Title: Midbrain reticular stimulating and generalized drive
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00097963/00001
 Material Information
Title: Midbrain reticular stimulating and generalized drive
Physical Description: vi, 46 leaves : illus. ; 28 cm.
Language: English
Creator: Vierck, Charles John, 1936-
Publisher: University of Florida
Place of Publication: Gainesville, Fla
Publication Date: 1963
Copyright Date: 1963
Subject: Brain   ( lcsh )
Behaviorism (Psychology)   ( lcsh )
Rats   ( lcsh )
Psychology thesis Ph. D   ( lcsh )
Dissertations, Academic -- Psychology -- UF   ( lcsh )
Genre: bibliography   ( marcgt )
non-fiction   ( marcgt )
Bibliography: Bibliography: leaves 33-41.
Additional Physical Form: Also available on World Wide Web
General Note: Manuscript copy.
General Note: Thesis - University of Florida.
General Note: Vita.
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Bibliographic ID: UF00097963
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: alephbibnum - 000537895
oclc - 13015795
notis - ACW1101


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August, 1963



Norma Lee


The author wishes to express gratitude to his

Chairman, Dr. Frederick A. Kins and to Dr. Robert L. King

for their valuable suggestions and assistance. Special

appreciation is also due Professors B. N. Bunnell,

D. C. Goodman, and W. B. Webb for their assistance as

members of the doctoral committee.



DDICATION LEDG .. . . . . . . . 1
A.CY U ;OW:LEDGME; NT S .i....... ... ii1

LIST OF TABLES . . . . . . . . v


IITRODUCTION .. . . . . . . . . 1

METHOD o . o o o e.. e. e o o o 7

Subjects . . . . . . . 7
Surgery . . . . . . . . 7
Stimulation . . * . . . 8
Electrocorticogr recording * * * 9
Gross observations . . . . . . 10
Behavioral tasks . . . . . . 10

Gross activity (GA) . . . . . 10
Fine activity (FA) . . . .. 10
Exploration (EX) . . . . 11
Food reinforcement (FR; FRS) . . 11
Sidman avoidance (AV) . . . . 12
Stimulation valence (VAL) . . .a 12

Behavioral testing procedures . . . 13
Histology . . . .. . . ... 14

RESULTS . . . . . . . . . . . 15

ECG activation . . . . . . . 15
Gross observations . . . . . . 17
Behavioral tasks o . a 19
Histology ..... ..... . . . 24

DISCUSSION . . . . . . . . 26

SUMMARY . .. .. ..... . . 32

BIBLIOGRAPHY . . . . . . . . . 33

APPENDIX * . . . . . 42


Table Page

1. Average differences between stimulation (S)
and non-stimulation (NS) scores for each
task and at each intensity of stimulation . 21

2. Treatment (S vs. NS) by treatment (uA) by
subjects analysis of variance, showing F-ratios
for the two main effects and the interactions
between these effects . . .... .. 22


Figure Page

1. Electrocorticogram recordings, illustrating
arousal induced by reticular stimulation . 16

2. Behavioral performance without stimulation
and as a function of increasing intensity of
stimulation . . . . . . . . 20

3. Diagrammatical representation of electrode
tip placements in the dorsolateral midbrain
reticular formation . . . . . . 25


The idea that reticular activation and generalized

drive might be functionally related has been stated by

Berlyn (1960), Dell (1958), Hebb (1955), Lindsley

(1951, 1958, 1961), malmo (1959), Morgan (1957), and
Schlosberg (1954). For example, Hebb (1955, p. 249)

has stated that ". .. arousal .. is synonomous with

a general drive state .," and Dell (1958, p. 196)

has remarked that "It [the reticular formation] provides

the active source of continued excitation referred to

by psychologists as 'motive', 'sensitizing component',
'drive', etc."

On the basis of psychological data, Brown (1953,
1961) has argued convincingly for the concept of drive

that includes an intensity dimension but no directive

function. A similar view has been expressed by Duffy

(1951, 1957). According to the theoretical formulations
of Hull (1943) and Spence (1956), an increase in nonspecific
drive level will heighten the vigor and/or efficiency of

ongoing behavior, regardless of the particular source of

the drive. Thus, if the reticular formation (RF) serves

as a central mechanism for generalized drive, direct

stimulation of this region should produce performance effects

identical to those brought about by manipulation of

motivational variables. That is, a wide variety of

behaviors should be affected by reticular stimulation

regardless of the source of drive employed during

training, and the nature of the performance change should

parallel that produced by an increase in drive from any

other source.

There is a great deal of indirect evidence support-

ing a correspondence of reticular activity with drive level..

Bolanger and Feldman (reported in I1almo, 1959), Duffy (1932),

Freeman (1948), Lindsley (1958, 1961), and Steiner (1962)

have shown a direct, though sometimes nonmonotonic, rela-

tionship between various physiological measured of arousal

level with performance level and/or drive.

