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Make-up of spinal cord circuits which process inputs from the femoral-saphenous vein

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Title:
Make-up of spinal cord circuits which process inputs from the femoral-saphenous vein
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Yates, Billy J., 1960-
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English
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ix, 116 leaves : ill. ; 29 cm.

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Subjects / Keywords:
Cats ( jstor )
Diameters ( jstor )
Interneurons ( jstor )
Micrometers ( jstor )
Nerves ( jstor )
Neurons ( jstor )
Skin ( jstor )
Spinal cord ( jstor )
Spinal ganglia ( jstor )
Veins ( jstor )
Afferent Pathways ( mesh )
Dissertations, Academic -- Neuroscience -- UF ( mesh )
Neural Pathways ( mesh )
Neuroscience thesis Ph.D ( mesh )
Saphenous Vein -- innervation ( mesh )
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bibliography ( marcgt )
non-fiction ( marcgt )

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Thesis:
Thesis (Ph.D)--University of Florida, 1986.
Bibliography:
Bibliography: leaves 106-115.
General Note:
Typescript.
General Note:
Vita.
Statement of Responsibility:
by Billy J. Yates.

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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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THE MAKE-UP OF SPINAL CORD CIRCUITS
WHICH PROCESS INPUTS FROM
THE FEMORAL-SAPHENOUS VEIN






By

BILLY J. YATES


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


1986




THE MAKE-UP OF SPINAL CORD CIRCUITS
WHICH PROCESS INPUTS FROM
THE FEMORAL-SAPHENOUS VEIN
By
BILLY J. YATES
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
1986


ACKNOWLEDGEMENTS
The studies presented in this dissertation could not have been
performed without the guidance, assistance and encouragement of a
small army of individuals. Much of the guidance and encouragement
came from the members of my committee: Floyd Thompson, John Munson,
Chuck Vierck and Paul Davenport. Special thanks go to Floyd, from
whom I learned many electrophysiological techniques and a great deal
of philosophy. I also could not have completed this work without the
advice concerning neuroanatomical techniques that I obtained from
A
Parker Mickle, Paul Reier, John Houle, Celeste Wirsig and Roger Reep.
I thank Alan Light for his suggestions regarding the staining of
neurons. I wish to express my utmost gratitude to Deb Dalziel, Bill
Hedden, Albis Acosta, Dan Orlando, Marty Yungmann, Linda Cheshire,
Josephs Cheong, Shoobha Daftary and Ellen Grygotis for superb
technical assistance.
Data were analyzed using the Statistical Analysis System (SAS,
SAS Institute Inc., Cary, North Carolina). Computing was done using
the facilities of the Northeast Regional Data Center of the State
University System of Florida.
Financial support was provided by NIH grant ROI HL 25619, U.S.
Air Force Aerospace Medicine Contract F33615-82-D-0627, the
Department of Neuroscience, the Department of Neurological Surgery,
the Division of Sponsored Research and the Center for Neurobiological
Sciences.
ii


TABLE OF CONTENTS
Page
ACKNOWLEDGEMENTS ii
LIST OF TABLES v
LIST OF FIGURES vi
ABSTRACT viii
CHAPTERS
I GENERAL INTRODUCTION 1
II TRACING OF AFFERENT PATHWAYS FROM THE FEMORAL-
SAPHENOUS VEIN TO THE DORSAL ROOT GANGLIA USING
TRANSPORT OF HORSERADISH PEROXIDASE 5
Introduction 5
Methods and Materials 6
Results 9
Discussion 19
III PROPERTIES OF SPINAL CORD PROCESSING OF FEMORAL-
SAPHENOUS VENOUS AFFERENT INPUT REVEALED BY
ANALYSIS OF EVOKED POTENTIALS 25
Introduction 25
Methods and Materials 26
Results 33
Discussion 41
IV PHYSIOLOGICAL PROPERTIES OF SINGLE SPINAL CORD
NEURONS ACTIVATED BY STIMULATION OF THE
FEMORAL-SAPHENOUS VEIN 44
Introduction 44
Methods and Materials 44
Results 49
Discussion 65
iii


V THE VARIABILITY IN THE RESPONSES OF NEURONS
LOCATED IN DIFFERENT LAMINAE FOLLOWING
STIMULATION OF THE FEMORAL-SAPHENOUS VEIN 70
Introduction 70
Methods and Materials 70
Results 72
Discussion 92
VI GENERAL DISCUSSION 101
REFERENCES 106
BIOGRAPHICAL SKETCH 116
iv


LIST OF TABLES
Page
2-1 Locations of cell bodies in the dorsal root
ganglia giving rise to femoral-saphenous
venous afferents 13
2-2 Conduction velocities of primary femoral-
saphenous venous afferents 23
4-1 Responses of spinal neurons receiving inputs
from the femoral-saphenous vein 53
5-1 Spontaneous firing rates of neurons located in
different laminae 78
5-2 Locations of neurons receiving convergent inputs
from the femoral-saphenous vein, muscle and skin .. 91
v


LIST OF FIGURES
Page
2-1 Cell bodies in the dorsal root ganglia labeled
by the application of HRP to the femoral-
saphenous vein 11
2-2 Diameters of cell bodies in the dorsal root
ganglia of the kitten giving rise to venous
afferents 16
2-3 Diameters of cell bodies in the dorsal root
ganglia of the adult cat giving rise to venous
afferents 18
3-1 Minimal spinal cord processing time for inputs
from muscle, skin and the femoral-saphenous vein .. 35
3-2 Potentials recorded along a typical electrode
track through L6 following stimulation of the
femoral-saphenous vein 38
3-3 The location of the focus of the short latency
negative waves elicited by stimulation of the
femoral-saphenous vein 40
4-1 The effects of stimulation of the femoral-saphenous
vein on the excitabilities of interneurons 51
4-2 Effects of stimulation of muscle, cutaneous and
venous afferents on the excitability of a single
neuron 59
4-3 The response amplitude versus stimulus intensity
relationship for cutaneous afferents 63
5-1 Examples of recording sites marked through the
extracellular iontophoresis of HRP 74
5-2 Locations of sites at which responses were recorded
from venous afferent-activated interneurons 77
vi


5-3 Locations of units excited or both excited and
inhibited by stimulation of femoral-saphenous
venous afferents 80
5-4 Procedures used to determine whether a unit is
activated at monosynaptic latencies by primary
femoral-saphenous venous afferents 83
5-5 Locations of units which were activated at
different minimal latencies following stimulation
of the femoral-saphenous vein 86
5-6 Locations of units activated for different
durations following the stimulation of the femoral-
saphenous vein 89
5-7 Reconstructions of venous afferent-activated
interneurons that were intracellularly stained
using HRP 94
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
THE MAKE-UP OF SPINAL CORD CIRCUITS
WHICH PROCESS INPUTS FROM
THE FEMORAL-SAPHENOUS VEIN
By
BILLY J. YATES
August 1986
Chairman: Floyd J. Thompson
Major Department: Neuroscience
Several characteristics of the afferents which conduct inputs
from the femoral-saphenous vein to the spinal cord and the neurons
within the cord which process these inputs were described.
Experiments using the retrograde transport of horseradish peroxidase
(HRP) from the vein to the dorsal root ganglia were performed to
determine the number, distribution and sizes of the cell bodies from
which the femoral-saphenous venous afferents originate. These
experiments showed that afferents arising along the entire length of
the vein project to very localized spinal levels and that these
afferents are few in number. Most of the cell bodies labeled by the
application of HRP to the femoral-saphenous vein were small in size
viii


(diameter less than 35 micrometers). However, some large cell bodies
(diameter greater than 50 micrometers) were also noted. It was
estimated that over 70% of the femoral-saphenous venous afferents are
C fibers; at least 13% were estimated to be A fibers. The largest
venous afferents were predicted to conduct action potentials at
approximately 60 m/sec.
The minimal spinal cord processing time for inputs elicited by
stimulating large fibers arising in the femoral-saphenous vein was
approximately the same as for inputs from skin; thus, di- or tri-
synaptic pathways appeared to be the shortest circuits separating the
primary afferents and motoneurons. Mappings of field potentials as
well as the recording of single unit activity from reconstructed
sites suggested that most of the first neurons activated by this
afferent input are found in lamina V. Thus, it is likely that many
of the large primary femoral-saphenous venous afferents terminate in
this region.
Recordings were done from single spinal neurons driven by large
venous afferents. The interneurons characterized were excited or
both excited and inhibited for long durations following stimulation
of the femoral-saphenous vein. These neurons were located
predominantly in laminae V and VII. Most neurons could also be
activated by stimulation of large muscle and cutaneous afferents.
Some of the neurons were intracellularly stained; the reconstructed
cells had large cell bodies and extensive dendritic fields.
ix


CHAPTER I
GENERAL INTRODUCTION
The study of the neural substrate of relatively simple reflex
pathways allows an investigator to gain insight into how the central
nervous system organizes and coordinates motor behavior. A spinal
reflex which appears to be produced by relatively simple central
nervous system circuitry is elicited when receptors in the walls of
the femoral-saphenous vein are stimulated (Thompson and Barnes, 1979;
Thompson and Yates, 1983, 1984, 1986; Thompson et al., 1982, 1983).
The femoral-saphenous venous afferents include fibers excited by
intravenous perfusion pressures as low as 2-3 mm Hg (Thompson et al.,
1983). Electromyographic recordings following low-threshold
electrical stimulation of these afferents showed that they can elicit
the simultaneous contraction of flexor and extensor hindlimb muscles
(Thompson et al., 1982). It is interesting that both flexors and
extensors are simultaneously excited by stimulation of afferents
arising from the femoral-saphenous vein, as other known hindlimb
reflexes involve the reciprocal facilitation and inhibition of
motoneurons. Stimulation of cutaneous afferents or other
exteroceptors generally results in flexor facilitation and extensor
inhibition (Sherrington, 1910; Lloyd, 1943a; Hagbarth, 1952).
Activation of muscle afferents typically produces the reciprocal
facilitation and inhibition of agonists and antagonists (Lloyd,
1


2
1943a; Laporte and Lloyd, 1952; Matthews, 1969). The unique
segmental reflex elicited by venous afferent stimulation has been
suggested to provide a rapid means of controlling the counterpressure
applied to the high-capacitance intramuscular veins; changes in this
counterpressure are potentially important in compensating for
orthostatic blood shifts (Thompson and Yates, 1983, 1984, 1986).
It is well established that other inputs from the internal
environment, in addition to those from the femoral-saphenous vein,
converge with somatic inputs in the spinal cord on interneurons that
are part of somatic reflex pathways (Downman, 1955; Evans and
McPherson, 1958, 1959, 1960; Evans, 1963). Spinal neurons which send
their axons into ascending tracts, including the postsynaptic dorsal
column pathway (Rigamonti and Hancock, 1974; Rigamonti et al., 1978),
the spinocervical pathway (Rigamonti and Michelle, 1977) and the
spinoreticular and spinothalamic pathways (Hancock et al., 1975;
Foreman and Weber, 1980; Blair et al., 1981, 1984; Milne et al.,
1981; Rucker and Holloway, 1982; Ammons et al., 1984; Rucker et al.,
1984), also receive visceral inputs. These inputs arise in part from
Pacinian corpuscles that sometimes discharge in phase with
cardiovascular pulsations (Gammon and Bronk, 1935), thermoreceptors
that are located at least in part along blood vessels (Thompson and
Barnes, 1970; Rawson and Quick, 1972; Riedel, 1976), vascular
receptors that respond to visceral movement or distention (Bessou and
Perl, 1960; Floyd and Morrison, 1974; Floyd et al., 1976),
proprioceptors found in the intercostal muscles and in the diaphragm


3
(Yamamoto et al., 1960; Remmers and Tsiaras, 1973; Davenport et al.,
1985; Marlot et al., 1985), mechanoreceptors located in the vagina
(Komisaruk and Wallman, 1977; Henry, 1983) and chemoreceptors
innervating blood vessels (Moore and Moore, 1933; Lim et al., 1962;
Potter et al., 1962; Besson et al., 1972). It would be of great
interest to compare the spinal cord circuits which process inputs
from the femoral-saphenous vein with the circuits that process other
inputs from the internal environment.
The present study examined several of the characteristics of the
afferents which conduct inputs from the femoral-saphenous vein to the
spinal cord and the neurons within the cord which process these
inputs. Experiments using the retrograde transport of horseradish
peroxidase from the vein to the dorsal root ganglia were performed to
determine the number, distribution and sizes of the cell bodies from
which the femoral-saphenous venous afferents originate. The results
of these experiments are described in Chapter II. In addition, field
potential mappings were performed to determine the location of the
first interneurons excited by the venous afferents. These mappings
also demonstrated the shortest pathway between the primary afferents
and motoneurons; this information is presented in Chapter III. The
electrophysiological properties of the venous afferent-activated
interneurons are discussed in Chapter IV. Interneurons with
different locations in the cord were shown to be affected by venous
afferent stimulation in different ways; the variability in how the
excitabilities of neurons in the different laminae of Rexed were
altered by vein stimulation is presented in Chapter V. Finally


4
Chapter VI considers the hypothesis that the characteristics of the
spinal cord circuitry which processes inputs from the
femoral-saphenous vein are similar to the characteristics of the
circuitry, described in previous studies, which processes other
inputs from the internal environment.


