; sm-hab--stria medullaris via habenlua; mfb--medial
forebrafn bundle; fnx--fornix; MIT--mamillothalamic tract.
Figure 4. Schematic representation of the proposed model of the physiological substrate of
personality.


Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy
FUNCTIONAL BRAIN SYSTEMS AND PERSONALITY DYNAMICS
By
David Lindquist
May 1985
Chairman: Dr. Harry Grater
Major Department: Psychology
A heuristic model of the physiological substrate of personality
structure was developed from a review of recent neurological and physio
logical psychology literature. The formulation of the model was based
on assumptions that the phylogenetic transition from instinctive
responding to self-determined behavior required the evolution of auto
matic neural mechanisms to insure that the organism would monitor the
environment for significant stimuli, be motivated to respond in
the presence of those stimuli, and mobilize the appropriate psycho
logical operations to determine the form of that response. Functional
elements of these basic mechanisms were identified and a functional
meta-system was outlined which would organize the elements and
optimize the utilization of lateralized cognitive processes in the
interest of assuring the emission of adaptive behavior.
The proposed model suggests that certain psychodiagnostic entities
might be classified as hyper- and hypo-dominance spectrum disorders
vi 1 i


38
style of the right hemisphere is not suited to solving problems or
making decisions, it is uniquely qualified to perceive the quality of
significance (as defined by the individual's experience) in complex
environmental stimuli. The evidence suggests that the human right
hemisphere attends to the overall pattern of stimuli, searches out
("apposes") associations which are correlated with important
stimulus configurations and collates them into percepts that have
meaning for the organism. Its associational processes are unen
cumbered by rules of logic and its perceptions uninfluenced by
expectations.
The thrust of Bogan's (1969b)"dual mind" thesis was a reaction
against the traditional concept of hemispheric dominance which
relegated the right hemisphere to the role of an "automaton" or
reserve organ (e.g., Henschen, 1926; Strong & Elwyn, 1943). A
basic assumption in the present work, however, is that evolutionary
pressures reguired in the development of an automatic environment
monitoring system in order to permit the transition from instinctual
to self-determined behavior. It appears that evolution solved
this problem by taking advantage of the fact that the human central
nervous system contains two relatively autonomous brains which could
be yoked together by the limbic system. Within this configuration
the left hemisphere may be seen as a problem-solving and response
generating system and the right hemisphere might be said to
function as the repository and librarian of the individual's
reinforcement history.


136
Springer and Deutch (1981) suggested that the left hemisphere
"assumes that what it sees encompasses everything there is" (p. 176).
This interpretation is supported by the fact that the speaking left
hemisphere of split-brain subjects is unaware of having lost half of
its sensory fields (Gazzaniga, 1970). Similarly, the subjects
in the experiment reported by Dimond et al. (1976), who viewed
films presented to only one hemisphere, were not aware that they
were blind in one visual field. In addition, they experienced
the stimuli as "centered" when in reality it was presented from
the left or right of the midline. Finally, it is worth noting in
this context that Korsakoff's psychosis (in which the patient is
unable to gain access to recent memory traces) is commonly referred
to as the "amnestic-confabulatory syndrome." The Korsakoff patient's
speaking left hemisphere produces false accounts when memory fails
to provide the facts. This material "is presented without awareness
of its distortions or of its inappropriateness, and ... is moti
vated in no other way than factual information based on genuine
data" (Talland, 1965, p. 50). In short, when deprived of expected
memory data, the left hemisphere believes that what it thinks is
true. Considerable evidence of confabulation by the isolated left
hemisphere in split-brain patients was reported by Sperry (1974).
Mental Health and Psychopathology
The model developed in the preceding sections defines the basic
functional elements of the personality structure. It remains to
describe the ways in which these interact to produce mental health
and psychopathology.


125
emotional experience by programming cognitive operations in the left
post-central association areas to analyze the situation (aided by
context information and generalized expectations obtained from the
right experiential memory system in steps two and three) and formulate
a behavioral response; (6) the behavioral response alters the
situation, allowing the termination of motivating and mobilizing
systems activity.
Evidence concerning the proposed model will be organized and
presented below in terms of its relevance to the arousal, emotional,
and cognitive components of the meta-system.
Lateralized Mobilization Processes
The functional meta-system model suggests that the initial
identification or categorizaton of an environmental stimulus will
normally take place in the right hemisphere which will then trigger
the mobilization of the left half of the brain to deal with that
stimulus. Substantial support for this postulate is provided by
data from investigations utilizing different experimental approaches.
Asymmetrical reaction time to laterally presented stimuli.
Hielman and Van Den Able (1977) tested the effect of neutral warning
signals (WS) presented to the left or right visual half-field
on right hand reaction time (RT) to a centrally presented light.
They found that a WS presented to the right hemisphere reduced
RTs of the right hand more than a WS presented to the left hemisphere.
Bowers and Hielman (1976) examined between-hand RT differences
to a binaurally presented tone which was randomly preceded by a
verbal or nonverbal WS stimulus presented visually to both hemispheres


186
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University Press, 1970.
von Knorring, L., Espvall, M., & Perris, C. Averaged Evoked Potentials,
Pain Measures, and Personality Variables in Patients with
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Walter, W. G., Cooper, R., Aldridge, V. J., McCallum, W. G., &
Winter, A. L. Contingent Negative Variation: An Electric
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Watson, R. I. Psychology: A Prescriptive Science. American
Psychologist, 1967, 22, 435-443.
Wechsler, A. F. The Effect of Organic Brain Disease on Recall of
Emotionally Charged versus Neutral Narrative Test. Neurology,
1973, 23, 130-135.
Wechsler, D. The Measurement and Appraisal of Adult Intelligence,
4th Ed. Baltimore, MD: W i 11 i ans & Wilkins, 1958.
Wechsler, D. WAIS-R Manual. New York: The Psychological Corporation,
1981.
Weiskrantz, L. Behavioral Change Associated with Ablation of the
Amygdaloid Complex in Monkeys. Journal of Comparative and
Physiological Psychology, 1956, 49, 381-391.
Weiskrantz, L. A Comparison of Hippocampal Pathology in Man and
Other Animals. In CIBA Foundation Symposium 58 (new series),
Functions of the Septo-Hippocampal System. New York:
Elsevier Excerpta Medica/North-Holland, 1979.
Weiskrantz, L., & Warrington, E. K. Verbal Learning and Retention
by Amnesic Patients Using Partial Information. Psychoneurology
Science, 1970, 20, 210-211.