Increased attention is presumably a correlate of

high drive states, and Galambos (1954, 1956), Granit (1955a,

1955b), Hagbarth and Kerr (1954), Hernandez-Peon et al.

(1956), Kerr and Hagbarth (1955), and Rasmussen (1946, 1953)
have demonstrated reticular control over sensory input

from a number of modalities. In this manner information

through one or several sense modalities can be inhibited

by means of reticular influences, thus reducing irrelevant

noiso and allowing focus of attention within one modality.

In addition, high drive states are characterized by

increased sensitivity and reactivity to environmental

stimuli. Fuster (1958), Fuster and Uyeda (1962), and

Isaac (1960) have reduced reaction time with direct

reticular stimulation, and Lindsley (1958, 1961) has

shown the same phenomenon with cortical desynchronization.
Reticular stimulation has also been shown to increase the

resolving power of the cerebral cortex (Lindsley, 1958,

1961) and to lower thresholds for tachistoscopic perception

(Fuster, 1958; Fuster and Uyeda, 1962).

A number of studies dealing with humoral and drug

effects on the RF and behavior have supported the drive

proposition. Bonvallot ct a,. (1954), Courville et al.

(1962) and Dell Lt al. (1954) have shown that adrenaline

stimulates the RF and producer cortical arousal. In

addition to the well known role of sympathetic activity

in emotional behavior, it has been demonstrated that

motivational states of asphyxia (Bonvallet ct al., 1954;

Dell et a., 1954) and hunger (Holzbsuer and Vogt, 1954)

increase adrenaline output. Also, Sawyer (reported by Dell,

1958) found that estrus decreased the threshold for cortical
arousal with RF stimulation. Bradley (1958) has produced

EEG activation with amphetamine and has localized its

effect in the reticular formation. A number of behavioral

studies have reported a facilitating effect of amphetamine

upon performance (Dews, 1958; Morse and Herrnstein, 1956;

Skinner and Heron, 1937; Uyeda and Fuster, 1962; and

Wentink, 1938), although exceptions exist (Kelleher and

Cook, 1959; and Owens, 1960). Chlorpromazine has been

shown by Killam and Killam (1958) to depress reticular

arousal, and Posluns (1962) presents data and cites 13

studies demonstrating inhibition of avoidance behavior and

activity with chlorpromazine.

AlthouGh the above evidence lends impressive

support to the drive hypothesis, very few studies have

tested the effects of direct reticular stimulation in

situations Thcre manipulation of deprivation time or

intensity of noxious stimulation have been utilized to

establish the concept of generalized drive. In fact,

the studies which most closely approach this procedure

have reported inhibitory effects of the stimulation.

Stimulation of various cerebral structures capable of

producing cortical arousal has been shown to inhibit or

impair both learning and performance of complicated choice

behaviors (Gliclman, 1958; rIahut, 1962; Milncr, 1954;

Proctor et al., 1957; Rosvold and Delgado, 1956; and

Thompson, 1958). The areas stimulated in these studies

were: medial and intralaminar thalamic nuclei, brain

stem RF, posterior hypothalamus, and caudate nucleus.

Though these results seem contrary to the drive hypothesis,

they may indicate interference with memory rather than

drive functions.

In order to study adequately the drive mechanism,

rate measures of simple or unlearned responses should be

used. In studies of this nature, Chiles (1954) found

inhibition of bar pressinS for food with medial thalamic

and posterior hypothalamic stimulation, but Mahut (1962)

found no effect on running speed or eating behavior with

thalamic and brain stem reticular stimulation. Grastyan

et al. (1956) applied stimulation to the posterior hypo-

thalamus which facilitated defensive conditioning and

inhibited alimentary conditioning. Bloch and Hcbb (1956)

abolished a simple avoidance response with intralaminar

thalamic stimulation, but reticular stimulation had no

effect. Sheer (1961) found no effect of posterior hypo-

thalamic stimulation upon a conditioned flexion "esponoe,

delayed reaction performance, or bar pressing on a VI

schedule for food. Ingram (1958) inhibited bar pressing

for food with posterior hypothalamic stimulation, but not

with caudate stimulation. Finally, to complicate the

picture further, Ehrlich (1963) found that tegnental

lesions decreased rate of bar pressing for food or water

but increased rate of running to obtain food.