CHAPTER II
TRACING OF AFFERENT PATHWAYS FROM THE FEMORAL-SAPHENOUS VEIN TO
THE DORSAL ROOT GANGLIA USING TRANSPORT OF HORSERADISH PEROXIDASE
Introduction
In the mammal, intracranial (Davis and Dostrovsky, 1985; Keller
et al., 1985; McMahon et al., 1985; Norregaard and Moskowitz, 1985),
extracranial (McMahon et al., 1985; Norregaard and Moskowitz, 1985),
and visceral (Bessou and Perl, 1960; Lim et al., 1962; Floyd and
Morrison, 1974; Floyd et al., 1976; Guilbaud et al., 1977; Vance and
Bowker, 1983; Barja and Mathison, 1984) blood vessels receive an
extensive afferent innervation. Anatomical studies have also
demonstrated the presence of large, myelinated fibers coursing along
the walls of large veins of the hindlimb and into adjacent nerve
trunks (Woollard, 1926; Hinsey, 1928; Truex, 1936; Polley, 1955).
These observations are further supported by experiments in which
electrical stimulation of carefully isolated segments of the
femoral-saphenous vein elicited (i) compound action potentials
recordable from the femoral nerve and dorsal root fibers entering the
lower lumbar spinal cord (Thompson and Barnes, 1979; Thompson and
Yates, 1986), (ii) evoked field potentials recordable from the lumbar
cord (Thompson and Barnes, 1979; Thompson and Yates, 1986), (iii)
changes in firing patterns recordable from single spinal motoneurons
(Thompson et al., 1982) and (iv) electromyographic activity
recordable from hindlimb muscles (Thompson et al., 1982). The
5


6
injection of fluid pulses into the femoral-saphenous vein or
longitudinal stretch of the vein wall similarly elicited field
potentials in the lumbar spinal cord and activity recordable from
ventral roots (Thompson et al., 1983).
The study presented in this chapter was performed in order to
gain insight concerning the projection pattern of these
femoral-saphenous venous afferents to the spinal cord. The
retrograde transport of horseradish peroxidase (HRP) from the
femoral-saphenous vein to the dorsal root ganglia was used to
determine the range of afferent sizes, the relative numbers of these
afferents, and the spinal cord segments to which they project.
Methods and Materials
HRP Application Procedures
Experiments were conducted on kittens (5-6 weeks of age) and
adult cats (weighing 2.5-3.8 kg). Animals were anesthetized using an
intraperitoneal (ip) injection of sodium pentobarbital (Butler, 38
mg/kg) or an intramuscular (im) injection of Ketaset (ketamine
hydrochloride, Bristol, 10 mg/kg) combined with Rompun (xylazine,
Haver-Lockhart, 0.5 mg/kg). Maintenance doses of sodium
pentobarbital (12 mg/kg) or Ketaset (5 mg/kg) were delivered as
necessary to maintain areflexia during the time that tissues were in
contact with HRP.
The skin of either of the hindlimbs was opened ventrally to
expose the femoral-saphenous vein and underlying muscle. In 3
kittens and 9 adult cats, one to three 10-15 mm segments of the vein


7
were separated from surrounding tissues and placed on rubber dams. A
segment of the proximal femoral vein (overlying the vastus medialis
and adductor femoris muscles) was always prepared for HRP
application; other HRP application sites were located distally and
were spaced 3-6 cm apart. The tissues adjacent to the isolated vein
segments were covered with a thick layer of petroleum jelly to
prevent spread of the tracer. Uptake of HRP was promoted by making
small tears in the outer vein wall with sharp forceps so that fibers
coursing there would be damaged. A 30% solution of Sigma Type VI HRP
in 2% dimethylsulfoxide was applied to the prepared vein segments;
after 3-4 hrs the enzyme was removed from the vein and the
application sites were irrigated with saline. Incisions were closed
using wound clips and the animals were returned to their cages and
allowed to recover.
As a control for these experiments, different HRP application
sites were used in 2 adult cats. In these animals segments of the
femoral-saphenous vein were separated from the adjacent muscle as
described above. However, the HRP solution was applied to the muscle
which lay directly underneath the 10-15 mm segment of vein that had
been dissected away; these tissues were the most likely to be
accidentally exposed to HRP during those experiments in which the
tracer was applied to segments of the femoral-saphenous vein. In one
experiment 2 application sites located 31 mm apart were used; in the
other 3 application sites separated by 20 and 22 mm were used. The
distribution of cells in the dorsal root ganglia labeled by this
procedure was compared with that of cells labeled after the enzyme
was applied to the femoral-saphenous vein.


8
Histological Procedures
After a recovery period of 50-60 hrs in kittens and 90-96 hrs in
adult cats, the animals were reanesthetized; kittens were
transcardially perfused with 250-500 ml of chilled saline containing
0.1% sodium nitrite and 1.5 units of heparin per ml followed by 2 1
of chilled 1.25% glutaraldehyde/0.6% paraformaldehyde fixative.
After an additional 30 min the fixative was replaced with chilled 0.1
M phosphate buffer (pH 7.4). Adult cats were similarly
transcardially perfused; however, twice the volume of solutions were
used than in the smaller animals.
The spinal cord and lumbosacral dorsal root ganglia were exposed
by dorsal laminectomy. The L3-S1 dorsal root ganglia (on the side
exposed to HRP) and the L6-S1 spinal cord segments were removed and
placed in 0.1 M phosphate buffer (pH 7.4). The tissues were cut into
50 micron sagittal sections using a vibratome (Oxford).
Histochemistry was performed using the chromagen tetramethylbenzidine
according to the procedures of Mesulam (1982). Tissues were
counterstained using neutral red, rapidly dehydrated and cleared with
xylenes. Sections were observed using both bright- and dark-field
illumination. Neuronal cell bodies in the dorsal root ganglia were
noted to be oval in shape (see Figure 2-1). Both the long (major)
and short (minor) axes of labeled cells were measured using an
eyepiece graticule. The mean of these two measurements was taken to
give the average cell diameter. In addition, the dimensions of
unlabeled cells in representative sections were also measured to
indicate the range of sizes of cell bodies present in the ganglia.


9
Controls for the Diffusion of HRP
In all experiments the ventral horn of the spinal cord was
scrutinized for the presence of labeled motoneuron cell bodies. In
those experiments in which HRP was applied to the muscle underlying
the femoral-saphenous vein numerous densely labeled motoneuron cell
bodies were noted. Thus, the presence or absence of labeled
motoneurons appeared to be an adequate indicator as to whether HRP
had diffused into the muscle underlying the femoral-saphenous vein.
If any labeled cell bodies were noted in the ventral horn of a
preparation in which HRP was applied to isolated segments of the
femoral-saphenous vein, the experimental results were discarded and
not analyzed further.
Results
Five of the 12 experiments (42%) in which HRP was applied to the
femoral-saphenous vein were successful; in these experiments labeled
neuronal cell bodies were present in the dorsal root ganglia and no
labeled motoneuron cell bodies could be detected in the spinal cord.
Examples of cell bodies in the dorsal root ganglia labeled by the
application of HRP to the femoral-saphenous vein are shown in Figure
2-1.
Following the application of HRP to the femoral-saphenous vein,
an average of 289 +/- 402 (mean +/- S.D.)^ labeled cell bodies in the
dorsal root ganglia was counted per experiment; an average of 182
*Other confidence intervals presented here also represent mean +/-
standard deviation.


Figure 2-1. Examples of cell bodies in the L5 and L6 dorsal
root ganglia labeled by the application of HRP to the
femoral-saphenous vein. Calibration bars represent 50
micrometers. Labeled neuronal cell bodies are denoted by
arrows.




12
+/- 232 cells was counted per vein segment exposed to HRP. The top
portion of Table 2-1 shows the locations of cell bodies in the dorsal
root ganglia labeled by the application of HRP to the vein. In 3
preparations more than 95% of the labeled cell bodies counted were
confined to the L6 dorsal root ganglion; in 1 preparation all of the
labeled cell bodies were confined to the L5 dorsal root ganglion; in
the other preparation 77% of the labeled cell bodies counted were
located in the L5 dorsal root ganglion, and the other 23% were
located in the L6 ganglion. In total 78% of the labeled cells
counted were located in the L6 dorsal root ganglion, 21% were located
in the L5 ganglion and less than 1% were located at other levels.
The numbers and locations of cell bodies in the dorsal root
ganglia labeled by the application of HRP to muscle underlying the
femoral-saphenous vein (bottom portion of Table 2-1) were quite
different from those described above. In one experiment 6285 labeled
cells were counted; in the other 6251 were counted. The cell bodies
labeled by the application of HRP to the muscle underlying the
femoral-saphenous vein had a more widespread distribution than did
the cell bodies labeled by HRP application to the vein itself. In
total 3.9% of the labeled cells counted were located in the L3 dorsal
root ganglion, 7.3% were located in the L4 ganglion, 8.6% were
located in the L5 ganglion, 21.2% were located in the L6 ganglion,
30.5% were located in the L7 ganglion and 28.5% were located in the
SI ganglion. It is reassuring that most of the cell bodies labeled
by the application of HRP to muscle underlying the femoral-saphenous
vein were located at levels caudal to those containing the cell
bodies labeled by HRP application to the vein itself.