153
All subjects were selected by diagnosis and were asked to
participate by their primary therapist. Each subject read an
informed consent statement (Appendix B) before being tested.
Instruments
The dependent measures in this study were scores on the Street
(1931) Gestalt Completion Test; the WAIS-R Object Assembly, Similari
ties and Information Subtests (Wechsler, 1981); and the Mini-Mult, a
71 item abbreviated form of the MMPI (Kincannon, 1968). The Street
Test and the Object Assembly Subtest have both been shown to be
differentially sensitive to right hemisphere cognitive processes and
right hemisphere damage impairs performance on these tests (Bogan,
Dezure, Ten Houten & March, 1972; Direnzi & Sninnler, 1966; Ogden,
1967; Matarazzo, 1972; Rapaport, 1951; Black, 1974). The Similarities
and Information Subtests of the WAIS-R have been shown to be differ
entially sensitive to the left hemisphere cognitive processes and
left hemisphere damage impairs performance on these tests (Bogan
et al., 1972; Ogden, 1967; Matarazzo, 1972; Rapaport, 1951). Bogan
et al. (1972) demonstrated that patients with partial sectioning
to the cerebral commissure show higher scores on the Street Test
and higher Street/Similarity ratios than patients with complete
sectioning of the commissures. This was interpreted by the authors
as indicating that the Street Test normally required right hemi
sphere processing.
Scales 3 and 4 from the MMPI were selected as representing
symptoms that are characteristic of the hypothesized hypo-dominance
spectrum disorder. High scorers on Scale 3 (hysteria) show a


ACKNOWLEDGEMENTS
I am deeply grateful to Max and Ruth Lindquist for their patience
and the opportunities they gave me. My debt to my undergraduate
mentor, Dr. Carol Van Hartesveldt, who taught me about science and
let me make my own mistakes, is also gratefully acknowledged.
I want to thank my doctoral committee members, Doctors Grater,
Morgan, Schauble, Suchman and Ziller, who were also my most respected
graduate teachers. Special thanks are due to David Suchman, who
first encouraged me in this project, and to my chairman, Harry Grater,
who shepherded me through the crises with grace and allowed me to
keeD my dignity.
The help of Dr. Bill Froming in the design of this study, and of
Mssrs. Denny Gies, Bill Baxter and Ted Shaw (of the North Florida
Evaluation and Treatment Center) and of Ms. Janet Despard (of Mental
Health Services, Inc.) in facilitating its execution, is much
appreciated. I am especially grateful to Ms. Cheryl Shaw, who typed
the manuscript, and without whose friendship and organizational help
this project would not have been completed.
Warmest heartfelt gratitude goes to my dear friends Gabriel
Rodriguez, Marshall and Laura Knudson, and David Kurtzman, whose
love sustained me through these difficult years.
I can never properly express my thanks to Ruth Lindquist, to whom
this piece of work is lovingly dedicated.
iii


159
Age
The mean age in the hyper-dominant group was x = 42.65 years
(SD = 13.73 years). The mean age in the hypo-dominant group was
x = 30.45 years (SD = 12.14 years). A student's t-test performed
on these means revealed significant differences between groups
(t = 2.12^g, p < 0.05, two-tailed).


107
stimulus identification) while the reduction mode would facilitate
the sequential information processing and abstraction of detail
associated with problem-solving and language processes.
A novel stimulus (information plus uncertainty) "disinhibits
and gains access to large number of neurons indiscriminantly
throughout the neocortex" (Kety, 1972, p. 71). Pribram and
McGuinness (1975) reviewed evidence which indicates that the source
of the generalized cortical arousal associated with orienting
is to be found in the serotonergic nuclei of the brainstem. The
searching and sampling of the orienting process is terminated
when the stimulus is identified and registered in memory. At
this point uncertainty is resolved. If the situation registered
in the neuronal model of the identified stimulus has no built in
"demand" characteristics the organism may habituate to the stimulus
and proceed to ignore it. However, if the neuronal model generates
important expectations related to the stimulus, then cortical
activation systems which facilitate perceptual and motor readiness
to respond will be engaged. In contrast to the indiscriminate
cortical arousal associated with orienting and perceptual augmenta
tion, these activation systems produce a highly organized pattern
of cortical activity. Pribram and McGuinness (1975) reviewed
evidence which indicates that the mechanism and pathways involved
in controlling these activation systems are dopaminergic, and that
their operation is reflected in the phenomenon of the contingent
negative variation, or "expectancy wave."


based on the form of dysfunction within the meta-system. The ability
of the model to predict membership in diagnostic categories was
tested by assigning 42 adult psychiatric inpatient and outpatient
subjects to hyper-dominant and hypo-dominant groups by diagnosis,
according to the constructs of the model, and comparing performance
on instruments shown to be sensitive to right and left cerebral hemi
sphere dysfunction. The Street Gestalt Completion Test and the Object
Assembly, Similarities, and Information Subtests of the Wechsler Adult
Intelligence Scale-Revised were administered to each subject. An
abbreviated form of the Minnesota Multi phasic Personality Inventory
(MMPI) was used to compare symptomology between groups.
Significant between-group differences (p < 0.001) in the ratios
of test scores sensitive to right versus left cognitive functioning
were found in the predicted directions, while the groups did not differ
in overall performance on the instruments. Significant differences
(p < 0.005) in the ratios of selected MMPI clinical scales, in the
predicted direction, provided further support for the hypothesized
relationship between lateralized cognitive functioning, symptomology,
and diagnosis.
It was concluded that the proposed model orovides tenable and
potentially useful operational definitions of personality functions
and psychopathology. Results were discussed in terms of their impli
cations for psychotherapeutic interventions and additional methods
to test the validity of the model.
IX


146
activation system. (The association between NE-mediated mobilization
processes involving the locus ceruleus and cingulate cortex and
the "hyper-indexing" of memories in obsessive-compulsive illness
was noted in an earlier section.)
Amnhetamine increases the amount of dopamine available at
synapses. In animals, increasing amounts of dopamine activation
results in a decrease in the ranges of behaviors emitted and an
increase in the frequency of a few behaviors, leading to stereo
typed motor sequences at high doses (Iverson, 1977). In humans,
chrnoic amphetamine abuse has been reported to result in stereotyped
repetitive behaviors, increased attention to detail, compulsive
disassembly of objects, ruminative preoccupation with intellectual
ideation, and features of hypervigilance difficult to distinguish
from paranoid schizophrenia (Ellinwood, 1967).
Witkin (1965) developed the concept of a field-dependence-inde
pendence dimension in perception. In the field-dependent mode, per
ception is dominated by the overall organization of the field, the
parts of which are experienced as fused. In field-independent
perceiving, parts of the field are experienced as discrete from
organized background (cf., perceptual augmenting and reducing).
Field-dependence and independence were dramatically related to
right and left hemisphere functioning, respectively, in an experiment
utilizing unilateral ECT as the independent variable and rod-and-
frame test scores as the dependent variables. Twenty-four subjects
were administered the rod-and-frame test shortly after admission to
the clinic and were randomly assigned to receive right or left ECT


121
maturity until relatively late in ontogeny. The progression in
species from dependence on primitive perceptions through secondary
and tertiary integration is also evident in the development of brain
function in human individuals ('ontogeny recapitulates philogeny1).
The primary projection areas are operational at birth; secondary
association areas are programmed for function (via "learning") after
birth (Penfield, 1975); the tertiary integration areas do not become
fully operative until the seventh year of life (Luria, 1973a).
(This developmental sequence appears to parallel Piaget's descriptions
of sensory-motor, concrete operational, and formal operational
stages.) During ontogeny, the lower systems in the hierarchies become
subordinate to the higher levels: elementary perceptions are fit
into learned schemes and, in adults, are coded into logical systems
(Luria, 1973a). Inadequate development at any one level will affect
functioning at higher levels.
The development in humans of manual dexterity and, later, of
speech required the specialization of certain secondary and tertiary
association areas. A tendency emerged for these operations to be
organized in one hemisphere of the brain (Luria's law of the
progressive lateralization of functions). At this point the functions
of the hemispheres began to differ radically.
Both hemispheres possess advanced circuitry with the capacity
for high level information processing. However, they are relatively
isolated from each other and the left is genetically prepared to
deal with language. The primary projection areas in the two hemi
spheres have identical roles, but the secondary and tertiary