In the present study, the midbrain RF was stimulated

while animals performed a variety of simple tasks that have

been shown to be sensitive to drive effects. Different

incentives were sampled in order to test the generality of

drive effects that may be produced. Motivational studies

that pertain to the tasks used are as follows: (a) Gross

locomotor activity is increased under conditions of food

deprivation (Hall and Hanford, 1954; Siegel and Steinberg,

1949; and Strong, 1957). (b) Fine activity as seen in the

groominG and scratching behavior of animals is decreased by
deprivation (Stro%:g, 1957). (c) The experiments of Alder-
stein and Fchrer (1955), Dashiell (1925), Fehrer (1956),
Thompson (1953), and Zimbardo and Hiller (1958) have
indicated that high drive increases exploration of a noval
environment. (d) Dinsmoor (1952) and Forster and Skinner

(1957) found that increases in deprivation increased the
rates of bar pressing on VI and FR schedules respectively.
Also, Webb and Goodman (1958) trained animals to press a
bar for food, then satiated the Ss and found thai covering

the floor with water increased the rate of bar pressing.

(e) Amsel (1950) and Boren et jl. (1959) have demonstrated
that increasing the intensity of shock or adding hunger
increases the vigor of avoidance behavior.


Sub etos

Twenty-four male, Long Evans, hooded rats of

approximately 70 days of age were obtained from Harland

Small Animal Industries. SurGory was performed on all

animals of this group, and eight of the rats underwent

part or all of the behavioral testing. The experimental

subjects were selected primarily on the basis of a lack of

forced motor movement from stimulation up to at least

150 uA of current. Other factors such as death and

dislodging of electrodes further limited the number of



At about 90 days of age, the subjects received

surgical treatment under nembutal anesthesia injected

intraperitoneally at a dosage of 60 mg/kg. Each rat was

also given 0.2 mg of atropine sulfate intraporitoneally

and 60,000 units of penicillin G intramuscularly. The

skin on the dorsal surface of the skull was incised and

the skull was entered with a 4!2 trephine. A bipolar

electrode of the type described by Valenstein etal. (1961)

was lowered stereotaxically into the RF through the

trephine opening, according to measurements obtained from

De Groot's coordinates (1959). The coordinates used for

the reticular electrodes of the eight experimental animals

were 2 mm lateral to the midline, 0.5 mm anterior to the

0-0 point, and 6 mm in depth from the surface of the

cerebral cortex. Variations of 0.5 mm in the anterior-

posterior and dorsal-ventral dimensions were introduced

in several animals. Reticular electrodes were placed

on the right side in 5 experimental Ss and on the left

side in 3 animals.

Three stainless steel electrodes, bent at a

90 degree angle 2 mm from one end, were hooked under the

skull through small drill holes for supradural electro-

corticogram (ECG) recording. The recording electrodes

were placed over the frontal, parietal and occipital

areas of the side opposite the stimulation electrode.

Stainless steel screws were placed in additional drill

holes to serve as binding posts for acrylic cement which

was used to attach the electrodes to the skull. The skin

was closed around the electrodes, leaving an exposed area

of acrylic. The wound, however, healed well, and infection

was not evident.


Electrical stimulation was delivered to the reticular

formation by a Grass model S4E stimulator which was set to

provide a train of biphasic pulses of 0.1 msec duration

at a rate of 60/sec. The stimulus was fed through an

Argonaut isolation transformer. Current and voltage

were monitored on a Tektronix Type 502 dual-beam oscillo-

scope, and readings were taken from the trailing dodge of

the distorted square wave. Calculated electrode resist-

ances ranged from about 20,000 to 30,000 ohms.

Electrocort icogragm recording

Connections to the cortical recording electrodes

were achieved by attaching miniature pin clutches soldered

to lead wire. Two Grass Model 5P5 polygraph channels were

utilized in obtaining records from frontal-parietal and

parietal-occipital leads. The average resistance between

electrodes was 20,000 ohms.

ECG testing was first carried out as the animals

recovered from surgical Ilembutal pentobarbitall sodium).

Later, at 250 and 400 days of age, the animals received

further testing after injection of 100 mg of Robaxin

(methocarbamol). Robaxin presumably acts upon internuncial

neurons of the spinal cord, thus facilitating relaxation

and drowsiness. Predominantly high amplitude, slow ECG

activity is obtained after injection of Robaxin, and

arousal from either reticular or natural stimulation

occurs more readily with this drug that it does with


Each animal received RF stimulation in an ascending

series of 0.5 volt steps, starting at 1.0 volt and ending

beyond the ECG activation threshold. This threshold was

defined as the point at which high amplitude, slow activity

was completely obliterated by low amplitude, fast activity.

After the initial determination of threshold, a number of

activation tests were carried out at voltages bracketing

the threshold.

Gross observations

On four different occasions, RF stimulation was

delivered in an open field for the purpose of observing

behavioral signs of arousal. Strength of current was

varied in ascending and descending stops of 0.5 volts.