13
Table 2-1. Percentage of cell bodies in different dorsal root
ganglia labeled by the application of HRP to the femoral-saphenous
vein (top) or to the tissues underlying the vein (bottom).
HRP Applied to the Femoral-Saphenous Vein
Experi
ment #
# of Sites
to which
HRP Was
Applied
# of
Cell
Bodies
Labeled
Percentage of Labeled Cell Bodies
Counted in Each Ganglion
L3 L4 L5 L6 L7 SI
1
1
391
0
0
77
23
0
0
2
2
953
0
0
3
96
1
0
3
2
45
0
0
0
100
0
0
4
3
51
0
0
0
100
0
0
5
3
7
0
0
100
0
0
0
SUMMATION

1447
0
0
21
78
1
0
HRP Applied to the Muscle Underlying the Femoral-Saphenous Vein
1
2
6251
0
4
10
25
36
25
2
3
6285
8
11
7
18
25
32
SUMMATION

12536
4
7
9
21
31
29


14
Figure 2-2 shows the diameters of cell bodies in the dorsal root
ganglia labeled by the application of HRP to the femoral-saphenous
vein of the kitten; the figure also shows the diameters of unlabeled
cells measured in representative sections to indicate the range of
sizes of cell bodies present in the ganglia. Figure 2-3 shows
similar data collected from adult cats. Data derived from kittens
and adult cats are shown separately because the cell bodies in young
animals were noted to be smaller than those in the adult. Cell
bodies in the kitten labeled by the application of HRP to the
femoral-saphenous vein had an average diameter of 25.4 +/- 8.6
micrometers. The majority (87.5%) of the labeled cells were small in
size (diameter < 35 micrometers); only 1.6% were large (diameter > 50
micrometers). A total of 694 unlabeled cells were measured in the
kitten for comparison with the labeled cells. Unlabeled cells had an
average diameter of 30.5 +/- 11.3 micrometers; this mean was
significantly larger than that for labeled cells by Student's t-test
(p < 0.0001). Seventy-three percent of the unlabeled cells were
small in size (diameter < 35 micrometers); 6.5% were large (diameter
> 50 micrometers).
The relative sizes of cell bodies in the dorsal root ganglia
labeled by the application of HRP to the femoral-saphenous vein and
unlabeled cell bodies were similar in the adult cat to the relative
sizes in the kitten. Cell bodies in adults labeled by the appli
cation of HRP to the femoral-saphenous vein had an average diameter
of 32.9 +/- 14.0 micrometers. Most (71.8%) of the labeled cells were
small in size (diameter < 35 micrometers); only 13.2% were large


Figure 2-2. Diameters of cell bodies in the dorsal root ganglia labeled by the
application of HRP to the femoral-saphenous vein of the kitten. The diameters of
unlabeled cells measured in representative sections to indicate the range of
sizes of cell bodies present in the ganglia are also shown.


KITTEN-)
Labeled Cells Unlabeled Cells
Diameter in jjm


Figure 2-3. Diameters of cell bodies in the dorsal root ganglia labeled by the
application of HRP to the femoral-saphenous vein of the adult cat. The diameters
of unlabeled cells measured in representative sections to indicate the range of
sizes of cell bodies present in the ganglia are also shown. Vertical dashed
lines indicate divisions into small, intermediate and large-sized cells according
to the scheme proposed by Lee et al. (1986).


ADULT CAT
(n =4)
Labeled Cells
Unlabeled Cells
Diameter in jjm


19
(diameter > 50 micrometers). A total of 1255 unlabeled cells were
measured in the adults for comparison with the labeled cells.
Unlabeled cells had an average diameter of 41.7 +/- 19.5 micrometers;
this mean was significantly larger than that for labeled cells by
Student's t-test (p < 0.0001). Forty-five percent of the unlabeled
cells were small in size (diameter < 35 micrometers); 24.1% were
large (diameter > 50 micrometers).
Both large and small cell bodies in the dorsal root ganglia of
cats were labeled by the application of HRP to the femoral-saphenous
vein; however, most of the labeled cells were small. Labeled cell
bodies were smaller, on the average, than other cell bodies located
in the ganglia; in addition, the percentage of large labeled cell
bodies counted in the ganglia was smaller than the percentage of
large unlabeled cell bodies. However, the anterograde transport of
HRP from the dorsal root ganglia to afferent terminals in the spinal
cord was far less extensive than the retrograde transport from the
vein to the ganglia, if it occurred at all. The transganglionic
labeling of intraspinal portions of afferents was never detected
using either bright- or dark-field illumination following the
application of HRP to the femoral-saphenous vein.
Discussion
Previous electrophysiological studies in the cat have shown that
electrical or mechanical stimulation of isolated segments of the
femoral-saphenous vein can elicit potentials recordable from the
lumbar spinal cord (Thompson and Barnes, 1979; Thompson and Yates,
1983, 1984, 1986; Thompson et al., 1982, 1983). This study extends


20
these findings by providing an anatomical demonstration of the
number, distribution and sizes of the cell bodies from which these
femoral-saphenous venous afferents originate. The selective labeling
of afferents arising in the walls of the femoral-saphenous vein in
these preparations was supported by several lines of evidence. No
labeling of spinal motoneurons could be detected in these studies,
suggesting that HRP had not diffused from the vein to adjacent
muscle. Further evidence suggesting that no spread of tracer
occurred is that the application of HRP to the tissues adjacent to
the femoral-saphenous vein produced labeling of cell bodies found
predominantly in dorsal root ganglia caudal to those containing
labeled cell bodies following application of HRP to the vein itself.
These data also suggest that the afferents which were labeled
terminated in the walls of the femoral-saphenous vein and were not
fibers of passage coursing along the vein but arising in muscle. It
would be very unlikely for all fibers of passage from muscle to be
sensory afferents; if fibers arising in muscle had picked-up HRP in
these experiments, the labeling of motoneuron cell bodies should have
occurred. In addition, if fibers of passage from muscle had
transported HRP, the distribution of labeled cell bodies in the
dorsal root ganglia should have been similar to that following the
application of tracer directly to the muscle. Since the cells
labeled by the application of HRP to the femoral-saphenous vein had a
different distribution, it is likely that they also had a different
origin. Similarly, it is unlikely that fibers of passage arising in
skin were labeled in these studies. The skin overlying the


21
femoral-saphenous vein in the intact animal is innervated by the
saphenous nerve; this nerve projects to several spinal levels
(Bernhard, 1953; Crouch, 1969). If fibers of passage from skin had
picked-up HRP in these experiments, a more widespread distribution of
labeled cell bodies in the dorsal root ganglia would have been
expected. It is also noteworthy that the cell bodies in the dorsal
root
ganglia
labeled
by
the
application of HRP
to the
femoral-saphenous
vein
were
significantly
smaller, on the
average,
than
other cell
bodies
in
the
ganglia.
If a general
class of
hindlimb afferents, and not a specific population, had taken up HRP
in these studies, the diameters of the labeled cells should have been
identical to those of other cells in the ganglia.
A recent study by Lee et al. (1986) has made it possible to
correlate the diameter of a cell body in a dorsal root ganglion with
the conduction velocity along the peripheral process arising from it.
These investigators made intracellular recordings from single cell
bodies in the dorsal root ganglia of adult cats using microelectrodes
filled with an HRP solution, determined the conduction velocity along
the peripheral process to the cell body, and then injected the tracer
into the neuron so it could later be visualized. They reported that
cell bodies less than 35 micrometers in diameter gave rise to
peripheral processes that carried impulses at less than 2.5 m/sec (C
fibers). Cell bodies greater than 50 micrometers in diameter were
reported to give rise to peripheral processes that carried impulses
at greater than 2.5 m/sec (A fibers). For these neurons there was a
linear correlation between the diameter of the cell body and the


22
impulse conduction velocity; this relationship was defined by the
following equation:
Conduction Velocity = (0.84 Diameter of Cell Body) 20.3.
However, there was no predictable relationship between the diameters
of intermediate-sized cell bodies (35-50 micrometers) and the
conduction velocities along their peripheral processes.
Most (71.8%) of the cell bodies labeled by the application of
HRP to the femoral-saphenous vein of the adult cat were smaller than
35 micrometers in diameter; these cell bodies would be expected to
give rise to slowly conducting peripheral processes. However, 13.2%
of the cell bodies labeled by the application of HRP to the vein were
larger than 50 micrometers in diameter; these cell bodies should give
rise to rapidly conducting peripheral processes. The estimated
conduction velocities along these processes are shown in Table 2-2;
the table also shows the predicted conduction velocities of other
afferents based upon the diameters of cell bodies that were not
labeled following the application of HRP to the vein. The largest
femoral-saphenous venous afferents were estimated to have conduction
velocities of approximately 60 m/sec. Fifteen percent of the cell
bodies labeled by the application of HRP to the femoral-saphenous
vein were intermediate in size (35-50 micrometers); no estimate could
be made of the conduction velocities along these afferents. Blood
vessels other than the femoral-saphenous vein have previously been
shown to be innervated by thermoreceptors (Fruhstorfer and Lindblom,
1983), chemoreceptors (Moore and Moore, 1933; Lim et al., 1962;
Potter et al., 1962; Besson et al., 1972) and slowly conducting


23
Table 2-2. Estimated conduction velocities along the peripheral
processes of large primary femoral-saphenous venous afferents and
other large primary sensory neurons determined from the diameters of
the cell bodies.
Conduction Velocity Percentage of Cells Falling into the Range
Range Femoral-Saphenous Other Afferents
Venous Afferents
20-30 m/sec
47
37
30-40 m/sec
36
32
40-50 m/sec
15
19
50-60 m/sec
2
10
> 60 m/sec
0
2


24
mechanoreceptors (Bessou and Perl, 1960; Floyd and Morrison, 1974;
Floyd et al., 1976). It is likely that the femoral-saphenous vein is
also innervated by one or more of these receptor types as well as by
rapidly conducting afferents that have not been reported to be
associated with other vessels.
Only a small number of afferents were labeled by the application
of HRP to the femoral-saphenous vein (an average of 182 per vein
segment exposed to the enzyme), and only a small fraction of the
afferents labeled were large. However, electrical activation of
A-alpha/beta femoral-saphenous venous afferents has been reported to
produce large field potentials recordable from the cord (Thompson and
Barnes, 1979; Thompson and Yates, 1986). For the anatomy and
electrophysiology to be in register, it would seem that the
divergence of the inputs from the femoral-saphenous vein to the
spinal cord is substantial. Such divergence could be accomplished by
either the extensive branching of the intraspinal portions of the
afferents or the activation of second-order spinal interneurons that,
in turn, make contacts with a large number of higher-order cells.
Visceral afferent inputs into the lower thoracic spinal cord of the
cat have previously been shown to exhibit a similar widespread
divergence (Cervero et al., 1984).


CHAPTER III
PROPERTIES OF SPINAL CORD PROCESSING OF FEMORAL-SAPHENOUS VENOUS
AFFERENT INPUT REVEALED BY ANALYSIS OF EVOKED POTENTIALS
Introduction
The study presented in Chapter II suggested that afferents with
both large and small diameters project from the femoral-saphenous
vein to the lumbosacral spinal cord of the cat. In most preparations
the L6 segment was the input cord segment for these primary
afferents. The study discussed in this chapter examined two
characteristics of the interneurons which link inputs along these
afferents with spinal motoneurons. The minimum time required for
activation of spinal motoneurons by the femoral-saphenous venous
afferents was measured to estimate the least number of interneurons
interposed between the primary afferents and motoneurons. In
addition, focal synaptic field potentials were mapped to determine
the location of the interneurons monosynaptically excited by
stimulation of the femoral-saphenous vein.
The field potential mapping techniques utilized in this study
were first described by Gasser and Graham (1933); the intracord
mapping of potential fields has since been performed in a number of
studies (Campbell, 1945; Austin and McCouch, 1955; Howland et al.,
1955; Coombs et al., 1956; Fernandez de Molina and Gray, 1957; Willis
et al., 1973; Fu et al., 1974; Beall et al., 1977; Foreman et al.,
1979) and has proven to be an effective preliminary method of
25


26
describing the spatial location of the interneuronal pools, activated
by a particular stimulus. The most prominent field potential
recordable from the dorsal horn is the result of ionic movements at
excitatory axo-dendritic and axo-somatic synapses (Willis, 1980).
When the synapses are activated, positive ionic current leaves the
extracellular space at the synapses (sinks) and reappears along the
axons of the interneurons (sources). Because the majority of dorsal
horn interneurons ventral to the substantia gelatinosa have ventrally
projecting axons (Matsushita, 1969, 1970), the dorsal horn takes on a
net negative charge and the ventral horn takes on a net positive
charge. The negativity of shortest latency is focussed in a region
which contains a high density of interneurons monosynaptically
excited by primary afferents (Yates et al., 1982).
Methods and Materials
Surgical Preparation of Animals
Data were recorded from 25 adult cats of either sex.
Intraspinal potential fields were recorded from 7 of these animals;
the minimum times between the arrival of the primary afferent volleys
along muscle, cutaneous and venous afferents and detectable
discharges in the ventral roots were determined in the other 18.
Animals were initially anesthetized with 3% Fluothane (halothane,
Ayerst laboratories) vaporized in a mixture of nitrous oxide and
oxygen. A tracheostomy was performed and either of the common
carotid arteries was cannulated; the other common carotid artery was
ligated. The tracheostomy allowed for artificial ventilation with a