178
Lewin, W. W. Observations on Selective Leucotomy. Journal of
Neurology, Neurosurgery, and Psychiatry, 1961, 24_, 37.
Liss, P. Avoidance and Freezing Behavior Following Damage to the
Hippocampus or Fornix. Journal of Comparative and Physiological
Psychology, 1968, 66, 193-197.
Lubar, J. F., & Perachio, A. A. One-Way and Two-Way Learning and
Transfer of an Active Avoidance Response in Normal and Cingulec-
tomized Cats. Journal of Comparative and Physiological
Psychology, 1965, 6£, 46-52.
Lubar, J. F., Perachio, A. A., & Kavanagh, A. J. Deficits in Active
Avoidance Behavior Following Lesions of the Lateral and Postero
lateral Gyrus of the Cat. Journal of Comparative and
Physiological Psychology, 1966, 62,~~25~3-269.
Luria, A. R. The Working Brain. New York: Basic Books, 1973a.
Luria, A. R. The Frontal Lobes and the Regulation of Behavior,
In K. H. Pribram & A, R. Luria (Eds.), Psychophysiology of
the Frontal Lobes. New York: Academic Press, 1973b.
Luria, A. R., Homskata, E. D., Blinkov, S. M., & Critchley, M.
Impairment of Selectivity of Mental Processes in Association
with Lesions of the Frontal Lobe. Neuropsychologia, 1967,
5, 105-117.
Lynch, G., Rose, G., & Gall, C. Anatomical and Functional Aspects
of the Septo-Hippocampal Projections. In CIBA Foundation
Symposium 58 (New Series), Functions of the Septo-Hippocamal
System. New York: Elsevier ExcerDta Medica North-Holi and,
1977.
MacLean, P. D. Psychosomatic Disease and the "Visceral Brain":
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Psychosomatic Medicine, 1949, 11., 338-353.
MacLean, P. D. The Triune Brain, Emotion, and Scientific Bias.
In F. 0. Schmitt (Ed.), The Neurosciences, Second Study
Program. New York- Rockefeller University Press, 1970.
Mahut, H. A Selective Spatial Deficit in Monkeys After Transection
of the Fornix. Neuropsychologia, 1972, JJD, 65-74.
Mahut, H., & Zola, S. M. A Non-Modality Specific Impairment in
Spatial Learning After Fornix Lesions in Monkeys. Neuropsychologia,
1973, 11, 255-269.
Mandell, A. J. & Knapp, S. Asymmetry and Mood, Emergent Properties
of Serotonin Regulation. Archives of General Psychiatry. 1979.
36, 909-916. ^


116
However, when the situation requires that the animal select from
more than one possible hypothesis, as in successive or simultaneous
discrimination problems, hippocampal animals are impaired relative
to controls (Kimble, 1961, 1963). Similarly, naiv hippocampal
animals can learn to bar-press for rewards under continuous rein
forcement (CRF) conditions but are impaired when they are transferred
to a partial or intermittent schedule (e.g., the DRL). Schmaltz
and Isaacson (1970) demonstrated that it is the change in schedules
that is debilitating: Hippocampal animals trained from the outset
using only the DRL contingencies were not impaired on that task.
Winocur and Mills (1970) found that previous experience impaired
the performance of hippocampal animals only when the preceding task
was related in some way to the new problem; unrelated training
did not interfere. Thus, similarity between situations actually
hinders hippocampal animals rather than helping them. Isaacson
(1974) interpreted such results as the inappropriate transfer of
strategies or the fixation of predominant behavioral dispositions,
but the present formulation would stress the hippocampal animals'
inability to select between competing memory traces because they
do not receive memory indexing information ath the cortex.
Five experimental paradigms have been used extensively in
investigations of the affect of limbic system manipulations on
learning and memory. In the passive avoidance task an animal is
punished for making a response (e.g., receives a shock while
drinking from an electrified water tube) and must learn to avoid the


118
Animals with hippocampal or septal area damage show identical
alterations in their performance of all of these tasks (Isaacson
et al., 1971). They are impaired in the acquisition of passive and
one-way active avoidance problems; they perform better than controls
on the two-way task; they show perseverative over-responding on the
FI operant schedule and fail to obtain reinforcements in the DRL
situation because of this over-responding tendency (Isaacson, 1974;
Grossman, 1976). Each of these will be considered in turn.
It was noted in the last section that electrically induced
disruption of amygdala function immediately after training in the
passive avoidance (PA) paradigm resulted in a learning deficit
for that task which mimicked the deficit found after amygdalectomy.
It was hypothesized that the deficit reflected the failure to encode
the negative attributes of that situation in memory. The evidence
indicates that the septal area and hippocamous participate in this
encoding process. Animals with septal lesions also show a deficit
in this task (McCleary, 1961). Grossman and his colleagues have
succeeded in selectively transecting the various connections of
the septum using a specially designed retractable wire encephalotome
(see the review by Grossman, 1976). They found no PA deficit
following transection of the fornix (FNX), the medial forebrain
bundle (MFB), or the stria medullaris (SM), but cutting the stria
terminalis (ST) duplicated the PA deficit seen after septal (and
after amygdalar or hippocampal) lesions. Thus, it appears that it
is the isolation of the amygdala from the septo-hippocampal inte
grating mechanism and the subsequent failure to encode the negative


59
(Such indifference to alcohol is especially interesting in light of
the fact that the brain damage in most of these patients was caused
by years of sustained, heavy drinking.)
Post-mortem examination of the brains of persons who suffered
from Wernicke-Korsakoff disease reveals lesions of certain anatomical
structures in the limbic system associated with the well-known
circuit of Papez. Such damage almost invariably includes, and may
be confined to, injury to the mammillary bodies, the relay for
hippocampal output on its way to the anterior thalamic nuclei
(Brierly, 1977).
A pure form of this memory disorder was the unfortunate consequence
of bilateral removal of the hippocampi in humans. In the 1950s
Scoville performed a series of experimental operations designed to
relieve the symptoms of chronic schizophrenia without the undesirable
side-effects of a complete frontal lobotomy. The surgical procedure
involved the resection of the medial surface of the temporal lobes
from 5.0 to 8.0 cm posterior to the tip of the lobe combined in
some cases with orbital undercutting. Thirty severely deteriorated
schizophrenics had undergone the operation, with slight improvement
in their conditions, when a purely temporal resection was performed
on a nonpsychotic epileptic patient whose seizures were unresponsive
to medication. When this patient, "case H. M.," recovered from
the operation it became apparent that he had developed a severe
amnestic disorder which resembled Korsakoff's psychosis and which
persisted at 14 years (Milner, Corkin & Teuber, 1969). Scoville
and Milner subsequently examined eight of the psychotic patients


60
who had undergone the operation and who were able to participate in
formal testing. They discovered "some generalized memory disturbance
in all patients with removals extending far enough posteriorly
to damage portions of the hippocampus and hippocampal gyrus (Milner,
1958, p. 112). The degree of memory impairment was more or less
proportional to the amount of these structures removed. Bilateral
resection of the uncus and amygdaloid nucleus alone did not result
in amnesia (Brierly, 1977), nor did removal of the gyri of the outer
aspects of the temporal lobes (Bailey, 1946). It has been concluded,
therefore, that "the structures necessary for normal memorizing are
the hippocampal formations within the temporal lobes, the manrnillary
bodies and, possibly, certain thalamic nuclei within the diencephalon"
(Brierley, 1977, p. 221); that is, the hippocampi,, their output
pathways and related projection sites.
There has been, as yet, no definitive explanation of either the
nature of the amnesic deficit described above or of its underlying
mechanisms. The fact that remote memory seems to be intact in
these patients has led many to believe the the hippocampal-
diencephalic structures are not involved in the process of recall,
although Brierley (1977) pointed out that such a conclusion is
unjustified in the absence of adequate pre-post evaluation of this
function. Milner (1966, ch. 5) attempted to account for the pairing
of a period of retrograde amnesia with an inability to learn new
material by hypothesizing that the establishment of a permanent
memory trace requires an extended period of "consolidation," which
is somehow disrupted in this syndrome. In Milner's view the deficit