The animals' reactions to external stimuli were tested

with and without RF stimulation by the following

manipulations: (a) The subjects were handled by the

experimenter; (b) a pencil was moved before the

subjects' faces to test visual following; (c) the

animals were lightly prodded and tapped with the pencil;

and (d) the experimenter clapped his hands to produce

an auditory startle response.

Behavioral tasks

Gross activity (GA):- The animals were placed in

an alleyway 24 in. long and 8 in. wide containing photocells

that projected across the midpoint of the long sides. A

count was obtained each time the rat interrupted the beam,

and number of counts was the criterion measure for activity.

Fine activity (FA):- Fine activity was measured in

a tilt cage consisting of an 8 in. diameter cylinder

pivoted at the center of its base. Four metal posts,

equidistantly placed on the periphery of the base and

several mm below the level of the pivotal post, mode

electrical contact with metal plates affixed to the base.

Opposed sets of contacts recorded on the same counter,

and an animal's score consisted of the total counts

registered on the two counters. This apparatus was ex-

tremely sensitive and would count out rapidly for small

movements ouch as those scon with grooming behavior.

Exploration (Ej:':- The purpose of this task was

to determine the number of times subjects would approach

a novel object and the amount of time they would spend

examining it. Scores for the task were obtained by

summing the number of approaches and the time spent

sniffing or touching the object. The enclosure u-t.- was

24 in. in diameter and contained an interestingg" object

that protruded from the center of the floor. The object

consisted of a 6 in. high metal spire emerginG from con-

centric cylinders 3/4 in. and 1 1/2 in. in heiGht. Follow-

ing each two days of testing, another part was added to

the spire to ma:1ntain novelty. First a spring was attached

horizontally from the top, then a yellow wire was hung

from the end of the spring, and finally a small bolt and

nut were attached to the free end of the wire.

Food roinforceaent (FR; FRS):- The animals were

trained to obtain 45 mg Noyes lab rat food pellets by

pressing a bar protruding from one wall of a 12 in.

square enclosure. Five subjects were trained on a variable

interval (VI-30) schedule with a range of five to 55 seconds,

and the remaining two animals were trained on a high fixed

ration(FR-150) that produced strain. The animals were

maintained on an 8-12 hour food deprivation schedule (FR)

in order to keep the motivational level low but effective

in maintaining stable rates of performance. All subjects

were also tested under food satiation (FRS) on an FR-5


Sidman avoidance (AV):- The animals were trained

to avoid shock to the feet by pressing a bar projecting

from one of the short walls of a 12 in. by 6 in. enclosure.

The apparatus was dimly illuminated through 1/2 in. holes

above the bar and in the top of the enclosure, and through

a translucent glass window in the wall opposite the bar.

The apparatus was programmed so that electric shock was

delivered through a grill if 30 seconds elapsed after a

bar press or a previous shock (SS-30-RS-30 schedule).

The shock was maintained for a maximum of five seconds and

could be escaped by a bar press. Four animals were given a

1000 cps tone and light through the window as warning

signals that preceded shock by five seconds and remained

on until shock was terminated.

Sti~ul'.tion valence (VAL):- A test of whether RF

stimulation was positively or negatively reinforcing was

given in which the animals could choose between stimulation

and no stimulation. A shuttle box was divided by a 2 in.

high barrier into two 9 in. by 9 in. compartments with

independent brass grill floors. The floors were pivoted

at one end and suspended by spring tension at the other so

that the weight of an animal tripped a microswitch. The

microswitches controlled relays that allowed recording of

the time spent in each compartment (the criterion measure)

and directed reticular stimulation to the animal only when

he was in a particular compartment. The stimulation

compartment was changed from day to day.

Behavioral testi g procedures

On all tasks except stimulus valence, the subjects

daily received ten minutes of testing with stimulation and

ten minutes without stimulation. Five minute stimulation

(8) and non-stimulation (US) periods were counterbalanced

in the order S-S-NS-S- or ISS--S-1S. Four consecutive days

of testing were given at each stimulation level with the

two counterbalanced orders being alternated over days.

The stimulation values used for testing were given in the

order 10, 25, 50, 75, and 125 uA, with occasional commission

of some levels. For stimulus valence testing, an ascending

series was given with two days at each current intensity,

followed by a descending series. The intensities of

current utilized with each task are indicated in Table 1

of the Results section.

If an animal exhibited consistent facilitatory

effects of the stimulation, he received an additional four

days of testing at the stimulation intensity and tas]:

showing the effect. Consistent facilitation was defined

as facilitation occurring on three days out of the four

days of testing on one stimulus intensity and task.

Behavioral testing was bogun at 120 and continued

to 400 days of a6e. The number of subjects run for all

tasks can be found in Table 2. The order of presentation

of tasks to the subjects was randomized except that gross

activity vwas the first task and stimulus valence was the

last. For food reinforcement and avoidance, the subjects

were trained to a stable level of performance before

stimulation testing was begun. Training prior to testing

was not required with the remaining tasks.