27
Harvard respiration pump; cannulation of the carotid allowed for
measurement of arterial blood pressure. The left cephalic vein was
also cannulated to allow the intravenous administration of drugs and
fluids. Animals were rendered decerebrate at the midcollicular level
by transection of the brainstem and aspiration of the portions of the
brain rostral to this level. Following decerebration, anesthesia was
discontinued; this occurred at least 4 hours prior to the beginning
of the recording session. The spinal cord was also transected at
T10. Animals were immobilized in a spinal unit (David Kopf);
paralysis was induced through intravenous injection of Flaxedil
(gallamine triethiodide, Davis-Geck, 2 mg/kg). Flaxedil was
readministered when any muscle contractions were noted. Spinal cord
segments L3-S1 were exposed by laminectomy, the dura mater was opened
and pinned away from the cord dorsum, and the exposed tissues were
covered with warmed mineral oil. Body and oil pool temperatures were
maintained at 37 degrees C using a heating pad and a heat lamp.
The skin of either of the hindlimbs was opened dorsally to
expose a segment of the femoral-saphenous vein. A 10-15 mm segment
of the vein was separated from surrounding connective tissue and
gently placed upon a silver bipolar stimulating electrode. A plastic
membrane was placed beneath the stimulated vein segment to prevent
accidental contact of the electrode with surrounding tissues. In
those experiments in which the minimum time between the arrival of
the primary afferent volley and a detectable discharge in the ventral
roots was determined, a segment of one or more of the following
nerves was also prepared, in a manner similar to that used to prepare


28
the vein, for bipolar electrical stimulation: the sural nerve,
posterior tibial nerve, gastrocnemius nerve, deep peroneal nerve,
plantaris nerve or deep posterior nerve (branch of the tibial
innervating foot musculature). Bipolar stimuli were sometimes also
applied to the skin overlying the femoral-saphenous vein. In
addition, the L6 and L7 ventral roots were cut at dural entry and
were placed across bipolar hook recording electrodes.
Stimulation Procedures
Bipolar electrical stimulation was used in these experiments to
excite the femoral-saphenous venous afferents. A number of previous
studies have also utilized electrical stimulation to activate the
venous afferents (Thompson and Barnes, 1979; Thompson and Yates,
1983, 1984, 1986; Thompson et al., 1982); these studies provided
considerable evidence that this stimulation only excited afferent
fibers arising in the vein wall. Application of ligatures to
portions of the vein immediately proximal and distal to the
stimulating electrode abolished the evoked potentials elicited by
passing current along that vein segment (Thompson and Yates, 1986).
This result was apparently due to the blockade of impulse conduction
in afferents coursing along the vein wall (Gelfan and Tarlov, 1956).
Stimulation of isolated segments of the femoral-saphenous vein
elicited a unique spinal reflex; the stimulation produced the
simultaneous excitation of hindlimb flexor and extensor muscles
acting on the thigh, crus and foot (Thompson et al., 1982).
Activation of cutaneous afferents produces the reciprocal excitation
and inhibition of flexors and extensors; activation of large muscle


29
afferents also elicits the reciprocal excitation and inhibition of
hindlimb muscles (see Chapter I). It is thus likely that the
afferents which were excited by the stimulation of isolated segments
of the femoral-saphenous vein belonged to a unique class arising in
the vein wall and were not fibers of passage coursing along the vein
but arising in muscle or skin. The studies presented in the last
chapter involved the application of HRP to isolated vein segments;
the procedures used to isolate portions of the femoral-saphenous vein
for exposure to this tracer and for electrical stimulation are
similar. The results presented in Chapter II suggested that only
afferents arising in the vein were labeled by the application of HRP
to the isolated vein segments; thus, muscle and cutaneous afferents
apparently were not coursing along these segments. It is clear that
electrical stimulation is a technique which allows for the selective
activation of femoral-saphenous venous afferent fibers.
During experiments in which intracord field potentials were
mapped, the prepared segment of the femoral-saphenous vein was
stimulated using single-shock square-wave pulses of current, 0.2 msec
in duration, repeated at a frequency of 1 Hz. Stimuli were delivered
by a digital stimulator (Frederick Haer Pulsar 6i). Stimuli 3 times
threshold for eliciting field potentials recordable from the cord
dorsum were used to excite the afferent fibers. This stimulus
intensity was chosen because previous studies had shown it sufficient
to elicit compound action potentials recordable from the dorsal roots
with A-beta components of maximal amplitude. The A-delta and C
fibers shown in Chapter II to also arise from the femoral-saphenous


30
vein were not excited by this stimulus intensity (Thompson and Yates,
1986); thus, a very synchronous spinal input was elicited. During
the experiments in which the minimal times required for activation of
spinal motoneurons were measured, the femoral-saphenous vein was
stimulated at an intensity 2-3 times that necessary to evoke field
potentials recordable from the cord dorsum. The hindlimb skin and
the sural nerve were stimulated at twice cord dorsum threshold.
Stimuli applied to muscle nerves were graded to evoke only
monosynaptic reflexes recordable from the ventral roots or to elicit
both monosynaptic and polysynaptic reflexes.
Recording Procedures
Intraspinal potential fields. Intraspinal field potentials
were recorded using tungsten microelectrodes insulated with epoxy or
glass except for a 10-20 micrometer tip which had initial impedances
of 0.5-4.0 megohms. Potentials were recorded at 0.2 mm intervals
from the cord dorsum to 4.4 mm in depth; electrodes were positioned
using a hydraulic microdrive (Narishige). Five tracks were conducted
across the L6 cord segment. Penetrations were optimally made at
midline and 0.5, 1.0, 1.5 and 2.0 mm lateral to midline; slight
alterations in the choice of penetration site were often necessary to
avoid puncturing large blood vessels. Potentials were referenced to
animal ground; the reference electrode was positioned in the wound
margin adjacent to
the
cord.
Potentials were
led into A.C.
preamplifiers (Grass
P511)
with a
time constant
of 250 msec and
subsequently into a
tape
recorder
(Crown-Vetter
model A) which
contained preamplifiers with a bandwidth of D.C. to 15 kHz. The


31
potentials were displayed on an oscilloscope and were averaged by an
Ortec computer. Records of averaged potentials were made with an X-Y
plotter (Hewlett-Packard).
A radiofrequency lesion was placed at the bottom of each track
to facilitate the histological localization of the track. The depths
of lesions were also analyzed as a control for tissue shrinkage by
fixative and other factors (such as cord dimpling) which might cause
depths measured from histological material to deviate from those
measurements read from the vernier of the hydraulic microdrive.
Cord dorsum potentials. Cord dorsum potentials were
recorded from L6 using silver ball electrodes and were referenced to
animal ground. The potentials were led into A.C. preamplifiers, etc,
as for potentials recorded from microelectrodes. Cord dorsum
potentials were recorded in these experiments for two reasons. The
measurement of cord dorsum potentials provided a means to determine
the latency at which input along the fastest-conducting afferents
arrived at the cord; several studies have shown that the initial
triphasic spike of the cord dorsum potential is a compound action
potential propagating through large primary afferent fibers (Austin
and McCouch, 1955; Howland et al., 1955). In addition, the amplitude
of cord dorsum potentials is known to vary in proportion to spinal
cord excitability (Bernhard and Roll, 1953; Gelfan and Tarlov, 1956;
Molt et al., 1978). Thus, the measurement of cord dorsum potentials
provided a control to assure that the level of spinal cord
excitability remained constant throughout the recording session and
that little cord damage was produced by electrode penetrations in


32
those experiments in which intraspinal field potentials were mapped.
Data recorded from animals in which the cord dorsum potentials varied
in amplitude during the recording session were not analyzed.
Compound action potentials. Compound action potentials were
recorded from the L6 and L7 ventral roots during those experiments in
which the minimal time required for an afferent input to activate
motoneurons was determined. The bipolar recording electrodes were
positioned in close proximity to the spinal cord in these
experiments. Compound action potentials were led into A.C.
preamplifiers, etc, as described above.
Histological Procedures
At the conclusion of intraspinal field potential mappings,
animals were perfused through the heart with 1 1 of heparinized
saline followed by 1.5 1 of 1.25% glutaraldehyde/1% paraformaldehyde
fixative. The L6 segment was removed, quick-embedded in
gelatin-albumin, cut into 40 micrometer sections with a freezing
microtome and stained with cresyl violet. Most electrode tracks
could easily be reconstructed from histological material; only data
recorded along reconstructed tracks were considered for analysis. If
fewer than 4 tracks were reconstructed in a particular animal, data
recorded from that animal were not analyzed. Identification of the
laminae in which the recording sites were located was done through
microscopic examination of histological material and by comparison
with published accounts of laminar organization (Rexed, 1952, 1954).


33
Results
Figure 3-1 shows the results of experiments in which minimal
times required for activation of spinal motoneurons by inputs from
muscle, skin, and the femoral-saphenous vein were examined. A
determination of this minimal time for femoral-saphenous venous
afferent input is illustrated in part "A" of the figure. This
latency represents a combination of conduction time along the
intraspinal portions of primary afferents, synaptic delay times, and
conduction time along the processes of spinal neurons. However, a
comparison of the central delay following venous afferent stimulation
with the delay following stimulation of cutaneous and muscle
afferents, the synaptology of which is well established (Brown,
1981), provides an estimate of the minimal number of synapses
interposed between the primary afferents and alpha-motoneurons.
The minimal times between the arrival of the primary afferent
volley and a detectable discharge in the ventral roots are indicated
in part "B" of Figure 3-1. The abscissa shows the afferent type
stimulated. Data regarding muscle afferents represent the pooling of
delays measured following stimulation of the following nerves:
posterior tibial, gastrocnemius, deep peroneal, deep posterior or
plantaris. The latencies from triphasic spike to onset of both
monosynaptic and polysynaptic reflexes recorded from the ventral root
are shown. Data regarding cutaneous afferents represent the pooling
of delays measured following stimulation of the sural nerve or the
skin overlying the femoral-saphenous vein. Latencies reported for
muscle and cutaneous input reflect the processing time by the input


Figure 3-1. Minimal spinal cord processing time for inputs from
muscle, skin and the femoral-saphenous vein. (A) The method
used to determine minimal spinal cord processing time. (B) A
comparison of these latencies. Abbreviations: CD, cord dorsum
potential; VR, potential recorded from the ventral roots; TPS,
triphasic spike.