115
Each of these systems evolved from more primitive mechanisms
which regulated the emission (in the presence of releasing stimuli)
of unlearned (instinctive) behavioral sequences related to the
survival functions of feeding, fighting, fleeing, and reproduction.
In their evolved form these systems allow the decoupling of stimulus
and response, thereby permitting the neocortex to determine behavior,
but they continue to insure that an adequate response is emitted.
Learning and Memory: Animal Studies
The model indicates specific functional/anatomical mechanisms
which could account for the sometimes puzzling deficits seen after
different limbic system ablations in experiments performed by
physiological psychologists.
Animals with hippocampal lesions are sometimes the same and
sometimes different from normal animals in the performance of
tasks for appetitive rewards (Isaacson, 1974, ch. 5). Such
conflicting results have made it difficult to define a specific role
for the hippocampus in memory functions. Studies of human amnesia
victims (reviewed in the previous section) indicate that their
amnesic deficit reflects an inability to select the appropriate
memory trace from the set of available traces, although memories
continue to be laid down in the cortex. A naive animal (with hippo
campal damage) in a strictly controlled laboratory situation may
store only one memory trace (or "hypothesis") for that situation
and not be troubled by competing memory traces. There have been
many reports that animals with hippocampal damage are able to
learn a simple discrimination problem (Isaacson, 1974, ch. 5).


20
Language and the Left Hemisphere
The prominent Soviet geneticist, Theodosius Dobzhansky, observed
that . while all other organisms become masters of their envi
ronments by changing their genes, man does so mostly by changing
his culture, which he acquires by learning and transmits by teaching"
(Dobzhansky, 1964, p. 145). Cultural evolution was made possible
by the development of language, but language is more than a special
form of social communication by which culture is transmitted: it
is the mechanism which produces the adaptive behavior patterns
that released the species from the constraints of biological
evolution. Speech is the fundamental tool of intellectual activity.
Language processes are essential in the operations of abstraction
and generalization, the basis of categorical thinking, and the
vehicle for organizing and regulating mental processes and behavior
(Luria, 1973).
The clearest example of functional brain asymmetry is the
lateralization of language to the left hemisphere, a fact long
established by two observations: deficits in language functions
(aphasia) following damage to the left hemisphere and the retention
of language following right hemisphere injury (right handedness is
assumed throughout the text unless otherwise noted). The patterns
of language deficit following circumscribed lesions provide the
most concrete evidence that is available about the mechanics of
consciousness and its product, cognition. The focus of the following
review is not on language itself, but on the evidence which indicates
that the active operations of consciousness which produce


114
to deal with the stimulus and might activate the pituitary-adrenal
"rapid alarm" system.
The Mobilization System
The cortical mobilizing system is based on the brainstem
reticular formation (RF). Signals from this mechanism lower the
activation threshold of neurons it projects to. The activity of
the RF is modulated by biochemically mediated forebrain systems
whose descending influences reach the RF through the hippocampus
(HPC) and amygdala (AMYG). Ascending RF pathways pass through the
hypothalamus and septal areas and terminate on the nonspecific
thalamic nuclei (e.g., ILTN).
Descending influences from the prefrontal lobe modulate the
activity of the thalamic reticular system and in this way direct
the activation of specific cortical systems. This is accomplished
through an interaction between the nonspecific thalamic nuclei and
the thalamic association nuclei. In this manner the frontal lobes
are able to recruit and program cognitive operations in the post-
central cortex in accordance with intentions.
The Memory, Motivating, and Mobilizing systems supply the neo
cortex with memory indexing information, qualitative and quantitative
emotional data, and activating impulses, respectively. Each of these
categories of information is essential for normal functioning.
Damage within any one of these functional systems results in a
different syndrome of specific clinical deficits. The manifestation
of these deficits may vary with the location of the damage within
the system.


150
where the motivational signals are experienced as subjective emotion.
When this pathway is disrupted (by excision of the left temporal
components) emotional responsiveness is lost. When the right
temporal components are removed, motivational signals are generated
only on the left side and neurotic symptoms result. Failure of the
right hemisphere's monitoring system to activate the right limbic
system results in the failure to mobilize the left hemisphere
(so right processes have increased influence on behavior) and lack of
motivation signals to guide left hemisphere processes.


52
concepts of memory phenomena that are pertinent to the interests
of the applied psychologist.
Experimental psychologists have traditionally approached
the study of memory by subdividing it into registration, retention,
and recall, attempting to isolate and measure these aspects and
the variables which affect them. Rapaport (1961) criticized this
methodology as artificial, insisting that these functions are in
extricably related and that such experiments merely demonstrate
how memory can_ function under given laboratory conditions, Working
from a psychoanalytic perspective, Rapaport preferred to treat
memory as an aspect of cognition. He argued that "actual memory
phenomena are encountered only in the context of thought processes;
at best the classical memory experiments could ignore this fact
and make us ignore it, but they could not produce memory phenomena
outside this context" (Rapaport, 1961, p. 6). He acknolwedged
the difficulties in determining the relation of indistinct entities
such as emotion and memory and attempted to clarify the psychoanalytic
viewpoint by suggesting that "memory is a motivated behavior
phenomenon and . emotions are motivating factors" (Rapaport,
1961, p. 8). This statement, however, appears to beg the question;
if memories are activated by emotions, then what initiates arousal
and affective processes?
The interaction of cognition, affect and memory in the etiology
and cure of psychopathology were central themes in the work, pub
lished in 1893 by Josef Breuer and Sigmund Freud, which gave
psychoanalysis its start. The emphasis on the significance of


183
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388-395.
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Frequency Discrimination and Generalization Following Lesions
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no
of externally provided symbols and aids (e.g., language and rules
of granmar, mathematical tables and formulas). The sequential
operations involved in abstract reasoning, logical analysis, and
propositional speech are made possible by the temporal acuity of the
left hemisphere. The orderly selection (programming) of these
operations depends on the integrity of the tertiary association
zones in the dorsolateral area of the prefrontal lobe.
Three types of information are required at these neocortical
levels in order to guarantee that they will perform their problem
solving tasks appropriately in survival situations. This informa
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above. Each of these systems has limbic, thalamic, and cortical
components (see Fig. 4).
The Memory System
Input to the hippocampal memory system consists of processed
sensory data and/or the products of ongoing cognitive activity.
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to the temporal lobe into the entorhinal cortex. Hippocampal
output emerges in the cingulate cortex as memory indexing information
which is passed on to the I PL where it facilitates access to memory
traces which are associated with the input stimulus. These associa
tions finally emerge in consciousness as formal percepts or concepts
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FUNCTIONAL BRAIN SYSTEMS AND PERSONALITY DYNAMICS
By
DAVID LINDQUIST
A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN
PARTIAL FULFILLMENT OF THE REQUIREMENTS
FOR THE DEGREE OF DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA


149
tend to be lateralized to the left half of the body (Galin,
Diamond & Braff", 1977) suggests a right hemisphere involvement
in these phenomena. The hysteric's characteristic indifference to
these manifestations is similar to that seen with right cortical
lesions, which has been interoreted here as the failue of the right
cortex to trigger subcortical motivation and mobilization processes
In sociopathic patients the CNV is absent or develos only
very low voltage (Cohen, 1974). The tendency for sociopaths and
hysterics to be field-dependent with a global/diffuse cognitive
style was noted earlier. This is consistent with the poorly articu
lated self-conceDts and value systems generally seen in these
personality types. The egocentricity and impulsiveness which
characterize these people tynifies the reflexive, context-bound
functioning of the right hemisphere. All of these trends indicate
that the left hemisphere is less involved in the organization of
behavior in these individuals.
Neurotic symptoms were found to be common following right
temporal lobectomy for epilepsy, while psychopathic disorders were
more frequent following this operation on the left side (Taylor,
1972). These findings provide significant support for the present
functional meta-system model which postulates that normal emotional
responsiveness depends on a motivation system pathway which extends
from the right amygdala (which is activated by the monitoring
system) to the left amygdala and on to the left prefrontal lobe


177
Kincannon, J. C. Prediction of the Standard MMPI Scales from 71
Items: The Mini-Mult. Journal of Consulting and Clinical
Psychology, 1968, 32, 319-325.
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Behavior and Conditioned Avoidance Responses in the Rat. Journal
of Nervous and Mental Diseases, 1958, 126, 57-63.
Kinsbourne, M. Hemispheric Specialization and Human Understanding.
American Psychologist, 1982, 37_ (4), 411-420.
Kintsch, W. Models for Free Recall and Recognition. In D. A. Norman
(Ed.), Models of Human Memory. New York: Academic Press, 1970.
Kluver, H., & Buey, P. C. Preliminary Analysis of the Temporal Lobes
in Monkeys. Archives of Neurology and Psychiatry, 1938, 42, 979-
1000. ~
Knorring, L. von. Visual Averaged Evoked Responses in Patients
Suffering from Alcoholism. Neuropsychobiology, 1979, 2, 233-238.
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81
mammiliothalamic tract in the cat reduced the retention of a two-way
active avoidance task and produced a less striking deficit in the
retention of a one-way active avoidance task. These findings were
later replicated in the rat (Krieckhaus, 1965). Thomas, Frey,
Slotnick and Krieckhaus (1963) studied the post-operative acquisition
of the two-way active avoidance learning task and reported mixed
results: four of their eleven cats were completely unable to learn
the task, while seven of them mastered the problem within the
number of trials required by normal animals. (The significance of
the distinction between acquisition and retention will be discussed
in the following section.)
Cingulate cortex. The cingulate cortex lies above the
corpus callosum on the medial side of the hemisphere, separated
from the neocortex above by the cingulate sulcus. It merges with
the hippocampal gyrus posteriorly and with the neocortex of the
frontal lobe anteriorly. As noted by Papez, the cingulate gyrus
receives its main afferent supply from the anterior nuclei of the
thalamus and projects to the hippocampus via the cingulum bundle.
Stimulation of the cingulate cortex also produces activity in the
prefrontal and orbitofrontal regions of the neocortex (Dunsmore &
Lennox, 1950). There are reciprocal connections with the anterior
and other thalamic nuclei (including the dorsomedial). A strong
projection to the interior parietal lobule (IPL) in the post-central
neocortex has been demonstrated in the monkey (Mesulam, Van Hoesen,
Pandya & Geshwind, 1977). These authors, using the horseradish
peroxidase technique, found that "the cingulate gyrus contained one


26
Human Consciousness, Self-Awareness, and Thought
In the 1960s the general belief in cerebral dominance gave
way to the idea of cerebral specialization due, in large part, to
the "split-brain" studies conducted on the cerebral commissurotomy
patients of Drs. Vogel and Bogan. In 1969 Bogan took the progression
a step further when he revived Wigan's (1844) notion of "the duality
of mind." Wigan (noting the anatomical duality of the brain,
autopsy findings of hemispheric atrophy in patients whose personality
was apparently intact, and introspective evidence of concurrent,
opposing trains of thought) argued that if one hemisphere can
sustain a mind, "it necessarily follows" that a man with two hemi
spheres must have two minds (p. 271). Bogan endorsed this concept
in the third of his influential "Other side of the Brain" papers
(Bogan, 1969a) and concluded that
Pending further evidence, I believe (with Wigan) that
each of us has two minds in one person. . Various
kinds of evidence, especially from hemispherectomy,
have made it clear that one hemisphere is sufficient
to sustain a personality or mind. We may then
conclude that the individual with two intact hemi
spheres has the capacity for two distinct minds.
This conclusion finds its experimental proof in
the split-brain animals who can be trained to per
ceive, consider, and act independently. (1969,
pp. 156-157)
The "dual mind" concept implies two relatively equal but
functionally independent entities which act as opposed forces in
the process of determining behavior. Bogan contributed the hypoth
esis that the two hemispheres utilize different "modes of thought"
in this process: "propositional" on the left and "appositional"
on the right (1969, p. 160)- These appealing ideas were enthusiastically


104
Biochemical and Electrophysiological Aspects of Cortical Mobilization
Processes
It appears that the NE and 5-HT systems interact to control the
energizing output of the reticular formation. At the brainstem
level stimulation of the raphe' (5-HT) nuclei produce calmness
and EEG patterns similar to those of normal sleep, while damage to
these nuclei result in increased motor activity (see Fawcett, 1975).
The raphe' nuclei are thought to be the primary site of action in
morphine-induced catalepsy (see Broekkamp & Lloyd, 1981). Kety
(1972) summarized pharmacological evidence which implicates NE in
the modulation of arousal and sedation. Evidence to be reviewed
below suggests that, at the neocortical level, the 5-HT system
supports a pattern of indiscriminate cortical arousal which facili
tates gross sensitivity at the expense of complex analysis. Another
system mediated by DA ssems to permit the focusing of attention and
may be the means by which the frontal lobes recruit and organize
cognitive activities in the post-central cortex.
Surgical ablation of the prefrontal lobes is sometimes performed
to relieve otherwise intractable pain. This operation is not a
specific cure for pain, however, but a cure for "suffering,"
the source of the pain and its threshold remain unaltered. It is
the person experiencing the pain that is altered by this psycho-
surgical intervention. Petrie (1952; 1960; 1967) found that the
changes in pain tolerance were paralleled by striking differences
in the subjectively perceived intensity of sensation other than
pain. She suggested that the personality and perceptual tendencies


144
ECT in relieving the symptoms of depression (haliday, Davidson &
Brown, 1968; Cronin, Bodly & Potts, 970).
While the dopaminergic activation system appears to predominate
the mobilization of the left hemisphere, the serotonergic arousal
system seems to be more imoortant in modulating the right hemisphere's
activity (see Gottfrieds, Perris & Roos, 1974). In their pharmaco
logical investigation, Mandell and Knapp (1979) found that lithium
treatment significantly reduced serotonin hemispheric asymmetry.
They hypothesized that the phenomenon of mood may be an emergent
property of asymmetrical serotonin regulation, and suggested that
varying degrees of serotonergic asymmetry accounted for the phase
being manic or depressive in bipolar affective disorders. (They
did not speculate on the direction of the asymmetry.)
Flor-Henry (1979) reported on a complex analysis of EEG data
taken from bipolar patients engaged in performing verbal and spatial
tasks (WAIS Vocabulary and Block Design subtests) and in neutral
conditions. He found abnormally high right parietal activity and
variability in depression which became bilateral in mania and
schizoaffective disorders. He also noted that, during spatial tasks,
depressives showed an increase in left temooral activity (versus
neutral conditions); they also showed an increase in right parietal
activity during verbal tasks. He suggested that these changes
indicated that comolex shifts of lateralized hemispheric specializa
tion were taking place. If this is the case, it is tempting to
speculate that such shifts might be caused by a reversal of the
normal inter- and intrahemispheric pattern of interaction between
the DA and 5-HT mobilization systems.