At the end of the experiment, the brain of each

animal, with the electrodes intact, was perfused and fixed

in formalin. The electrodes were then removed, and the

brains were embedded in celloidin. Serial, coronal sections

of 30 micra thickness were cut and appropriate sections

were stained with cresyl violet.


ECG activation

Reticular stimulation of all animals proved capable

of changing high amplitude, low frequency eloctrocortical

activity to the low, fast, desynchronized pattern typical

of activation. Cortical activation thresholds as deter-

mined under surgical anesthesia averaged 120 uA. Testing

at 250 days of age after injections of Robaxin revealed

a lower mean threshold of 40 uA in seven of the animals.

The range for these animals was from 25 uA to 60 uA, and

threshold for the eighth rat was 260 uA. The thresholds

of four animals were obtained at 400 days of age, also

under Robaxin, and an average rise intthreshold of 25 uA

was observed. The latency of observable activation was

typi.:Jlj about one second, and return to high voltage,

slow waves usually required approximately two seconds

after termination of stimulation. Several animals received

stimulation for five minutes and activation was maintained

throughout the interval with the usual latency of return to

high, slow activity. Samples of reticular induced activation

are presented in Fig. 1.

S0 0r-

43 Pc p. o L0

S0 :4 0l L -P
Z Cr43 )
ma 'o a A

t 0 .4 OO
tH W m
0 : ri4)
tO -1 M

000 iU
0D 0 ) 0
(-44- 0 'd 4+

B ) >O Q 0>

l 04 rd -r I
x Cr d 0

q0 0 ok *,
o S 0 H
0 ,C 0 m


Pt .' 4 C) W.-i 0
G)1 F4 O4
5: CO5

ad rldo*

0 d 00 N
*68 a3 OCc,

Gross observations

The first grossly observable reaction to RF

stimulation typically occurred at a threshold of approxi-

mately 50 uA and appeared to be a generalized "alortins".

This response involved some or all of the following com-

ponents: (a) a halting of grooming or perambulation,

here referred to as "freezing", (b) forward movements of

the pinnae, (c) vigorous movement of the vibrissae and

sniffing, (d) exaggerated, irregular respiration,

(e) fixation of the head with apparent visual staring, and

(f) occasional grinding of the teeth. At 75 uA, external

stimuli were presented and produced responses which deviated

only slightly from those elicited in the non-stimulation

animal. The subjects reacted as usual to handling with no

exaggerated struggling. A loud auditory stimulus (hand

clapping) produced normal startle responses. Orienting

was occasionally seen when a pencil was slowly moved

in front of the animals or when it was used to deliver

a tap to their backs. More commonly, however, the visual

and tactual stimuli were ignored with a relative absence of

visual following or turning toward the side tapped.

Within a range of 100 to 250 uA responses were

elicited that appeared to be of a "fearful" nature. This

involved an increase in muscular tone, freezing into a

huddled posture, and occasional bursts of activity in which

the animals backed up to a wall or into a corner. Alerting

was pronounced, and the So appeared to shrink from an

imaginary threat. Several animals would alternately huddle

in a corner and vigorously back across the floor to another

wall or corner. The animals seemed to ignore visual and

tactual stimuli and did not attack or avoid objects placed

before them. Auditory startle responses were observed.

Upon handling, strong attempts wore made to escape the

restraint, and biting sometimes occurred.

When the fear response became intense, within a

range of 150 to 250 uA, five of the animals leaped

spectactularly up the sides of the enclosure. These

jumps did not appear to be directly forced by the stimula-

tion. The responses were not stereotyped, coordination was

excellent, and the jumps in all cases were well directed

toward the top edges of the box. Also, it was discovered

that leaps did not occur when the subjects were stimulated

while on the laboratory floor, free from restraining walls.

In this latter case vigorous running occurred as if designed

to escape the vicinity of stimulation.

Forced motor movements were seen between 150 and

250 uA. Six animals exhibited a contralaterally directed

concavity of the body and turning of the head. In three of

these cases, contralateral circling and/or leaning appeared,

but the other three subjects turned Ipsilaterally. Thresholds

for the responses observed in these animals usually appeared

in the order: alerting, fear, forced movement, and jumping.

At threshold current for forced movement, the circling

and body concavity could be compensated, and the movements

were interrupted by return to normal standing posture.

Jumping behavior usually occurred concurrently with forced

movement but also appeared in animals not showing the latter.