MINIMAL PROCESSING TIME (MSEC)
35
V. A. V. A. MUSCLE CUTAN. MUSCLE
L6-L7 L6'L6 (polysyn) (monosyn)


36
cord segment for the stimulated afferents (L7 for the stimulated
muscle nerves and sural nerve and L6 for the skin overlying the
femoral-saphenous vein). Two central delay times following
stimulation of the femoral-saphenous vein are reported: the latency
separating the L6 triphasic spike and activity recorded from L6
ventral roots as well as the latency separating the L6 triphasic
spike and activity recorded from the L7 ventral roots. Thick bars
indicate mean minimal spinal cord processing time; thin bars indicate
the standard error. Stars indicate values shown to differ
significantly from all others by an analysis of variance procedure in
combination with either a Waller-Duncan K-ratio T test or Duncan's
multiple range test (alpha=0.05). The delay separating the triphasic
spike elicited by muscle afferent stimulation and the monosynaptic
reflex recorded from the ventral root was significantly shorter than
all others; the delay separating the L6 triphasic spike elicited by
femoral-saphenous venous afferent stimulation and ventral root
activity recorded from L7 was significantly longer than all others.
Figure 3-2 shows a series of waveforms recorded following
femoral-saphenous venous afferent stimulation at various depths along
a typical raicroelectrode track through L6. The potentials were
recorded as a series of negativities near the cord dorsum or in the
dorsal gray matter (waveforms A-D). As the microelectrode tip
advanced into the ventral horn, the negative waves decreased in
amplitude and eventually reversed in polarity (waveforms E and F).
Figure 3-3 shows the focus of the short latency negative waves
elicited by stimulation of the femoral-saphenous vein. The dots


electrode track through L6
Figure 3-2. Potentials recorded along a typical
following stimulation of the femoral-saphenous vein.


lmm
L
10 msec


Figure 3-3. The location of the focus of the short latency negative waves
elicited by stimulation of the femoral-saphenous vein. Symbols are explained in
the text.


2.6 MSEC
1 MM


41
indicate points where maximal-amplitude potentials were recordable;
the lines surrounding the dots indicate the area in which potentials
80% or more of maximal could be recorded; the arrows at the top of
both diagrams indicate the location and orientation of electrode
tracks used to collect data; the time over each diagram indicates
latency from the onset of the triphasic spike of the cord dorsum
potential. In most experiments (5 of 7), the initial negativities
were focussed in the neck of the dorsal horn; this region corresponds
to Rexed's lamina V (Rexed, 1952, 1954). In 4 of 7 experiments, the
initial negativities were also focussed, along the medial-to-lateral
axis, roughly in the center of the gray matter; however, foci both
medial and lateral to this point were also noted. It is likely that
the inconsistencies in results observed in a few experiments were due
more to limitations in the field potential mapping technique than to
physiological variability between animals. If a microelectrode
failed to pass directly through a potential generator during a
recording session, the spatial location of that potential generator
could appear displaced from its actual location.
Discussion
The latency separating evoked activity recordable from the
dorsal and ventral roots of L6, the input segment for
femoral-saphenous venous afferents, following stimulation of the
femoral-saphenous vein (4.4 msec) was approximately the same as the
latency noted following stimulation of cutaneous afferents (4.3
msec). Minimal spinal cord processing time, estimated in the same
way for inputs from muscle which reach alpha-motoneurons through


42
polysynaptic circuits, was only slightly shorter (3.8 msec). It is
known that the fastest circuits which link cutaneous inputs and
muscle inputs with interneurons and then motoneurons are di- and tri-
synaptic (Lloyd, 1943b, 1943c). These data are consistent with a di-
or tri- synaptic circuit being the shortest pathway between primary
femoral-saphenous venous afferents and alpha-motoneurons.
Following electrical stimulation of the femoral-saphenous vein,
negative waves of the cord dorsum potential can be recorded in spinal
animals for 20 msec or more (Thompson and Yates, 1986). These waves
reflect the activation of dorsal horn interneurons (Coombs et al.,
1956; Fernandez de Molina and Gray, 1957; Willis et al., 1973; Fu et
al., 1974; Beall et al., 1977; Foreman et al., 1979; Willis, 1980;
Yates et al., 1982). In addition, maximal ventral root action
potentials elicited in spinal animals by femoral-saphenous venous
afferent stimulation have a duration of 20-25 msec (Thompson et al.,
1982). Thus, although relatively short di- and tri- synaptic
circuits interconnect venous afferents and motoneurons, much higher
order circuits also appear to exist.
The minimal latency separating L6 dorsal root activity and L7
ventral root activity elicited by FVA stimulation was significantly
longer (by at least 1.7 msec) than other processing times estimated.
Increased intraspinal conduction distances can account for only a
portion of this longer latency; the conduction time along primary
femoral-saphenous venous afferents from L6 to L7 is less than 1 msec
(Thompson and Yates, 1986). It is likely that at least one
additional synapse is interposed between femoral-saphenous venous


43
afferents and L7 motoneurons than is interposed between the afferents
and L6 motoneurons.
A distribution of intracord potential fields similar to that
produced by stimulation of the femoral-saphenous vein (negativities
in the dorsal horn and positivities in the ventral horn) is also
elicited by stimulation of other afferent types. Previous studies
have shown that the negative waves of shortest latency in the dorsal
horn can be attributed to monosynaptic excitation of interneurons by
primary afferents (Coombs et al., 1956; Fernandez de Molina and Gray,
1957; Willis et al., 1973; Fu et al., 1974; Beall et al., 1977;
Foreman et al., 1979; Willis, 1980; Yates et al., 1982). Since the
largest negative waves of shortest latency were recorded from Rexed's
lamina V, it appears that most of the first interneurons excited by
the large venous afferents are found in this region.


CHAPTER IV
PHYSIOLOGICAL PROPERTIES OF SINGLE SPINAL CORD NEURONS
ACTIVATED BY STIMULATION OF THE FEMORAL-SAPHENOUS VEIN
Introduction
The data presented in Chapter III suggested that at least one or
two interneurons are interposed between large (A-beta) primary
femoral-saphenous venous afferents and motoneurons. It was also
concluded that the first interneurons activated by these large
afferents are located in Rexed's lamina V. The study discussed in
this chapter examined the responses of single spinal cord neurons
following stimulation of the femoral-saphenous vein. In addition,
the patterns of convergence of inputs from muscle and skin on the
venous afferent-activated interneurons were examined. This study was
conducted to show whether venous afferent inputs were highly
processed by spinal interneurons and, if so, whether these neurons
were modality-specific.
Methods and Materials
Surgical Preparation of Animals
Experiments were performed on 19 unanesthetized, decerebrate
cats with spinal cords transected at T12 and 5 intact cats
anesthetized using alpha-chloralose. The two preparations were used
because a comparison of the data obtained from each type would
suggest whether an elicited input had strong or weak effects on the
excitabilities of the neurons being studied. It is known that the
44


45
excitabilities of spinal interneurons in intact animals anesthetized
using alpha-chloralose are lower than in decerebrate-spinal
preparations. The lower excitabilities in these anesthetized
preparations are due to the influences of descending supraspinal
inputs to the spinal interneurons as well as to direct effects of the
anesthetic on the neurons (Alvord and Fuortes, 1954; Balis and
Monroe, 1964; Brown, 1981). If spinal neurons are driven in both
decerebrate and anesthetized preparations by a particular input, it
is likely that the input has powerful effects on these neurons.
However, if spinal neurons are activated by the input in
decerebrate-spinal preparations, but not anesthetized preparations,
it is likely that the input has only weak influences on the
excitabilities of these neurons.
Anesthesia was induced in the group of animals which would not
be decerebrated using the intravenous injection of alpha-chloralose
(Merck, 75 mg/kg). Maintenance doses of the drug (25 mg/kg) were
routinely delivered every four hours. However, the electrocardiogram
and arterial blood pressure were continuously monitored during the
experiment; if either the heart rate or blood pressure increased,
indicating that the animal was recovering from anesthesia, additional
alpha-chloralose was administered until both of these measurements
stabilized at their original level. The other animals were rendered
decerebrate using the procedures described in Chapter III.
Most of the surgical procedures used to prepare cats for single
unit recordings were similar to those described in Chapter III for


46
preparing animals for the recording of field potentials. Several
additional procedures were also used to provide better mechanical
stability for recording. A dural hammock was formed by placing
stitches through the dura and then by suspending small weights from
the threads. In this way the cord was lifted away from underlying
vertebral bodies and pulsating blood vessels. In many animals the
bladder was catheterized so that it would remain empty; this helped
to stabilize the blood pressure and prevented blood pressure surges
due to a full bladder. A bilateral pneumothorax was also used to
minimize movements of the spinal cord during respiration. In
addition to these procedures for increasing mechanical stability,
holes were made in the arachnoid and pia overlying the L6 spinal cord
using sharp forceps. This procedure permitted the insertion of
microelectrodes into the cord without breaking the tips.
Stimulation Procedures
Bipolar electrical stimulation was used to activate
femoral-saphenous venous afferents, cutaneous afferents coursing in
the sural nerve and muscle afferents running in the posterior tibial,
deep posterior, lateral gastrocnemius and hamstring nerves.
Square-wave pulses of current, 0.2 msec in duration, were used to
excite afferent fibers. While data were being collected, stimulus
frequencies of 1 Hz were used. However, stimulation frequencies up
to 100 Hz were used to differentiate afferent responses from
responses generated by interneurons (see below). Throughout the time
microelectrode penetrations were being done, the femoral-saphenous
vein was continuously stimulated at an intensity 3 times that


47
necessary to evoke a field potential recordable from the cord dorsum.
As discussed in Chapter III, this stimulus intensity provides a
maximal A-beta volley to the spinal cord but does not excite the
A-delta and C afferents. High current intensities were not used in
an attempt to recruit the small-diameter venous afferents due to
concerns regarding stimulus spread. Nerves were stimulated at
intensities up to 10 times the threshold for eliciting a cord dorsum
potential; the minimal current intensity for affecting the
excitability of a unit was also determined.
Recording Procedures
Intracellular and extracellular single unit recordings were done
using glass micropipette electrodes filled with 3 M KC1 or 8% HRP in
tris-buffered 0.15 M KC1 (pH 7.3). Electrodes filled with 3 M KC1
had initial impedances of 15-30 megohms; electrodes filled with the
HRP solution
had
initial impedances
of
30-90
megohms.
Microelectrodes
were
positioned using
a hydraulic
microdrive
(Narishige). The
L6
dorsal columns ipsilateral
to the
stimulated
vein and nerve segments were usually impaled by the electrodes;
however, a few penetrations were also made through the L6 dorsal root
entry zone. The microdrive was angled sharply towards the midline
during these latter penetrations. When electrodes filled with 3 M
KC1 were used to record responses, only units located at 700-1500
micrometers depth were studied. At these depths, the electrode tips
should have remained in the dorsal horn or in the most superficial
portion of lamina VII (Rexed, 1952, 1954); thus, no motoneurons
should have been encountered. In addition, the units were tested for


48
their responses to high frequency stimulation; it is known that
action potentials recorded from primary afferents show constant
latency with high stimulation frequencies, whereas the latencies of
responses recorded from interneurons are altered by high frequency
stimulation (Yates et al., 1982). If responses recorded from a unit
were invariant in latency with 100 Hz stimulation, these responses
were not analyzed. The use of electrodes filled with the HRP
solution allowed recording sites to be marked either by the
intracellular staining of the characterized neuron or by the
extracellular iontophoresis of the tracer; the procedures used to
reconstruct recording sites are discussed in Chapter V. Only data
recorded from units located outside of lamina IX were analyzed.
All potentials were referenced to animal ground; the reference
electrode was located in the wound margin adjacent to L6. Potentials
were led into a microelectrode pre-amplifier (Winston model 1090A)
and subsequently into an A.C. amplifier (Grass P511) with a time
constant of 250 msec, tape recorder (Vetter model D) with a 10 db
bandwidth of 0-2500 Hz and oscilloscope. The direct output of the
microelectrode pre-amplifier was also led into an oscilloscope and
into the tape recorder. Analysis of data was facilitated by the time
histogram unit of an Ortec computer (model 4621); the latter unit
generated poststimulus time histograms from the data. Hardcopy
records of histograms were generated using an X-Y plotter
(Hewlett-Packard).
At the conclusion of single unit recordings from 11 animals,
small bundles of the L7 dorsal rootlets were cut proximally and


49
placed across a silver bipolar recording electrode. The sural and
posterior tibial nerves were then stimulated using the same current
intensities applied to activate interneurons during the single unit
recordings. Potentials were led into an A.C. amplifier, tape
recorder and oscilloscope. Potentials were also averaged using the
Ortec computer. In addition, the conduction distances from the nerve
stimulation sites to the dorsal root recording site were measured.
These data were used to determine the conduction velocities of the
afferents activated by the minimal current intensities necessary to
alter the excitabilities of interneurons.
Results
Responses of Interneurons Following Stimulation of the Femoral-Saph
enous Venous Afferents
Data were recorded from 81 neurons activated by stimulation of
the femoral-saphenous vein in decerebrate cats; detailed analyses
were made of the data recorded from 66 units. While only the latter
data were useful in quantitative determinations of such parameters as
the onset latency of evoked activity in neurons, etc., all of the
data were useful in determining the patterns of convergence of
femoral-saphenous venous afferent input with inputs from muscle and
skin. In anesthetized animals, data were recorded from 14 units; the
responses of all of these neurons following femoral venous afferent
stimulation were characterized in detail.
Figure 4-1 illustrates the types of responses recorded from
interneurons following femoral-saphenous venous afferent stimulation
at an intensity 3 times that necessary to evoke field potentials


Figure 4-1. The effects of stimulation of the femoral-saphenous vein on the
excitabilities of interneurons. The top traces show 16 superimposed oscilloscope
sweeps. The bottom traces show poststimulus time histograms generated from 64
consecutive sweeps. The stimulus was delivered at the onset of the traces.