143
the evidence suggests that these entities may be divided into hyper-
and hypo-dominance spectrum disorders. Such a grouping should have
important theoretical implications for the treatment of these
disorders.
Schizophrenia and the Affective Disorders
The neurological correlates of schizophrenia and the affective
disorders have been reviewed by Tucker (1981) and Flor-Henry (1979)
The evidence suggests that these major disorders involve a severe
disruption of intrahemispheric functioning in addition to the
disturbances in interhemispheric information flow which is seen
here as contributing to neurotic and characterological illnesses.
The finding, noted earlier, that left temporal lobe epilepsy
often evolves to a schizophreniform pattern of symptomology is
paralleled by evidence of left hemisphere EEG abnormalities in
schizophrenia (see Tucker, 1981; Flor-Henry, 1979). The evidence
suggests that this abnormal cortical mobilization is mediated by
the dopaminergic activation system. Disruptive overactivation
of the left hemisphere, and right-ear deficiencies in temporal
discrimination tasks observed in schizophrenics were both normalized
following administration of chlorpromazine (Serafetinides, 1973;
Gruzelier & Hammond, 1976), a drug which blocks dopamine receptors.
Bruder and Yozawitz (1979) reported that patients with affec
tive disorders showed left-ear deficiency on dichotic listening tasks
which were correlated with the level of symptomology, indicating
right hemisphere dysfunction. This is consistent with findings
that unilateral right ECT and bilateral ECT were superior to left


171
Cohen, B. D., Berent, S., & Silverman, A. J. Field Dependence and
Lateralization of Function in the Human Brain. Archives of
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A Study of Memory Disturbance and Relief from Depression. Journal
of Neurology, Neurosurgery, and Psychiatry, 1970, 3, 705-713.
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Brain, 1953, 76, 509-512.
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the Regulation of Emotional States. Human Physiolooy, 1975, 1,
394.
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L'Endephale, 1951, 43, 193-200.
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Response from Right and Left Hemispheres. Nature, 1976, 261,
690-692.
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Unilateral Cerebral Disease. Journal of Nervous and Mental
Diseases, 1966, 142, 515-525.
Dobzhansky, T. Heredity and the Nature of Man. New York: Siqnet,
1964.
Doty, R. W. The Unilateral Engram Acta. Neurobiol. Exp., 1973, 33
711-728. ~
Dunsmore, R., & Lennos, R. Stimulation and Strychninization of
Supracallosal Anterior Cingulate Gyrus. Journal of Neuro
physiology, 1950, _13, 207-214.
Edinger, H., & Siegal, A. Functional Aspects of the Hippocampal-
Septal Axis. In J. F. DeFrance (Ed.), The Septal Nuclei.
New York: Plenum Press, 1976.
Efron, R. The Effect of Handedness on the Perception of Simultaneity
and Temporal Order. Brain, 1963a, 86, 216-284.
Efron, R. Temporal PerceDtion, Aphasia, and Deja Vu. Brain, 1963b,
86, 403-424.
Elithorn, A., Gilthero, E., & Slater, E. Leucotomy for Pain.
Journal of Neurology, Neurosurqery and Psychiatry, 1958, 21.
249-261.
Ellinwood, E. R. Amphetamine Psychosis: Description of the Individuals
and Process. Journal of Nervous and Mental Disease. 1967.
144, 272-283. ~


127
They suggested that this hypoarousal might be a factor in the
indifference and other mental status anomalies that are common in
these patients.
Asymmetrical biochemical and electrophysiological processes.
Evidence reviewed in an earlier section delineated two separate
cortical mobilization systems which co-exist in each hemisphere.
A diffuse arousal system is based on the serotonergic (5-HT)
neurons of the median raphe nuclei located in the core of the
reticular formation. An attention focusing activation system
utilizes the catecholamines NE and DA which are derived from the
more laterally placed locus ceruleus and nuclei in the peri
aqueductal gray. The diffuse, indiscriminate arousal of cortical
neurons produced by the 5-HT system results in increased stimulus
reactivity and other phenomena which facilitate the primary objective
of orienting processes (i.e., stimulus sampling and identification).
Conversely, the focusing of attention produced by the NE-DA
activation system is essential to the processes of analysis and
response generation. The proposed model would suggest, therefore,
that 5-HT related mobilization processes would predominate in
the right hemisphere and NE/DA related processes would be more
evident in the left. Gottfries, Perris, and Roos (1974) found that
increased levels of the serotonin metabolite 5-hydroxyindoleacetic
(5-HIAA) in the cerebrospinal fluid was significantly and positively
correlated with the amplitude of auditory evoked potentials from
the right, but not the left hemisphere in a group of mixed
psychiatric patients. On the other hand, increased levels of the


164
psychopathological phenomena with important implications for the
practice of psychotherpay. Such a fully validated model would
permit an integrated approach to diagnosis and treatment planning.
An analysis of the ordering of events in a pathological psychological
process might allow specification of the relationships betv/een
significant experiences (recorded in the right hemisphere system),
the conditions and manner in which these (and associated generalized
expectations) are elicited by stimuli, and the way in which those
are interpreted and/or dealt with by the left hemisphere system. This
information would allow the goals of the treatment process to be
operationalized and indicate intervention points and methods. Such
a plan might specify, for example, that a specific corrective emotional
experience or metaphorical association of specified generalized
expectations to an identified stimulus, followed by a well-defined
cognitive behavior modication and a particular social skill enhance
ment would be the most efficient intervention package. Finally,
biofeedback techniques might find new significance in both the
processes of diagnosis and treatment.


128
NE/DA metabolite homovanillic acid (HVA) in the CSF was significantly
and positively correlated with the amplitude of evoked potentials
from the left, but not from the right hemisphere. These findings
indicate that an identical stimulus triggers mobilization processes
associated with orienting in the right hemisphere and with problem
solving/response-generating in the left.
Bilateral Interaction in Emotion and Cognition
The model outlined above specifies that the quality of signifi
cance is encoded along with context information in neuronal
models (memories) which are stored in the neocortex. When such a
model is activated by an environmental stimulus this coded informa
tion causes the amygdala to generate an emotional response which
will motivate the organism to respond appropriately. This cortical-
limbic system interaction was apparent in a unique experiment
performed by Doty (1973) in which a macaque monkey was prepared
with a unilateral amygdalectomy combined with sectioning of the
optic tract on the opposite side, thus blinding the hemisphere with
the intact amygdala. This was followed by sectioning of all the
forebrain commissures except for the posterior one-third (the splenium)
of the corpus callosum. The splenium was ensnared by a ligature
which was left protruding through the skull. After recovering from
surgery the animal was placed in a large enclosure. When a man
entered the enclosure the animal responded with normal fear and
fled. However, when the commissural transection was completed
(by pulling the snare) the animal's behavior was dramatically
different; it became docile and would actually approach and


67
that do not lend themselves easily to verbal encoding.
. . Thus within the sphere of learning and memory
there is a double dissociation between the effects of
these two lesions. (Milner, 1971, p. 274)
Butters and Cermack (1974) focused on the specifically verbal
aspects of the amnesic disorder in Korsakoff patients and noted
that, during learning tasks, these patients did not react to
changes in semantic categories on successive lists. They concluded
that the amnesic deficit was due to the patient's
inability to encode verbal information along semantic
or meaning dimensions. . Korsakoff patients do not
spontaneously employ semantic encoding strategies,
but rely on basic acoustic and associative categorizations.
If the Korsakoff patient is instructed to encode semanti
cally, he will do so, but in an impaired manner.
(pp. 74-75)
Butters and Cermack assumed that the Korsakoff patients'
deficient utilization of "meaning" in learning tasks was a specifically
verbal (i.e., left hemisphere) phenomenon. However, Gazzaniga and
his colleagues produced evidence which suggests that the right
hemisphere may play an important role in imparting "meaning" to
verbal memory processes. These authors tested patients with partial
or complete section of the cerebral commissures for recall of two
lists of paired-associate nouns. On presentation of the second
list each patient was instructed to "form a 'picture in his mind'
of the two items interacting in some unusual or amusing way"
(Gazzaniga, Risse, Springer, Clark & Wilson, 1975, p. 12). Patients
with partial sectioning of the cerebral commissures showed marked
improvement with the imagery instructions but none was seen where
there was complete section of the hemispheric interconnections.