Behavioral tasks

The predominant effect of reticular stimulation

was to inhibit ongoing behavior, as shown by lower scores

during stimulation. This effect was consistent among

subjects and was significantly present for GA, EX, FA, FR,

and AV. As seen in Fig. 2 and Table 1, the amount of

inhibition increased from 10 uA through 125 uA, and the

majority of significant effects were obtained at the higher

intensities. The stimulation also became increasingly

noxious as current was increased. Table 1 gives the

average difference score (minus signs indicating inhibition

and plus signs indicating facilitation) for each task and

at all current levels. In Table 2, F-ratios are given in

analysis of stimulation effects over all currents, effects

between the several stimulation intensities, and the inter-

action effect between the two treatments.



43 0 .............. ............... ... .......

2 0O. .-..., *.

Sso- I
... ......... ......

..50 ... .............0..
c 350- -... *
S230- .
o 190 .........................
o I O .......... .
15 0 -.*
S I00- ...... ..OO
60 ........... .................. ......... ..........
10 25 50 75 125
Stimulalion Current In L.A

Fig. 2. Behavioral performance without
stimulation and as a function of increasing Inten-
sity of stimulation. Each point represents an
average group score over four days of testing.
Stimulation scores are connected by broken lines,
and non-stimulation scores are joined by solid
lines. Statistical probability of less than 5%
as determined by the randomization test for matched
pairs (Siegel, 1956) is indicated by the symbol (*).


Average differences between stimulation (3) and non-
stimulation (NS) scores for each task and at each intensity
of stimulation. Hinus (-) signs indicate greater number of
bar presses or activity and exploration counts during the
lS periods. Plus (+) signs indicate the opposite.
Hinus signs for valence of stimulation indicate more time
spent without stimulation. Abbreviations are as follows:
GA gross activity, EX exploration, FA fine activity,
FR food reinforcement, FRS satiated food reinforcement,
AV Uidman avoidance, VAL valence of stimulation, and
D average difference scores.

10 uA 25 uA 50 uA 75 uA 125 uA

GA D 7.4 5.9 15.2 24.7* 33.3*

EX D 13.3 32.3*

FA D -328.31*

FR D 13.4 37.4* -127.3 -201.9*

FRS D 1.4 + 4.2 14.2

AV D + 10.7 24.0 37.9* 39.3 -113.4'

VAL D 15.3 63.7 75.8 -114.9*

indicates statistical probability of less than 5% as

matched pairs

determined by the randomization test for
(Siegel, 1956).


Treatment (S vs. 113) by treatment (uA) by subjects analysis
of variance,a showing F-ratios for the two main effects and
the interactions between these effects. The abbreviations
are as in Table 1.

I1 S-JNS (df) uA (df) S-1S5 x uA (df)

GA 8 5.53 (1/7) 0.65 (4/28) 2.17 (4/28)
EX 7 5.50 (1/6) 2.54 (1/6) 2.30 (1/6)

FR 7 6.75* (1/6) 2.41 (3/18) 2.34 (3/18)

FRS 5 0.05 (1/4) 1.60 (2/8) 0.30 (2/8)

AV 7 6.04* (1/6) 0.04 (4/24) 3.80*(4/24)

a The statistical analysis was taken from
Lindquist (1953).
indicates statistical probability of less than 5%.

Undoubtedly, the attainment of statistical signifi-

cance of reticular inhibition has been made difficult by

several features of the experimental design. For example,
the variability of scores was increased for the food rein-

forcement and Sidman avoidance tasks by changing the

conditions for some of the animals. This was done in an

attempt to find a procedure that might be sensitive to

facilitatory effects of stimulation. Also, the variability
of scores was probably increased by testing the animals,

in most cases, under low levels of drive. Weak current

levels were chosen with the expectation that facilitation

would occur at these levels if at all. Preliminary testing

with several animals at current values just below threshold

for jumping and forced movement indicated that these values

produced dramatic inhibition of performance. In view of

these factors, a nonparametric, randomization test was

used for the analysis of differences between S and VS

scores at each current level of each task.

As seen in Table 2, significant inhibition of bar

pressing over all current intensities was found for avoid-

ance and food reinforcement. In addition, the significant

interaction for avoidance was produced by an increasing

number of ND bar presses from 10 to 125 uA and a decreasing

number of S presses. This same trend was observed with all

other tasks except FRS.

Retests of animals showing consistent facilitation

over four days at a particular current level and task did

not reproduce the facilitation in any case. The average

of six retest scores was -28.2. No one animal or task

was repeatedly represented in the reruns. All but one

instance of original facilitation occurred at 10 or 25 uA.

One animal that was tested for 16 days on FR at 10 uA

exhibited a consistent and significant facilitatory effect

of stimulation. This animal did not show the facilitation

in any other situation and, in fact, was markedly inhibited

by the stimulation in many cases.

In general, valence of stimulation scores revealed

an increasing aversiveness of the stimulation from 10 to

75 uA. Only one animal preferred the stimulation, and this

effect was prominent only at 25 uA, changing to aversive-

ness at 50 and 75 uA. The only consistent effect observed

for this animal at 25 uA was inhibition of exploration.