Burst
Burst +
Inhibition


52
recordable from the cord dorsum. A burst of action potentials was
recorded from 42 out of 66 (64%) of the venous afferent-driven
neurons following stimulation of the femoral-saphenous vein in
decerebrate animals. The other 24 neurons (36%) responded to vein
stimulation with a combination of one or more bursts of action
potentials and a period during which the firing rate of the unit was
depressed below the spontaneous rate. The neurons studied in cats
anesthetized using alpha-chloralose responded in similar ways to
femoral-saphenous venous afferent stimulation; half of the 14 neurons
characterized were excited by vein stimulation, and the other half
were both excited and inhibited.
The quantitative aspects of the response properties of these
neurons activated by stimulation of the femoral-saphenous vein are
summarized in Table 4-1. The mean spontaneous firing rates of the
neurons were similar in decerebrate and anesthetized animals (15.5
+/- 16.7 spikes per second in the former and 14.3 +/- 20.3 spikes per
second in the latter). Clearly, most of the units examined had
substantial ongoing activity in the absence of stimulation; however,
a number of units showed no recordable spontaneous firing. Twelve of
the units (18%) characterized in decerebrate cats were silent in the
absence of stimulation; 5 of the units (36%) in anesthetized animals
exhibited no spontaneous firing. The occurrence of these silent
units is important, since a unit which does not fire spontaneously
will be classified as only being excited by elicited inputs, whether
or not it receives inhibitory inputs. It is only possible to show
that the firing rate of a unit decreases below the spontaneous level


53
Table 4-1. Responses of spinal interneurons to electrical stimulation
of the femoral-saphenous venous afferents, posterior tibial nerve and
sural nerve. All confidence intervals represent mean +/- standard
deviation. Onset latencies were measured from the onset of the
triphasic spike of the cord dorsum potential. All means except those
for spontaneous firing rate are in msec; mean spontaneous firing rates
are in spikes per sec.
Decerebrate Animals
'Burst'
'Burst + Inhi-
Both Types
Units
bition' Units
Combined
Spontaneous
13.5 +/- 17.2
19.1 +/- 15.6
15.5 +/- 16.7
firing rate
Onset latency of
(n = 42)
(n = 24)
(n = 66)
activity from
13.5 +/- 19.6a
5.6 +/- 4.9a
10.6 +/- 16.3
vein stimula
tion
(n = 42)
(n = 24)
(n = 66)
Duration unit
was excited by
34.0 +/- 65.0
59.7 +/- 82.7
43.3 +/- 72.4
vein stimula
tion
Duration unit
was inhibited
(n = 42)
(n = 24)
41.3 +/- 26.4
(n = 66)
by vein stimu
lation
(n = 24)
Total duration
excitability
was altered by
34.0 +/- 65.0a
101 +/- 88.6a
58.4 +/- 80.6
vein stimula
tion
(n = 42)
(n = 24)
(n = 66)
Total duration
excitability
was altered by
102 +/- 186
89.8 +/- 73.7
95.8 +/- 138
tibial nerve
stimulation
(n = 11)
(n = 11)
(n = 22)
Total duration
excitability
was altered by
42.6 +/- 48.0
45.0 +/- 30.4
43.7 +/- 40.0
sural nerve
stimulation
(n = 10)
(n = 8)
(n = 18)
^Values for 'burst' and 'burst + inhibition' units were significantly
different by Student's t-test (p < 0.05).


54
Table 4-1continued.
'Burst'
Units
Anesthetized Animals
'Burst + Inhi- Both Types
bition' Units Combined
4.0 +/- 9.6
(n = 7)
24.6 +/- 23.4
(n = 7)
14.3 +/- 20.3
(n = 14)
6.8 +/- 6.2
(n = 7)
4.0 +/- 4.2
(n = 7)
5.4 +/- 5.3
(n = 14)
17.4 +/- 19.8
(n = 7)
18.5 +/- 20.8
(n = 7)
43.6 +/- 63.3
(n = 7)
17.9 +/- 19.4
(n = 14)
17.4 +/- 19.8
(n = 7)
62.1 +/- 66.4
(n = 7)
38.5 +/- 51.9
(n = 14)
18.6
(n = 1)
75.1 +/- 81.1
(n = 4)
63.8 +/- 64.6
(n = 5)
5.1 +/- 5.9
(n = 3)
81.5 +/- 68.6
(n = 5)
52.9 +/- 65.3
(n = 8)


55
following stimulation if the unit fires spontaneously. Thus, some of
the units with a low spontaneous rate, classified in this study as
being only excited by venous afferent stimulation, may indeed also
have been inhibited by this input.
Table 4-1 also shows the onset latencies of the responses
elicited by venous afferent stimulation and the durations that the
excitabilities of the units were altered by this stimulation. The
variances in these values were enormous. The changes in the
excitabilities of neurons elicited by stimulation of the
femoral-saphenous vein had an average onset latency of 10.6 +/- 16.3
msec following the onset of the triphasic spike of the cord dorsum
potential in decerebrate animals and 5.4 +/- 5.3 msec in anesthetized
animals. These means were shown to be significantly different by
Student's t-test (p < 0.039). The triphasic spike is a compound
action potential propagating through the largest active primary
afferent fibers (Austin and McCouch, 1955; Howland et al., 1955;
Yates et al., 1982); thus, the onset latency measured from the
triphasic spike represented the onset latency of activity following
the arrival of femoral-saphenous venous afferent input at the cord.
In decerebrate cats, there was also a significant difference (p <
0.016) in the onset latency of elicited activity in the units excited
by venous afferent stimulation and the units both excited and
inhibited by this input; the mean latency was 13.5 +/- 19.6 msec in
the former and 5.6 +/- 4.9 msec in the latter. A similar significant
difference in the onset latency of activity elicited by venous
afferent stimulation was not shown between units of the two types in
anesthetized cats


56
The excitabilities of the units receiving inputs from the
femoral-saphenous vein were modulated for long durations following
vein stimulation. The mean duration the excitabilities of the
neurons were altered by vein stimulation was 58.4 +/- 80.6 msec in
decerebrate animals and 38.5 +/- 51.9 msec in anesthetized animals.
In decerebrate animals there was also a significant difference in the
duration that units excited by venous afferent stimulation and
neurons both excited and inhibited by this stimulation were affected
by inputs from the femoral-saphenous vein (p < 0.0008).
Convergence of Inputs from the Femoral-Saphenous Vein with Inputs from
Muscle and Skin on Single Spinal Neurons
The effects of stimulation of the posterior tibial nerve, the
largest muscle nerve of the hindlimb, and the sural nerve, the
largest cutaneous nerve of the hindlimb, on the excitabilities of
units receiving input from the femoral-saphenous vein were also
studied. The posterior tibial nerve innervates a number of muscles
of the crus and foot, including the triceps surae, tibialis
posterior, flexor hallucis longus, lumbrical muscles and interosseus
muscles. The sural nerve innervates skin overlying the foot and
ankle (Jefferson, 1954; Crouch, 1969). In decerebrate animals, 78
out of 80 (98%) of the venous afferent-activated units tested could
also be activated by stimulation of the tibial nerve at an intensity
10 times threshold for eliciting a cord dorsum potential; 58 out of
78 (74%) of the tested units were also driven by sural nerve
stimulation at an intensity 10 times that necessary to evoke a cord
dorsum potential; 56 out of 77 (73%) of the neurons were activated by


57
stimulation of both nerves. In anesthetized animals, 9 out of 13
(69%) of the venous afferent-activated units examined could also be
activated by stimulation of the tibial nerve; 10 out of 13 (77%) of
the tested units were also driven by sural nerve stimulation; 7 out
of 13 (54%) could be activated by stimulation of both nerves.
Figure 4-2 shows poststimulus time histograms generated from
data recorded from a single unit following stimulation of the
femoral-saphenous vein, the sural nerve and the tibial nerve. The
activity elicited by tibial nerve stimulation had an average onset
latency of 6.6 +/- 10.3 msec following the triphasic spike produced
by stimulation of this nerve in decerebrate animals and 3.9 +/- 2.8
msec in anesthetized animals. The activity elicited by sural nerve
stimulation had an average onset latency of 8.2 +/- 5.4 msec
following the triphasic spike produced by stimulation of this nerve
in decerebrate animals and 4.4 +/- 4.9 msec in anesthetized animals.
The venous afferent-activated neurons were excited, inhibited or both
excited and inhibited by stimulation of the sural and tibial nerves.
Most of these units (37 out of 54 or 69%) studied in decerebrate
animals were both excited and inhibited by stimulation of the tibial
nerve; 16 units (30%) were excited by tibial nerve stimulation and 1
neuron (2%) was inhibited by stimulation of this nerve. In
contrast, most of these units (22 out of 37 or 59%) were only excited
by sural nerve stimulation in decerebrate cats. Only 13 units (35%)
were both excited and inhibited by sural nerve stimulation; 2 cells
(5%) were inhibited by the stimulation of this nerve. In
anesthetized animals, 40% of the tested units were excited by tibial


Figure 4-2. Poststimulus time histograms showing the effects of
stimulation of the femoral-saphenous vein, posterior tibial nerve
and sural nerve on the excitability of a single spinal neuron. The
histograms were each generated from 64 consecutive sweeps. The
stimulus was delivered at the onset of the traces.


