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Lateral Asymmetry of H-Reflex Recovery Curves in Cat: Evidence for a Spinal

Motor Asymmetry
Üner Tan a
a
Medical Faculty, Institute of Physiology, Atatürk University, Erzurum, Turkey

Online Publication Date: 01 January 1984

To cite this Article Tan, Üner(1984)'Lateral Asymmetry of H-Reflex Recovery Curves in Cat: Evidence for a Spinal Motor
Asymmetry',International Journal of Neuroscience,24:1,45 — 52
To link to this Article: DOI: 10.3109/00207458409079533
URL: http://dx.doi.org/10.3109/00207458409079533

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 24. pp. 45-52 0 1981 Gordon and Breach Sceince Publishers. lnc.
Printed in the United Kinsdom

LATERAL ASYMMETRY OF H-REFLEX

RECOVERY CURVES IN CAT: EVIDENCE FOR A
SPINAL MOTOR ASYMMETRY
UNER TAN
A tatiirk University, Medical Faculty, Institute of’Physiology, Erzurrrm, Turkey
(Received February 16, 1984)

Recovery curve of the Hoffmann reflex from the left and right sides was studied in lightly anesthetized
cats before and after spinalization. According t o the differences in the H-reflex recovery curves from
the left and right lateral gastrocnemiussoleus nerves, three populations of intact animals were found:
right dominant. with higher recovery curve on the right side (31.8%), left dominant with higher re-
covery curve on the left side (36.4%), and ambilateral with no dominance (31.8%). After spinalization
the dominance remained the same in the two first groups and dominance appeared in 5 out of the
7 cats of the last group. These results provide evidence for a bilateral asymmetry of alpha motoneuron
excitability in cats, which can be used as a model to study the possible mechanisms of spinal motor
asymmetry.

It is well known that the two hemispheres of the human brain ar asymmetric in
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regard to morphological and functional properties. Such a lateralization in the spinal

cord is also conceivable, since the brain and spinal cord are functionally interrelated
systems. Goode et al. (1980) used the Hoffmann reflex (H-reflex) recovery curve to
investigate the lateral differences in the excitability of alpha motoneurons. They
found that the recovery curves measured from the left and right leg differed from each
other significantly in normal human subjects. These authors concluded that this
asymmetry is related in part to cortical laterality. However, no evidence was provided
f0r.a spinal motor lateralization. The aim of the present work was to examine whether
there is a spinal motor lateralization in cats. If so, an animal model would be available
to study the possible mechanisms of the lateral differences in the motoneuronal excita-
bility more extensively than in man. A preliminary account of these experiments has
been presented elsewhere (Tan, 1983).

MATERIALS AND METHODS

The experiments were performed on 20 cats. They were anesthetized with ether during
preparation. The hindlimbs and the hip regions were denervated except the nerves to
the lateral gastrocnemius-soleus (LG-SOL) muscles. The LG-SOL nerves were
severed from their distal ends and prepared for eliciting and recording the H-reflexes.
The first lumbar spinal segment was exposed to spinalize the animal at a later stage of
the experiment. The spinal segments from L6 to SIwere exposed by laminectomy to
ascertain the reflex nature of the H-response by cutting the dorsal roots at the end of
each experiment.

Correspondence Address: Professor Dr. h e r Tan, Atatiirk universitesi, Tip Fakiiltesi, Fizyoloji,
Kiirsiisii Bqkanl, Erzurum, Turkey.
45
 46 is. TAN

A polyethylene cannula was inserted into the right subclavian artery to monitor thc
blood pressure during the experiment. When the blood pressure dropped under
90 mm Hg, Macrodex was infused through another cannula inserted into the right
subclavian vein. Rectal temperature was maintained at about 38°C by means of a
heating device. After preparation was completed, the animal was fixed to a metal
frame and all wound edges along with pressure points were infiltrated with procaine.
Ether was discontinued after these procedures. Anesthesia was then maintained on
i.v. pentobarbitone (0.05 mg/kg/h) infusion and N20/02mixture (60%, 40%).
Stimulating electrodes were placed on the right and left LG-SOL nerves proximally.
Recording electrodes were placed on the distal ends of these nerves. Square wave
pulses werc applied to the exposed nerves in 1Osec intervals. The corresponding
reflex responses were recorded; 16 responses were averaged and stored on a magnetic
tape. The stimulus strength eliciting maximal H-reflex was established. Merent
volleys were simultaneouslyrecorded from the entry zone of dorsal root L7 to establish
whether the left and right sides of the spinal cord receive a symmetric sensory input.
The left and right reflexes were tested alternatively. At least six trials were made in
each animal, which were then spinalized and the same procedure repeated.
The ratio of the amplitude of the test H-reflex (H2)to the amplitude of the H-
reflex elicited by the conditioning shock (HI), expressed as a percentage (H2/H1%),
was estimated. The H-reflex recovery curves, H2/H1%versus stimulus interval, were
plotted for the right and left soleus nerves. All the animals were tested for paw
preference by presenting food. We examined whether the cats required to draw food-
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towards themselves used their right or left forelimb or both of them. Difference bet-
ween the mean amplitudes of unconditioned H-reflex from the right and left soleus
nerves was tested by using z or Student’s t-test. Analysis of variance was used to
determine whether the H-reflex recovery curves from the right and left sides differ
from each other significantly.