Two animals revealed no marked seeking or avoidance of the

stimulation at any current level but distributed their

choices randomly. Both those animals were considerably

inhibited by stimulation on several tasks, primarily at

75 uA and above, but on one occasion at 25 uA.


The reticular electrode placements, as verified

histologically, are diagrammatically presented in Fig. 3.

The range of placements was 5 to 6 1/2 mm ventral to the

cortex, 1.5 to 2.5 mm lateral to the midline, and 0.5 to

1.5 mm anterior to the 0-0 point. The placements were

closely grouped, and all were within the dorsolateral

reticular substance.

Fig. 3. Diagramuatical representation of
electrode tip placements in the dorsolateral mid-
brain reticular formation. Dorsal-ventral and
lateral dimensions are indicated in m. The draw-
ing is taken from De Groot (1959, p. 37), and the
level of section is 0.5 mm anterior to the 0-0
point. Distances anterior to the 0-0 point are
indicated by solid circles (0.5 mm), open circles
(1.0 mm), and a square (1.5 mm).


The grossly observable alerting effect has been

noted by a number of investigators and appears to be an

invariant response to reticular stimulation of moderate

to high intensity. It is interesting though, that the

alerted animals in this study were loss responsive to

external stimuli. This was seen as a diminution in visual

following, orienting to a tactual stimulus, and exploration

of a novel stimulus. The alerting may be comparable to the

arrest reaction produced by Hunter and Jasper (1949) to

intralaminar and medial thalamic stimulation. The thalamic

reaction seems to be more extreme, however, since Hunter

and Jasper saw no auditory startle response or visual


The forced contralateral head turning and body

concavity is contrary to the ipsiversive tegnental response

that has frequently been reported (see review by Skultety,

1962). Turning or circling was variable in the present

study and does not remove the discrepancy. However,

Skultety (1962) found contralateral turning with dorsolateral

midbrain placements in the cat that correspond roughly with

those of the present study. Eased on his work and that of

Sprague and Chambers (1954) with lower brain sten placements,

Skultety proposed a medial (ipsiversive)-lateral (contra-

versive) division that is supported in part by the present


Behaviors very similar to the fear reaction of

this study have been evoked by a number of investigators

from diencephalic and midbrain structures (Delgado, 1955;

Delgado et al., 1954, 1956; Fernandez De Molina and Hun-

sperger, 1959, 1962; Masserman, 1941; Roberts,(19%8, 1962;

Segundo et al., 1955; and Spiegel et al., 1954). Roberts

and Delgado et al. have shown that animals can learn to

avoid stimulation of "alarm" points in the dorsomedial and

lateral thalamus, the tectal area, the medial leaniscus,

and the inferomedial hippocampus, though avoidance learning

was not obtained from "flight" points in the posterior

hypothalamus. If tegmental stimulation produces a true

fear reaction, as these studies suggest, then it would

seem that avoidance performance should have been facilitated

in the present study.

The inhibition of reticular stimulation upon

performance was consistent, with the exception of one

animal showing facilitation on the VI-30 schedule at 10 uA.

Valence of stimulation was not determined for that animal

because of early death, and neither his electrode placement

nor behavior on other tasks give a clue to the reason for

this isolated finding. Roticular stimulation became

increasingly noxious and increasingly inhibitory as current

was increased, but a causal relationship should not be

assumed. Two animals demonstrated behavioral inhibition

with no evidence of escaping the stimulation. It is also

interesting to note that significant inhibition occurred

with fine activity and food reinforcement at 25 uA, which

was below the average ECG arousal threshold. This is not

particularly surprising, however, since that stimulation

was delivered to an awake, aroused animal.

It is improbable that reticular induced inhibition

is specific to certain parameters of the electrical stimulus.

A wide range of current intensities was sampled in the

experiment proper, and frequencies from 1 to 300/sec

were used in preliminary testing on gross activity.

Facilitatory effects were not observed throughout the

frequency range. In moot cases, tegmental stimulation

did not abolish activity, exploration, or operant behavior

but decreased rate to a degree related to the intensity

of stimulation. That is, the correspondence of amount of

inhibition with current value was not the result of

averaging data of an all-or-none nature.

Studies showing facilitation of performance with

drug intervention or spontaneous, generalized activation

are not conclusive evidence of reticular mediated drive,

since the cerebral structures involved in the initiation

of this arousal are not definite. The studies of Lindsley,

Fuster and Isaac, demonstrating enhancement of reaction time,

perception, and cortical receptivity with direct RF stimu-

lation, are more difficult to reconcile with the present

study. Unless species differences are a crucial factor,

it appears that reticular stimulation facilitates perceptual

processes and inhibits performance, and reasons for this

apparent contradiction must be sought.