59


60
nerve stimulation, 40% were both excited and inhibited, and 20% were
inhibited. In the same animals, half of the units examined were
excited by sural nerve stimulation, and the other half were both
excited and inhibited by the stimulation of this nerve. There were
no obvious
simple
relationships between how
a
neuron
responded
to
stimulation
of
the
femoral-saphenous
vein and
how it
responded
to
stimulation
of
the
tibial and sural
nerves.
The excitabilities
of
the neurons were modulated for long durations following stimulation
of the sural and tibial nerves; these durations are shown in Table
4-1.
To better determine the patterns of convergence of muscle
afferent input on femoral-saphenous venous afferent-activated spinal
interneurons, the effects of stimulation of 3 muscle nervesthe
hamstring nerve, the lateral gastrocnemius nerve and the deep
posterior nervewere tested on 17 units in decerebrate cats. The
nerves were stimulated using current intensities 10 times that
necessary to evoke a field potential recordable from the cord dorsum.
The hamstring nerve innervates 3 flexors of the thigh, the
semimembranosus, the semitendinosus and the biceps femoris; the
lateral gastrocnemius nerve innervates 2 extensors located in the
crus, the lateral gastrocnemius and the soleus; the deep posterior
nerve innervates a number of muscles located in the foot (Jefferson,
1954; Crouch, 1969). All of the 17 venous afferent-activated units
examined could also be driven by stimulation of the deep posterior or
the lateral gastrocnemius nerve; 12 of the units could also be driven
by stimulation of the hamstring nerve. It was clear that muscle


61
inputs from many parts of the hindlimb and from both flexors and
extensors converged on single interneurons also receiving inputs from
the femoral-saphenous vein.
Following recordings from many interneurons, small bundles of
the L7 dorsal rootlets were cut proximally and placed across a silver
bipolar recording electrode. The tibial and sural nerves were then
exposed to the same current intensities which were necessary to alter
the excitabilities of interneurons. Compound action potentials,
recorded from the dorsal rootlets, elicited by these stimulus
intensities were analyzed to determine the conduction velocities of
the excited fibers. In most cases, low stimulus intensities elicited
A-alpha and A-beta volleys, whereas higher stimulus intensities also
elicited A-delta volleys; the results of a typical study are shown in
Figure 4-3. This figure shows the effects of different stimulus
intensities applied to the sural nerve on the amplitudes of 3
components of the evoked dorsal root action potential. In
decerebrate cats, 18 out of 29 units (62%) for which an analysis was
done were driven by stimulus intensities applied to the tibial nerve
which only elicited A-alpha and A-beta components in the compound
action potentials recorded from the dorsal roots. The other 11 units
(38%) were activated by current intensities that produced A-delta
volleys as well as A-alpha and A-beta volleys. Sixteen out of the 17
units (94%) for which an analysis was done were driven by stimulus
intensities applied to the sural nerve which only elicited A-alpha
and A-beta components in the compound action potentials recorded from
the dorsal roots. Only 1 unit was driven by current intensities that


Figure 4-3. The response amplitude versus stimulus intensity relationship for
cutaneous afferents. (A) The relationship between stimulus intensity and the
peak amplitudes of 3 components of the dorsal root action potential evoked by
sural nerve stimulation. (B) A maximal-amplitude compound action potential; the
waveform represents the average of 64 consecutive sweeps.


An'OOc


64
elicited A-delta as well as A-alpha and A-beta volleys. The results
in anesthetized animals were quite similar. Six out of 7 units (86%)
for which an analysis was done were driven by stimulus intensities
applied to the tibial nerve which only elicited A-alpha and A-beta
components in the compound action potentials recorded from the dorsal
roots. The other unit was activated by current intensities that
produced A-delta volleys as well as A-alpha and A-beta volleys. Two
out of the 3 units for which an analysis was done were driven by
stimulus intensities applied to the sural nerve which only elicited
A-alpha and A-beta components in the compound action potentials
recorded from the dorsal roots. The other unit was driven by current
intensities that elicited A-delta as well as A-alpha and A-beta
volleys. However, at minimal stimulus intensities necessary to evoke
A-delta dorsal root volleys, the A-alpha and A-beta volleys were
often still increasing in amplitude. This was especially true for
dorsal root action potentials elicited by stimulation of the tibial
nerve. Thus, in most cases in which both A-delta volleys and volleys
along larger afferents were elicited by stimulus intensities
minimally necessary to activate a neuron, it was impossible to
conclude which fiber group carried the input which activated the
unit. Nonetheless, it was clear that large muscle and cutaneous
afferents were included in the population which carried inputs to the
spinal cord that altered the excitabilities of neurons also driven by
inputs from the femoral-saphenous vein.


65
Discussion
Venous Afferent Effects on the Excitability of Spinal Neurons
The response properties of the neurons driven by stimulation of
the femoral-saphenous vein were similar to those reported by others
for units activated by electrical stimulation of cutaneous and muscle
afferents. These previous studies have shown that most spinal cord
neurons, especially those in decerebrate preparations, have a
substantial spontaneous firing rate. However, units which do not
fire in the absence of stimulation have also been reported (LeBars et
al., 1975; Light and Durkovic, 1984). In addition, the onset latency
and duration of activity elicited in spinal neurons by venous
afferent stimulation are included in the range of values reported
following electrical stimulation of large fibers in the hindlimb
nerves (Feldman, 1975).
Neurons were either excited or both excited and inhibited by
stimulation of the femoral-saphenous vein. In this study, as well as
in previous studies, spinal interneurons were also shown to be
excited or both excited and inhibited by electrical stimulation of
muscle and cutaneous afferents (Hongo et al., 1966; Hillman and Wall,
1969; Wagman and Price, 1969; Feldman, 1975). Units both excited and
inhibited by venous afferent stimulation are likely to receive inputs
through several separate pathways, some responsible for the
excitation, others for the inhibition. The excitation could either
be due to direct inputs from primary afferents to the unit or to
inputs from higher-order spinal neurons driven by venous afferent
stimulation. The inhibition is also likely due to inputs from


66
venous afferent-driven neurons to the unit; however, the neurons
responsible for the inhibition are likely to be different from the
neurons that are responsible for the excitation. An alternate
possibility is that the inhibition is due to the venous
afferent-induced presynaptic inhibition of the tonically active
afferents responsible for the high spontaneous firing rate observed
in these neurons. However, the presynaptic inhibition of primary
spinal afferents has a time course of 200 msec or more (Eccles et
al., 1954, 1962, 1963a, 1963b; Eccles and Krnjevic, 1959; Wall,
1962). The period of inhibition produced by venous afferent
stimulation in most units had a time course of 50 msec or less; in
many units the duration was less than 20 msec. Thus, it is likely
that the inhibition recorded from most of the units following venous
afferent stimulation is due to postsynaptic inhibition, and not
primary afferent depolarization. However, in the few units that
were inhibited for long durations following venous afferent
stimulation, it is possible that primary afferent depolarization also
contributed to the reduced firing rate. Data recorded from one such
unit are shown in Figure 4-1.
Since units that were both excited and inhibited by venous
afferent inputs were likely to receive these inputs through a number
of pathways, the venous afferent influences on the excitabilities of
these units appear to be powerful. Units that responded in this
manner following venous afferent stimulation were common, suggesting
that venous afferent inputs, in general, have a powerful impact on
the excitabilities of spinal cord neurons. Units both excited and


67
inhibited by venous afferent inputs also probably are involved with
the integration and processing of these inputs, although the purpose
of this integration cannot be determined in studies using electrical
stimulation. In contrast, units that are only excited by an afferent
input are more likely to have a role of relaying the input without
significantly processing it. These latter units may fire for a long
duration after receiving the afferent input, however, thereby
prolonging its effects in the nervous system.
Stimulation of the femoral-saphenous vein had prolonged effects
on the excitabilities of the units examined in both decerebrate and
anesthetized animals. This is further evidence that the inputs
carried by the large femoral-saphenous venous afferents have a potent
influence on the excitabilities of spinal neurons.
Convergence of Inputs on Single Neurons
Stimulation of A-beta femoral-saphenous venous afferents
activated many spinal interneurons that could also be driven by
electrical stimulation of muscle and cutaneous afferents. Previous
studies of the properties of single spinal interneurons that received
noxious inputs from the internal environment showed that widespread
convergence of inputs from muscle and skin occurred on these units as
well (Pomeranz et al., 1968; Hancock et al., 1973, 1975; Gokin et
al., 1977; Guilbaud et al., 1977; Foreman and Weber, 1980; Milne et
al., 1981; Blair et al., 1981, 1984; Cervero, 1982; McMahon and
Morrison, 1982; Rucker and Holloway, 1982; Takahashi and Yokota,
1983; Rucker et al., 1984). Pomeranz et al. (1968, p. 528), in fact,
"failed to detect cells which responded only to visceral afferents"


68
in the thoracic spinal cord. Thus, spinal neurons which processed
venous afferent inputs were similar to those that process noxious
inputs from the viscera, in that both types of neurons received
widespread convergent inputs from muscle and skin.
Comparison of Responses in Decerebrate and Anesthetized Animals
The responses of spinal neurons following venous afferent
stimulation differed somewhat in decerebrate and in anesthetized
preparations. These differences could be explained by a generalized
lower neuronal excitability in the latter animals. The onset latency
of the activity elicited by venous afferent stimulation was
significantly shorter in anesthetized cats than in decerebrate cats.
Inputs to neurons mediated through multi-synaptic connections are
more likely to be affected by a lowered state of excitability than
inputs mediated through connections comprised of only a few synapses.
Thus, neurons activated at long latency would be expected to be more
affected by anesthesia than neurons activated at short latency. The
mean onset latency of elicited activity in the neurons in
anesthetized animals would accordingly be expected to be shorter than
in decerebrate animals.
However, many of the properties of the venous afferent-activated
units were similar in both anesthetized and decerebrate preparations.
In both groups of animals the convergence of venous, muscle and
cutaneous inputs on single neurons was common. This suggests that
muscle and cutaneous inputs have powerful influences on the
excitabilities of these units. It was also common for units to be
both excited and inhibited following venous afferent stimulation in


69
the two types of preparations. As discussed above, it is likely that
these units which responded to inputs from the femoral-saphenous vein
in a complex manner are involved with the integration and processing
of the inputs. Thus, it appears that venous afferent inputs are
highly processed in the spinal cord by interneurons.


CHAPTER V
THE VARIABILITY IN THE RESPONSES OF NEURONS LOCATED IN DIFFERENT
LAMINAE FOLLOWING STIMULATION OF THE FEMORAL-SAPHENOUS VEIN
Introduction
The data presented in Chapter IV suggested that spinal neurons
respond in a wide variety of ways following the electrical activation
of A-beta femoral-saphenous venous afferent fibers. Some neurons
were excited by stimulation of the femoral-saphenous vein; others
were both excited and inhibited. Some units were driven within a few
milliseconds following stimulation of the vein; others were driven
only at long latency. The excitabilities of some neurons were
affected only a few milliseconds by vein stimulation; the
excitabilities of others were modulated for over 100 msec. In
addition, most neurons driven by stimulation of the femoral-saphenous
vein could also be activated by stimulation of muscle and cutaneous
afferents; however, a few units were not activated by the stimulation
of hindlimb nerves. It would be of interest to determine whether
the neurons that responded in similar ways following stimulation of
the femoral-saphenous vein had similar locations in the spinal cord.
The study discussed in this chapter examined this possibility.
Methods and Materials
Experiments were performed on 12 unanesthetized, decerebrate
cats with spinal cords transected at T12 and 5 intact cats
anesthetized using alpha-chloralose. The responses recorded from
70


71
these animals were described in Chapter IV; this chapter will
describe the locations of the units with different response patterns
following stimulation of the femoral-saphenous venous afferents,
tibial nerve and sural nerve. Surgical procedures used to prepare
animals for recordings and the techniques used to record single unit
responses were described in detail in Chapter IV. The data described
in this chapter were recorded using microelectrodes filled with an 8%
solution of HRP in tris-buffered 0.15 M KC1 (pH 7.3).
After a neuron had been characterized extracellularly, an
attempt was made to penetrate the element studied. If the attempt
was successful, HRP was iontophoretically injected with 1-3 nA
positive-going rectangular pulses, 500 msec in duration, at 1 Hz.
Current was injected through the electrode for 7.5-15 min and was
delivered through the companion bridge unit (Winston BR-1) to the
microelectrode pre-amplifier. This bridge unit had been
custom-modified by the manufacturer to make it capable of presenting
high voltages. If the characterized neuron could not be impaled, HRP
was iontophoresed extracellularly using 5 nA pulses for 20-30 min. A
blank electrode was inserted into the opposite side of the cord
parallel to each track which yielded data. Once positioned, the
blank electrode was broken using scissors so that only 1-2 mm
protruded above the surface of the cord. These electrode tips were
left in place through the fixation process; they produced easily
observed tracks in the tissue which were used to help identify
sections from which data were recorded. A rostrocaudal distance of
at least 1 mm separated sites of HRP iontophoresis.