37
the social environment is critically important to survival throughout
the phylum. Campbell (1974) noted that animal vocalizations and
signals are "emitted only in the presence of the appropriate stimu
lus" (p. 349) and warned against equating these vocalizations with
human speech: "the signals . are generated or motivated by the
phenomenon of emotion, and find their neurological origin not in the
cortex but in the limbic system of the brain" (1974, p. 348). The
cortical organization of these functions in the right brain of humans
appears to mirror that of language in the left hemisphere with
comprehension and expression utilizing anatomical areas homologous to
Wernicke's area and Broca's area, respectively (Ross & Mesulam, 1979).
Right hemisphere responses might achieve direct expression in circum
stances where control by Broca's area in the left hemisphere is
impaired or attenuated, a case in point being the clearly enunciated
emotional exclamations of the frustrated Broca's aphasic. Similarly,
poorly defined and undifferentiated emotionally generated behavioral
impulses (e.g., approach, avoidance) might also achieve motor
expression in the absence of adequate left hemisphere control.
Hughlings Jackson (1864) suggested that if the "faculty of
expression" was proven to be lateralized in the left cerebral
hemisphere it would then be reasonable to expect that its corre
sponding opposite, perception, might be lateralized to the right.
Although the concept of mental "faculties" has given way to an
appreciation of complex functional systems, the role of the right
hemisphere within those systems might, in a broad sense, be said to
conform to Jackson's prediction. While the information processing


135
emotional signals which motivate the organism to respond appropriately
in life or death situations (e.g., fight or flight). These signals
must be transmitted via the left amygdala to the left frontal
lobe before they can achieve conscious status. The subjective
experiences which accompany these "primitive" emotional states are
intrinsically negative, and must be in order to insure a prompt and
vigorous response. The task of evaluating the significance of
environmental stimuli ("monitoring") is relegated to the right
hemisphere in humans, the left being occupied with language functions
which require an incompatible mode of information processing. If
the right hemisphere is damaged and the right amygdala receives
little or no information, then no motivating emotional signal is
generated to be passed on to the left side. In the absence of
these signals the conscious left hemisphere is indifferent to the
stimuli it perceives. If the cortex is mobilized in the absence
of motivational inputs the resulting subjective state might be
characterized as euphoria. Conversely, if a damaged left hemisphere
is receiving emotional input from a normal (or disinhibited) right
hemisphere, and the damage diminishes the left side's ability
to resolve or cope with such input, the resulting experience might
be characterized as catastrophic. In the case of damage to the
right hemisphere's highest, and final, level of information
processing (the IPL), the left would not only fail to receive
emotional/alerting signals but would be deprived of processed
sensory data about the left half of the body and space as well,
from the left hemisphere's point of view these stimuli would cease
to exist.


94
produce pleasure and permit the satisfaction of basic
needs. Similar effects are produced pharmacologically
by potentiation of the noradrenergic (or blockade of
the serotonergic) activity of the median forebrain
bundle. On the other hand, electrolytic lesions of
the median forebrain bundle cause severe deficits in
goal-directed behavior and the loss of consummatory
reactions; again, similar effects are produced by
pharmacological blockade of noradrenergic function or
potentiation of serotonergic function. (Stein et al.,
1972, p. 82)
These authors suggest further that the NE system produces positive
feedback which facilitates ongoing behavior that was rewarded in the
past and the 5-HT system produces negative feedback which terminates
behavior that was unsuccessful or punished. Experimental evidence
supports their contention that these systems also mediate learned
behavior. Wise, Berger, and Stein (1970) demonstrated that drugs
which deplete central stores of serotonin, (and drugs which produce
blockade of serotonergic receptors) significantly reduced a shock-
induced elevation of brain serotonin levels significantly increased
the conditioned behavioral suppression. None of the drugs effected
the behavior of unshocked control animals. The complementary role
of NE was evident in a study where monkeys received bilateral
stimulation or lesions of the locus ceruleus (the source of the
dorsal NE system). Redmond (1979) reported that bilateral locus
ceruleus stimulation elicited behaviors which were associated with
increased fear. Animals with bilateral locus ceruleus lesions
subsequently rose in their social hierarchies because they lost their
fear of previously dominant animals (and humans): they showed
increased motor activity, increased social aggressiveness, and
decreased retreat from threat from pre-lesion levels while retaining


APPENDIX A
HYPO- AND HYPER-DOMINANCE SPECTRUM DISORDERS
BY DSM III DIAGNOSTIC CLASSIFICATION
Hyper-dominance Spectrum
Disorders
295.30 Schizophrenia, paranoid
type
300.40 Dysthymic disorder
300.21 Agoraphobia with panic
attacks
300.22 Agoraphobia without
panic attacks
300.23 Social phobia
300.49 Simple phobia
300.01 Panic disorder
300.30 Obsessive-compulsive
disorder
300.02 Generalized anxiety
disorder
300.00 Atypical anxiety
disorder
301.00 Paranoid personality
disorder
301.40 Compulsive personality
disorder
Hypo-dominance Spectrum
Disorders
300.81 Somatization disorder
300.11 Conversion disorder
307.80 Psychogenic pain disorder
300.70 Hypochondriasis
300.71 Atypical somatoform
disorder
300.12 Psychogenic amnesia
300.13 Psychogenic fugure
300.14 Multiple personality
300.60 Depersonalization
disorder
300.15 Atypical dissociative
disorder
312.31 Pathological gambling
312.32 Kleptomania
312.33 Pyromania
312.34 Intermittent explosive
disorder
312.35 Isolated explosive
disorder
312.39 Atypical impulse control
disorder
301.50 Histrionic personality
disorder
301.70 Antisocial personality
disorder
301.60 Dependent personality
disorder
301.81 Narcissistic personality
disorder
165


50
is critically important to survival throughout the phylum.
Campbell (1974) noted that animal vocalizations and signals are
"emitted only in the presence of the appropriate stimulus" (p. 349)
and warned against equating these vocalizations with human speech:
"the signals . are generated or motivated by the phenomenon
of emotion, and find their neurological origin not in the cortex
but in the limbic system of the brain" (1974, p. 348).
Discussion
It appears that basic human emotional exnerience is an
emergent property of the functioning of mechanisms that originally
served to regulate the emission of SDecies-specific behaviors
which were elicited directly by releasing stimuli in survival-
related situations. The functional system which evolved in humans
decouples stimulus and response. Thus, in humans, it is the emotional
experience evoked by a stimulus--rather than the stimulus itself--
that is the primary motivating factor (reinforcer) which ultimately
determines behavior. Further, this emotional experience might be
most properly considered to be a part of the experiencing person's
environment, since that experience is involuntary and has the power
to condition the person's response. These processes appear to
have their functional impact in the left hemisphere, where the
formulation of behavioral responses occurs.
The physiological mechanisms which motivate adaptive behavior
in humans are centered on the amygdala which integrates information
from the internal and external enviroments in order to attach
emotional significance to stimuli. This structure is involved in the