It is probable that stimulation in the reticular

substance near the medial lemniscus and collicull produces

sensory effects including pain. In fact, Spiegel and

Wycis (1961) have stimulated the reticular formation of

humans subsequent to midbrain spinothalamic lesions, and

elicited reports of pain which was projected to the opposite

side. Such pain could elicit the fear reaction, and either

or both factors would serve as distracting influences to a

performing animal. Even in an avoidance situation, bar

pressing to avoid shock might extinguish during stimulation

periods, since relief from centrally elicited pain and fear

is not achieved. It would seem that such distracting

influences should have also impaired performance in Fuster's

study of tachistoscopic perception, but the nature of the

task and the response required may play an important role.

Fuoter's animals were well trained, and the response was

a discrete occurrence following upon observance of only

one of the briefly illuminated objects. Possibly a dis-

tracting stimulus must be intense to inhibit this reaction

as compared to repetitive, sequential, or complex responses,

which may be disrupted by relatively mild stimulation.

Also, the arrest of activity and alerted posture resulting

from RF stimulation might conceivably enhance perception but

antagonize maintenance of a steady rato of responding.

Another source of explanation is offered by the

inverted U hypothesis of arousal proposed by Malno (1958).

He has reviewed evidence indicating that intense rctivation

impairs performance on tasks measuring rate of performance

and vigilance. Two factors may be of importance here:

(a) The effect has not been demonstrated with tasks involv-

ing discrete trials widely spaced in time, and (b) it may

be difficult to produce subtle increases in activation with

electrical stimulation. The latter point, however, was

not supported by the ECG activation records obtained in

this study. ECG arousal did not appear to be an all-or-none

phenomenon, but as stimulating current approached threshold,

amount of low voltage, fast activity increased until high

voltage, slow activity was obliterated.

Several other possibilities should be considered

for future research. It is conceivable that direct chemical

stimulation of the RF would more closely simulate natural

arousal than electrical stimulation. According to Rinaldi

and Himwich (1955), and Stoiner and Himwich (1962), arousal

can be obtained with adrenergic stimulation in the pontine

reticular formation, and cholinergic stimulation in the

mesodiencephalic activating system.

Also, in view of recent evidence suggesting anatomi-

cal and functional differentiation within the brain stem RF

(Batini et al., 1959a, 1959b; Batsel, 1960; Bonvallet and

Bloch, 1961; Brodal, 1957; Feldman and Waller, 1962;

Huttenlocher, 1961; Jouvet, 1961; and Schlag et al., 1962),

a more detailed exploration of specific reticular areas

with electrical stimulation may be of value. In this

regard, it should be noted that Fuster found facilitatory

effects with dorsal midbrain placements comparable to those

of the present study. Nevertheless, there remains the

possibility that separate reticular areas differentially

express perceptual and motivational functions.


Eight animals were stimulated within the dorso-

lateral midbrain reticular formation while performing

in situations designed to be sensitive to variations in

drive level. Amount of gross activity, fine activity,

exploration, and bar pressing to obtain food and avoid

shock was suppressed as an increasing function of stimulat-

ing current from 10 to 125 uA.

Electrocortical activation was seen in all animals

with reticular stimulation, and grossly observable alerting,

fear, and forced motor reactions were described. The reward

value of stimulation was tested in the majority of animals,

and an increased tendency toward escape of the central

stimulus was seen with increasing amperage.

It was concluded that direct midbrain reticular

excitation does not produce the expected increase in

motivational level that would be evidenced as facilitation

of a wide variety of activities.


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Charles John Vierck, Jr. was born July 6, 1936

in Columbus, Ohio, and attended public school there until

graduation from Upper Arlington High School in 1954. As

an undergraduate the author attended Ohio State University

and the University of Florida. In January, 1959, he received

the B.S. degree from the University of Florida with a group

major in biology, chemistry, and psychology. The M.S.

and Ph.D. degrees were also taken at the University of

Florida in June, 1961, and August, 1963. His area of

gnecialization is physiological psychology.

The author served as Research Assistant at the

University of Florida from 1959 to 1961 and at Vanderbilt

University in the summer of 1961. In 1961-1963, he was

supported by the National Institute of Mental Health as

a Research Follow. The author is married to the former

Norma Lee IIiginson of Jacksonville, Florida.

This dissertation was prepared under the direction
of the chairman of the canditato's supervisory committee
and has been approved by all members of that committee.
It was submitted to the Dean of the College of Arts and
Sciences and to the Graduate Council, and was approved
as partial fulfillment of the requirements for the
degree of Doctor of Philosophy.

August 10, 1963.

Dean, College of Arts and Sciencs

Dean, Graduate School

Supervisory Committee:

IT /'~,PI'


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