RESULTS

Lateral Asymmetry in the H-refix Recovery Curves in Intact and Spinal Cats
Right Dominance. The H-reflex rcovery curve of the right LG-SOL was found to be
significantly higher (ptO.O1) than that of the left LG-SOL in seven cats (31.8%).
The slope of the recovery process was about the same. This statistically significant
difference persisted after spinalization. Four of these animals showed right paw and
the others left paw preference. Figure 1A illustrates the average H-reflex recovery
curves measured from the right (0)and left (0)LG-SOL nerves of seven intact cats.
The recovery curve of the right H-reflex was found to be significantlyhigher than that
of the left H-reflex (F= 1O9.2>Ft;0.01=6.63;df=450). In these experiments the right
H-reflex was signisCantly larger than the left H-reflex estimated by z or StudeDt’s
t-test. As seen in B, a similar difference between the right anci left recovery curves
was observed following spinalization (F=20.9>F=6.85; df= 120).

Lefr dominance. In six intact cats (36.4%), the H-reflex recovery curve measured
from the right LG-SOL nerve was found to be significantly lower than that from the
left LG-SOL nerve. This asymmetry persisted after spinalization (p<O.Ol). These
results were summarized in Figure 2. As seen in A, the average recovery curve from
the right LG-SOL nerve was significantly lower than that from the left LG-SOL nerve
 SPINAL MOTOR ASYMMETRY 47

(F=233.3>Ft,o.ol. =6.63; dJ=405). The mean amplitude of H-reflexes from the

right side was significantly larger than those from the left side in three cats. The
opposite was found in the remaining three cats. Four of these animals showed right
paw preference, and two left paw preference. In Figure 3B, the recovery curves of the
same animals were plotted after spinalization, which did not change the elevation of
left recovery curve (F= 8.3>Ft,o.01= 7.3; df= 36).
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'"1
(%) B.SPINAL
T T
80 -
___ _ - - - -- - I
60- 6,

0 I I I I I 1

0 50 100 150 200 2501ms)300

FIGURE 1 H-reflex recovery curves from the right and left LG-SOL nerves in right dominant
cats. A. intact cats;B. the same animals after spinalization. Abscissa: interval between two successive
shocks (msec). Ordinate: amplitude of the test reflexas percentage of the first one (HZ/HIO/@. Open
and closed circles: average H-reflex recovery curves from the right and left LG-SOLnerves, respec-
tively. Vertical lines correspond to standard deviations. All the data were taken from seven experi-
ments (df=450).
 48 ti. TAN

Ambilaterality in H- Rrfles Recovery Curves it1 Ititact Cats

No significant difference was established between the right and left H-reflex recovery
curves in seven intact cats (31.8%). Three of them showed right paw preference and
the other left paw preference. The mean amplitude of the H-reflexes from the right
side was significantly larger than those from the left side in four cats, and the opposite
was the case in three cats. Figure 3 summarizes the results of recovery curves from
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100-
6.S P I N A L _---I
(%I - - -- ___-----
80-
/
60-

40-

I I I I i

0 50 10 0 150 200 25O(ms) 300

FIGURE 2 H-reflex recovery curves from the right and left LG-SOL nerves in left dominant
animals. Abscissa and ordinate as in the previous figure. Open and filled circles: the average H-
reflex recovery curves-from the right and left LG-SOLnerves, respectively. A. intact cats; B. the
same animals after spinalization.
 SPINAL MOTOR ASYMMETRY 49

seven cnls, indicating no diFference in the right and left recovery curves (F=O. 14<t,o.ol
=3.04; clf=478).

Efect of Spinalization on Ambilaterality of Recovery Curces

The ambilaterality in the H-reflex recovery curves did not change in two cats after
spinalization. However, spinalization revealed a right dominance in three and left
dominance in two ambilateral cats. Figure 4A shows the average H-reflex recovery
curves from the ambilateral cats. There was no significant difference between the
= df= 98).
recovery curves from right and left LG-SOL nerves ( F = 0 . 0 4 ~ F t ~ o . 0 13.94;
After spinalization (B) the H-reflex recovery curve from the right LG-SOL nerve was
significantly higher than that from the left LG-SOL nerve ( E = 26.2>Ft;0.01=7.94;
df=22). In these three animals .the right reflex was significantly larger than the left
one.