21
adaptive behavior are part of a functional system that is lateralized
in the left hemisphere of the brain.
Aphasia: Anatomy and Syndromes
Three general types of aphasia have been defined and traced
to different anatomical structures in the left hemisphere. Through
an analysis of these syndromes it is possible to deduce the outline
of a functional system in which these separate areas work together
to produce the complex function of language. The following summary
is based on reviews by Gardner (1975) and Zaidel (1978).
Broca's aphasia, resulting from damage to the inferior, post
frontal zones of the left hemisphere, is an expressive disorder.
Utterances are difficult to initiate and speech is painfully slow
and labored. Patients are usually able to name objects and to repeat,
having less trouble with words that are familiar and concrete.
Although they manage to convey their meanings in a peculiar,
"telegraphic" style, they are unable to produce a fully formed
sentence. Their productions consist almost entirely of substantives;
grammatical parts and forms are absent or impoverished. Nouns are
usually delivered in the singular and verbs appear in their simplest,
noninflected form. Parts of speech that are purely grammatical in
function (conjunctions, prepositions, articles, adverbs) are exceeding
ly uncommon. The same deficit pattern is evident in the patient's
reading and writing.
Although patients with Broca's aphasia may have difficulty in
unravelling complex grammatical relationships (e.g,, "The lion
was eaten by the tiger: which animal is dead?"), their comprehen-


185
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Neurology, 1980, 191, 515-517.
Van Hoesen, G. W. The Differential Distribution, Diversity, and
Sprouting of Cortical Projections to the Amygdala in the
Rhesus Monkey. In Y. Ben-Ari (Ed.), Inserm Symposium No. 20,
The Amygdaloid Complex. New York: Elsevier/North-Holland
Biomedical Press, 1981.


100
in Fig. 2). The septo-hippocampal axis has received the most
attention to date.
Based on their micro-electrode recording studies Edinger and
Siegal (1976) suggested that
Activity in adjacent groups of hippocampal efferent
pyramidal cells could result in the discharge of a
septal neuron. The activation of this neuron will
excite the inhibitory interneuronal network [within
the septal area] to block activity in other portions
of the septum. The septum therefore acts as a
filtering mechanism to Dermit transmission of informa
tion only from that portion of the hippocampus whose
pyramidal neurons are most active at that point in
time. Information transmitted from less active areas
of the hippocampus will be suppressed by the inhibitory
network of the septum, (p. 249)
The septal area appears to have a similar effect on the interneuron
population of the hippocampus. Lynch, Rose and Gall (1977)
conclude from their anatomical investigations that "septal contacts
on [hippocampal] interneurons modulate their activity and thus
allow the 'biasing,' in effect, of the response of the hippocampal
neurons to their excitatory inputs . the septal projections may
serve to modulate the response of the hippocamous to the activa
tion of its massive afferent input from the entorhinal cortex"
(p. 17). Thus, the septal area is in a position to integrate the
activity of the brain's memory, emotion, and arousal systems and
the available evidence supports the suggestion of such a role
for this structure. This arrangement would permit the amygdala,
acting through the septal area, to focus the hippocamoal system's
memory search on associations which have immediate (emotional)
significance for the organism.


11
Stimulation of secondary motor areas produces smooth, coordinated
movements and ablation of these areas may result in the loss of
ability to perform skilled motor acts (apraxia) (Noback & Demerest,
1972).
The systems described thus far are typical of all mammals and
become progressively more elaborate and efficient in the higher
primates. Evolution proceeds generally by modifying and elaborating
existing hardware, allocating new functions to tissues which are in
some way pre-adapted to assume the new tasks (Campbell, 1974).
The development of higher mental functions in man is correlated with
bilateral anatomical expansion of two cortical areas which are
adjacent to the secondary association areas described above. These
regions subserve the highest level of organization in the hierarchies
and are called teritary association areas (Luria, 1973a). Both
areas are involved in what Penfield (1975) referred to as "trans
actions of the mind."
The inferior parietal lobule (I PL) (areas 23; 39, 40) lies at
the anatomical confluence of the secondary association areas in the
post-central cortex. This area is the "association cortex of
association cortexes" (Geshwind, 1979). Here processed information
from the surrounding sensory association areas is further integrated
and synthesized. The area is called "supramodal" because its
individual units can only be excited by the simultaneous stimulation
of two or more sensory modes (Luria, 1973a). Information processing
at this level is "abstract" in that it is independent of a particular
sensory modality (cf. Osgood, 1953). This ability allows the


119
attributes of an experience in memory which produces the PA
deficit (rather than "perseveration" or a "loss of inhibitory
control" as assumed by many authors, e.g., McCleary, 1966). That
the hippocampus (HPC) is essential in this encoding process is
indicated by the finding that animals trained on the PA task before
bilateral HPC ablation were not impaired in the retention of the
learning after surgery (Wishart & Mogunson, 1970), while such lesions
impair the post-surgery acquisition of that problem (Isaacson &
Wickelgren, 1962).
Transection of the FNX, but not of the ST or MFB, duplicated
the pattern of responding seen on the FI schedule after septal and
HPC damage, and partially replicated the disinhibitory affects of
such lesions in the DRL paradigm (Grossman, 1976). Grossman believed
that these results were "due specifically to a disruption of septo-
hippocampal connections" (1976, p. 405). The present formulation
would emphasize the role of the septum in modulating hippocampal
control of the Mobilization system via the RF. Septal lesions
block the theta rhythms normally seen in the HPC following and
alerting stimulus (Smithies, 1966). Failure of the HPC to inhibit
the RF would be expected to result in increased tonic mobilization
and "over-responding," even when such behavior is inefficient.
The puzzling superiority of animals with HPC or septal lesions
over controls in acquiring two-way active avoidance learning
(Isaacson, Douglas & Moore, 1961; King, 1958) is probably due in
part to the peculiarities of the shuttle-box situation which gives
an advantage to animals with a PA deficit. Grossman (1976)


42
before the organism can habituate to a stimulus). In contrast
to the indiscriminate arousal associated with orienting, these
latter processes require the focusing of attention.
The mobilization of selective attention ("activation") appears
to be reflected in the contingent negative variation (CNV) or
"expectancy wave" (Tecce, 1970). The CNV is a special form of
cortical evoked response which consists of a spreading wave of
negative potential that appears whenever there is a contingent
relationship between two stimuli. Negativity develops when brain
tissue is maintaining a readiness for processing (Pribram &
McGuinness, 1975). Thus, the CNV appears whenever the organism
is expecting to perform a perceptual or motor act. The negativity
becomes abruptly positive when that act is executed (Walter, Cooper,
Aldridge, McCall urn, & Winter, 1964). High amplitude CNVs are related
to greater efficiency of perceptual and motor responses; concentra
tion facilitates the CNV while inattention, boredom, or fatigue
decrease it (Cohen, 1974). Elithorn et al. (1958) postulated
that frontal lobe injuries somehow damaged the mechanism under
lying anticipatory sets. It is interesting, in this light, that
the CNV generally appears in the prefrontal lobes and sweeps
posteriorally over the post-central cortex.
Patients with frontal lesions are unable to sustain their
attention. While intelligence, as measured by standardized tests,
may be unimpaired, these individuals are highly distractable and
cannot carry out purposeful activity which is normally directed by
intentions (Luria, 1973a). Luria pointed out that patients with