72
After at least an hour following the last marking attempt, the
animal was perfused through the carotid artery with 1 1 of
heparinized saline containing 0.1% sodium nitrite followed by 2 1 of
2.5% glutaraldehyde/1.2% paraformaldehyde fixative. The L6 cord was
removed and stored overnight in the fixative. The tissue was cut
into sections 50 micrometers thick using a vibratome (Oxford). The
sections were then incubated for 30 min at room temperature in a
solution comprised of 0.05% 3, 3* diaminobenzidine tetrahydrochloride
and 0.1% hydrogen peroxide in 0.1 M tris buffer at pH 7.6. The
sections were then rinsed in 0.1 M phosphate buffer (pH 7.4), mounted
onto slides, dried, counterstained using 1% neutral red, dehydrated,
cleared in xylene and coverslipped. Two-dimensional camera lucida
reconstructions were made of stained neurons and extracellular
recording sites.
Pugh and Stern (1984) have shown that HRP can be very reliably
used to label extracellular single unit recording sites. Examples of
histologically reconstructed recording sites are shown in Figure 5-1.
Injection site diameters ranged from 50 to 250 micrometers. In
addition to the visible brown reaction product in the extracellular
space, internalization of HRP by neurons provided positive
identification of the center of the injection site which was readily
distinguishable from damaged vascular or neural elements mechanically
produced by the electrode.
Results
Fifty sites at which responses were recorded from venous
afferent-activated neurons were reconstructed; 36 of these


Figure 5-1. Examples of recording sites marked through the extracellular
iontophoresis of HRP. The injection sites were easily identified by the presence
of brown reaction product in the extracellular space. Most of the injection
sites were light in color towards the borders, but were much darker towards the
center. Neuronal elements in the center of the injection site typically also
contained HRP reaction product. The region of the injection site which was
darkest in color, and which contained neurons that had internalized HRP, was
assumed to be the location from which extracellular responses had been recorded.
Calibration bars represent 50 micrometers.


74


75
reconstructed recording sites were in decerebrate animals and the
other 14 were in anesthetized animals. The locations of the
recording sites are shown in Figure 5-2. In decerebrate cats,
neuronal activity elicited by stimulation of the femoral-saphenous
vein was recorded from Rexed's laminae IV-VIII and X; in anesthetized
cats, venous afferent-elicited neuronal activity was recorded from
laminae V and VI and from the dorsal portion of lamina VII. Most of
the recording sites were located either in lamina V (11 out of 50) or
in the most superficial portion of lamina VII, dorsal to the
dorsalmost border of lamina VIII (15 out of 50).
Units located in the different laminae had similar spontaneous
firing rates; these values are shown in Table 5-1. However, the
units which were excited by venous afferent stimulation and the units
both excited and inhibited by vein stimulation were highest in
density in different regions of spinal gray matter. The locations of
the units of the two types are shown in Figure 5-3. Most of the
units in the dorsal horn (12 out of 20 or 60%) were both excited and
inhibited by stimulation of the vein. In contrast, most of the units
in the ventral horn (22 out of 30 or 73%) responded to venous
afferent stimulation with only a burst of action potentials. Only 2
units located deeper than the dorsalmost border of lamina VIII were
both excited and inhibited by venous afferent stimulation; clearly,
units of this latter type were rare deep in the ventral horn.
An investigation was also done to determine the locations of the
neurons driven at monosynaptic latencies by femoral-saphenous venous
afferents. The cord dorsum potential elicited by venous afferent


Figure 5-2. Locations of sites at which responses were recorded from venous
afferent-activated interneurons.


* *>
- DECEREBRATE ANIMALS
- ANESTHETIZED ANIMALS
IX


78
Table 5-1. Spontaneous firing rates of neurons located in different
laminae. Confidence intervals represent mean +/- standard deviation.
All spontaneous firing rates are in spikes per second.
Lamina of Mean Spontaneous Range of Spontaneous
Rexed
n
Firing
Rate
Firing Rate
IV
2
17.5 +/-
21.1
2.5 32.4
V
11
8.8 +/-
11.7
0.0 33.3
VI
7
15.6 +/-
12.9
1.1 37.2
VII
22
20.8 +/-
21.0
0.0 60.0
VIII
5
9.9 +/-
9.6
0.0 20.8
X
2
5.0 +/-
7.1
0.0 10.0


Figure 5-3. Locations of units excited or both excited and inhibited by
stimulation of femoral-saphenous venous afferents.


80


81
stimulation was used to estimate this latency at which units were
first activated by inputs from the vein. The cord dorsum potential
is comprised of an initial triphasic spike, a series of negative
waves and a slow positive wave. The triphasic spike is a compound
action potential reflecting activity in the largest afferent fibers
excited by the stimulus, the negative waves are generated by dorsal
horn interneurons, and the positive wave is produced by the
presynaptic inhibition of afferent fibers (Austin and McCouch, 1955;
Howland et al., 1955; Eccles et al., 1963a, 1963b; Willis, 1980;
Yates et al., 1982). Thus, the latency of the first negative wave of
the cord dorsum potential corresponds to the latency at which
interneurons are being directly excited by large primary afferent
fibers. Neurons excited at the same latency as that of the peak of
the first negative wave of the cord dorsum potential, or at shorter
latencies, were assumed to receive monosynaptic, or at least
relatively direct, inputs from from primary femoral-saphenous venous
afferents. Figure 5-4 illustrates these procedures used to determine
which units received direct inputs from the venous afferents. The
average latency of the peak of the first negative wave of the cord
dorsum potential was 2.67 +/- 0.71 msec from the onset of the
triphasic spike in decerebrate animals and 2.06 +/- 0.85 msec in
anesthetized animals.
Based on these criteria, 8 of the 50 units with recording sites
reconstructed were classified as being directly excited by the
primary venous afferents; 4 of these units were in anesthetized cats,
and the other 4 were in decerebrate cats. The locations of these


Figure 5-4. Procedures used to determine whether a unit is activated at
monosynaptic latencies by primary femoral-saphenous venous afferents. The
lefthand portion of the figure compares the latency of the peak of the first
negative wave of the venous afferent-elicited cord dorsum potential with the
onset latency of activity evoked in a unit. The cord dorsum potential is the
average of 32 consecutive sweeps; unit activity is shown by a poststimulus time
histogram generated from 64 consecutive sweeps. Further responses recorded from
the same unit are shown in the lefthand portion of Figure 4-1. The first
negative wave of the cord dorsum potential is preceded by an intramedullary
spike, which is produced by action potentials propagating into the afferent
termini (Austin and McCouch, 1955). The righthand portion of the figure shows
the location of the unit from which activity was recorded.


Spike
IX


84
units are shown in part "A" of Figure 5-5; most (6 out of 8 or 75%)
were located in lamina V, 1 was located in lamina VI and the other
was located much more ventrally in lamina VII. These units driven at
short latency by venous afferent stimulation in decerebrate animals
had a mean spontaneous firing rate of 8.2 +/- 7.2 spikes per second
and were activated at a mean latency of 1.8 +/- 0.72 msec following
the triphasic spike of the cord dorsum potential. Two of the units
studied in decerebrate animals were excited by venous afferent
stimulation, and the other two were both excited and inhibited. The
excitabilities of these units were modulated for 32.6 +/- 44.2 msec.
The spontaneous firing rates of the units driven by venous afferent
stimulation at short latency in anesthetized animals were much lower
than those in decerebrate cats; the mean rate was only 0.24 +/- 0.28
spikes per second. Two of the units in anesthetized animals were
silent in the absence of stimulation; it was impossible to determine
whether these units were both excited and inhibited by venous
afferent stimulation or only excited. It is only possible to show
that the firing rate of a unit decreases below the spontaneous level
following stimulation if the unit fires spontaneously. The other two
units in anesthetized cats were both excited and inhibited by vein
stimulation. These four units were activated at a mean latency of
1.06 +/- 0.55 msec following the triphasic spike of the cord dorsum
potential; the excitabilities of the units were modulated by venous
afferents for a mean duration of 20.4 +/- 12.8 msec.
The locations of the units driven at longer latencies by the
stimulation of the femoral-saphenous vein are shown in parts "B" to


Figure 5-5. Locations of units which were activated at different minimal
latencies following stimulation of the femoral-saphenous vein. (A) Locations of
units driven at monosynaptic latencies. (B-E) Locations of units activated at
longer latencies. The time over each diagram indicates latency following the
onset of the triphasic spike of the cord dorsum potential.


-DECEREBRATE
812 msec
< 3 msec
c
3-8 msec
-ANESTHETIZED
00
o>


87
"E" of Figure 5-5. Within 3 msec following the triphasic spike of
the cord dorsum potential, but at latencies longer than that of the
peak of the first negative wave of the cord dorsum potential, a few
units located in laminae VI and VII were activated by vein
stimulation. At slightly longer latencies, many units located in the
ventral horn were activated. However, units in the dorsal horn were
also activated at these longer latencies. The onset latency of
neuronal activity produced by venous afferent stimulation ranged to
over 12 msec following the arrival of the afferent volley at the
cord; neurons activated at latencies longer than 12 msec following
the triphasic spike were located both in the dorsal and ventral horn.
Figure 5-6 shows the durations that the excitabilities of units
were modulated following stimulation of the femoral-saphenous vein;
neurons were activated from less than 10 msec to over 100 msec. The
excitabilities of units located in the dorsal horn and in the
superficial portion of lamina VII were modulated for both long and
short durations. However, most of the units located deeper than the
dorsalmost border of lamina VIII were only activated for short
durations (less than 50 msec). The excitabilities of units located
in the dorsal horn and superficial portions of lamina VII of
decerebrate cats were modulated for a mean duration of 74.8 +/- 86.1
msec following vein stimulation, whereas units located deeper were
activated for only an average duration of 26.2 +/- 44.5 msec. These
means were shown to be significantly different by Student's t-test (p
< 0.035).


Figure 5-6. Locations of units activated for different durations
following the stimulation of the femoral-saphenous vein.


89
-DECERE BRATE ^-ANESTHETIZED
<10 msec 10-49 msec
50-100 msec >100 msec


90
As discussed in the last chapter, most of the venous
afferent-driven units could also be activated by stimulation of the
tibial and sural nerves; however, a few units were not driven by the
stimulation of hindlimb nerves. Table 5-2 shows the locations of the
venous afferent-driven neurons that also received convergent inputs
from muscle or skin. In anesthetized animals most of the units which
did not receive convergent inputs from muscle or skin were located in
lamina V, the region which contains most of the neurons driven at
shortest latency by stimulation of the femoral-saphenous vein.
Accordingly, 2 of the 3 units in anesthetized animals that were not
driven by stimulation of the sural nerve were activated at
monosynaptic latencies by stimulation of the femoral-saphenous vein.
The other unit was driven by vein stimulation at a fairly short
latency (3.6 msec following the onset of the triphasic spike of the
cord dorsum potential). Two of the 4 units in anesthetized animals
that were not driven by stimulation of the tibial nerve were
activated at monosynaptic latencies by stimulation of the
femoral-saphenous vein. The other two units were activated at 3.6
and 3.8 msec following the triphasic spike of the cord dorsum
potential. Only one unit studied in decerebrate animals, located in
lamina VII, could neither be activated by stimulation of the sural
nerve or the tibial nerve; this unit was driven at monosynaptic
latencies by stimulation of the femoral-saphenous vein. However, the
other four units characterized in decerebrate cats that were not
driven by sural nerve stimulation were activated by venous afferent
stimulation only at long latency (11.5 +/- 1.9 msec following the


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