DISCUSSION

To study the motor asymmetry in human subjects in relation to cerebral lateralization,

Goode et al.(l980) used the H-reflex recovery curve from the left and right leg. The
H-reflex is currently used in clinical investigations. T o obtain results comparable with
this kind of research, the H-reflex was used in this work to establish a possible spinal
motor asymmetry in cat.
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According to Mayer and Mawdsley (1965), an H-reflex can be elicited i n normal

anesthetized cats only rarely. In the present work, the H-reflex could be elicited in
all intact cats by using light pentobarbital anesthesia. The reflex nature of the recorded
H-wave was proven by cutting the appropriate dorsal roots. The above authors were
also unable to record an H-wave in the acute spinal state, and therefore, used chronic
spinal cats. No spinal shock was observed in this work, and an H-reflex could be
100,
I%) I
80-

6 0-

40-

FIGURE 3 Ambiiaterality in the H-reflex recovery c w e s in intact cats. Open and filled circles
correspond to the H-reflex recovery curves from the right and left sides, respectively. Abscissa and
ordinate, as in the previous figures.
 50 ti. TAN
elicited in the acute state as easily as before spinalization. These methodological
advantages made possible any comparison of the lateral asymmetry of the H-reflex
recovery curves in both the intact and the spinal state.
Mayer and Mawdsley (1965) have found that a different late wave can be recorded
in intact cats. This wave shared many characteristics of an H-reflex and had a long
latency. However, it persisted following proximal sciatic nerve section. This late
potential is different from the H-wave recorded in intact cats in this work, since it had
a shorter latency and disappeared after cutting the appropriate dorsal roots.
I n intact cats, three populations were distinguished according to differences in the
H-reflex recovery curves from the left and right side: right dominant, left dominant,
and ambilateral cats. There was no statistically significant difference(p=O.OI) between

A. INTACT
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801
60

'3
20

U 60 80 100 120 1
FIGURE4 Right spinal dominance in ambilateral cats revealed by spinalization. A. Ambilaterality
in the H-reflexrecovery curves in three intact cats. B. Elevation of the right recovery curve following
spinalization. Open and closed circles correspond to the right and left recovery curves, respectively.
Abscissa and ordinate, as in the previous figures.
 SPINAL MOTOR ASYMMETRY 51

the left and right recovery curves in 3 I .8 ”/, of the animals (ambilateral cats). In seven
cats (31.873, the right recovery curve was significantly higher than the left one,
indicating a right dominance. I n the remaining 8 cats (36.4%), the left recovery curve
was higher than the right one, indicating left dominance. The above described
three groups were almost equally distributed in regard to lateralization. However,
asymmetries were found in the majority of animals (68.2 %).
The recovery curve of the H-reflex has been shown to reflect the excitability of alpha
motoneurons (Magladery & McDougal, 1950; Magladery, 1955; Zander-Olsen et al.,
1967; Taborikova & Sax, 1968, 1969). Several factors including almost all of the
spinal inhibitory mechanisms may contribute to the depression in the recovery curve
of H-reflex. These mechanisms were not subjected to an analysis in the present work,
because the main purpose of this study was first to establish a lateralization in the
spinal motor system. The mechanisms of this asymmetry will be analyzed at a later
stage of the experiments. However, the effects of the muscle afferent discharge due to
the antagonistic muscle stretch caused by the conditioning stimulation, the changes in
the afferent muscle discharge caused by the conditioning reflex contraction, and the
effects of cutaneous stimulation can be excluded because of the extensive denervation
and recording from the cut LG-SOL nerve instead of intact muscle. Nevertheless, a
left shifted recovery curve may be compatible with less inhibition, i.e., elevated ex-
citatory state of alpha motoneurons. Consequently, it can be concluded that the
excitability of alpha motoneurons is distributed asymmetrically in the right and left
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dominant animals.
There was no correlation between paw preference and lateralization of the moto-
neuronal excitability. In normal human subjects, Goode et af. (1980) found differences
between the H-reflex recovery curves measured from the right and left leg. In con-
formity with the above results, they established that seven subjects had higher recovery
curves in the right leg, and four in the left leg, although all of them expressed right
hand preference. Thus, there is no correlation between preferred handedness and the
recovery curve asymmetry in man and cats. However, it was not examined whether
the H-reflex side differences are relatcd to paw preference of the forelimb in other
motor tasks.
It is well known that the spinal motor system is under the control of supraspinal
structures, and the spinal cord has a plasticity at least at the early stages of develop-
ment. Therefore, it is conceivable that a spinal motor lateralization develops in
parallel with cortical lateralization. To examine this hypothesis, the H-reflex recovery
curves were also measured in spinal cats. It was found that 18 of 20 cats (90%)
showed persistent lateral asymmetry of alpha motoneuron excitability. Ten of them
(56.6%) showed right spinal dominance, and eight left spinal dominance (44.4%).
These results suggest that there is a pure spinal motor lateralization in cats.
In summary, the present work provided evidence for a spinal motor lateralization
in cats. One possible explanation of this finding is an inherent asymmetry, since the
asymmetric depression of the H-reflex recovery curves was observed in both the
intact and the spinal state. Moreover, a left or right dominance appeared in ambi-
lateral cats following spinalization. Handedness in man was regarded as the outcome
of a process of maturation governed by heredity; in other words, laterality is deter-
mined genetically (Annett, 1964, 1967). However, there may be factors peculiar to
the human milieu which facilitate and channelize the handedness. In contrast, a motor
laterality in animals cannot be affected by social factors. An animal model is now
available to study the possible mechanisms of a spinal motor asymmetry and the
interrelationships between the cerebral and spinal lateralizations.
 52 u. TAN
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