Lateralized alpha-motoneuron excitabilities

during lying and standing of healthy

individuals in relation to Parkinsonian rigidity
Sibel Karaca1, Meliha Tan1, Uner Tan2
1
Department of Neurology, Adana Teaching and Research Center, Faculty of Medicine, Baskent University,
Adana, Turkey, 2Department of Physiology, Medical School, Çukurova University, Adana, Turkey

Objectives: To elucidate mechanisms of Parkinsonian rigidity by assessing excitability of alpha-

motoneurons innervating right and left soleus muscles in healthy controls and Parkinson’s disease (PD)
patients with rigidities in the right, left and both legs.
Published by Maney Publishing (c) W. S. Maney & Son Limited

Methods: One group of 45 controls was recruited and 60 PD patients in three groups: rigidities,
predominantly in the right, left and both legs. H-reflex (H) and muscle response (M) were recorded from
right and left soleus muscles during stimulations of the posterior tibial nerve at the popliteal fossa while lying
and standing. The H/M ratio was taken as an index for motoneuron excitability.
Results: Mean H/M ratios were significantly different on the right and left sides, modified by postural
changes in controls and PD patients. Analysis of variance showed that in healthy subjects the H/M ratio
was: standing.lying (right), lying.standing (left). In right leg rigidity patients, the H/M ratio was greatest
during standing, and smallest during lying. In left leg rigidity patients, the H/M ratios on the right and left
sides were equally independent of posture. In controls, left H/M.right while lying, ,right while standing. In
right leg rigidity patients, right H/M.left, but ,right in left leg rigidity patients, independent of posture.
There was no side difference in patients with rigidity in both legs.
Conclusions: (i) motoneuron excitability may show side and postural differences in healthy individuals and
PD patients; (ii) posture may be associated with lateralized motoneuron excitability in these subjects; and
(iii) Parkinsonian rigidity may have spinal motor origins.
Keywords: Parkinson’s disease, H-reflex, Motoneuron excitability, Posture, Laterality

Introduction individuals were not compared with those obtained

In 1910, Paul Hoffmann first described a reflex in
humans that is the electrical equivalent of the deep
tendon reflex. In this reflex the muscular primary
afferents are stimulated electrically and the monosy-
naptic reflex response from the homonymous synergic
alpha-motoneurons is recorded.1 Since its discovery the
Hoffmann reflex – or the H-reflex (H) – has frequently
been used to assess alpha-motoneuron excitability in
pathological and physiological conditions.
Studies using this reflex have shown that recovery
curves of the soleus H-reflexes exhibit side differences
in healthy individuals, with the alpha-motoneuron
excitability being higher on the left side than the right
in right handers, and vice versa in left handers.2
However, the subjects were lying face down during
these investigations, and the motoneuron excitability
modulations during different postures of healthy
Correspondence to: Uner Tan, Department of Physiology, Medical School,
Cukurova University, 01330 Adana, Turkey. Email: unertan37@yahoo.
com
with the subjects lying down. Handedness has also
been shown to be associated with motor symptom
asymmetry in Parkinson’s disease (PD),3,4 with some
inconsistent results.5
The soleus alpha-motoneuron excitability assessed
by the H-reflex amplitude in healthy individuals was
shown by other authors to vary with posture. For
instance, the H-reflex amplitude was smaller when
standing than when prone;6 the soleus H-reflex was
suppressed during standing compared to that under
unloading7 and prone,8 but there was no difference
between H-reflex amplitudes when sitting or lying
prone.9,10 However, more appropriate methods to
assess the motoneuron excitability, such as the H-
reflex/muscle response (H/M) ratio or the H-reflex
recovery curve, were not used either by these or by
other authors.
The intrinsic excitability and the presynaptic
inhibition of the alpha-motoneurons should be kept
constant if the H-reflex amplitude is to be used to

ß W. S. Maney & Son Ltd 2011

976 DOI 10.1179/1743132811Y.0000000024 Neurological Research 2011 VOL . 33 NO . 9
 Karaca et al.
estimate the motoneuron excitability. The H-reflex
amplitudes must be normalized to take into account
variations due to skin resistance, subcutaneous fat
levels, and the location of the stimulus electrode
before any comparison of the H-reflex amplitudes can
be made. This is done by taking H/M.11
The Hmax/Mmax ratio is commonly used as a
dependent measure when data are being collected
on more than one occasion.11 For these reasons we
used the Hmax/Mmax as an index for the alpha-
motoneuron excitability changes during postural
conditions in healthy individuals and cases with PD,
instead of H-reflex amplitudes alone as used in the
other studies.
The alpha-motoneuron excitability assessed by the
H-reflex amplitude was also reported with regard to
Parkinsonian rigidity.12,13 These studies, however,
Published by Maney Publishing (c) W. S. Maney & Son Limited
also used the H-reflex amplitude as an index for the
motoneuron excitability, without considering a more
appropriate method such as the H/M ratio or H-
reflex recovery curve. Despite these methodological
inadequacies, the above studies suggested an increase
in the alpha-motoneuron excitability as the cause of
the rigidity in PD patients.
In light of the above mentioned H-reflex studies in
healthy individuals and patients with PD, with some
minor but important methodological inadequacies,
we studied the alpha-motoneuron excitabilities asse-
ssed by the H/M ratios in healthy individuals and
patients with PD, to determine if there are bilateral
differences and modulations in alpha motor neuron
excitabilities produced by postural changes.
Consequently, we expected to gain more insights
into the laterality of the final common pathways in
healthy individuals compared to PD patients exhibit-
ing rigidities exclusively in the right, left or both legs.
Studying the alpha-motoneuron excitabilities on the
right and left sides during lying and standing of
healthy individuals and patients with PD –for the first
time in the scientific literature – would give further
insight into the pathophysiology of the Parkinsonian
rigidity. Moreover, the results obtained from this
study might also shed light on the outcomes of the
therapeutic interventions applied to PD patients with
or without drugs.
The specific aims of this study were: (i) to compare
the extensor alpha-motoneuron excitabilities in heal-
thy individuals (controls) with PD patients, using the
H/M ratios as a motoneuron excitability index with
regard to side-related differences; (ii) to gain a more
direct insight into the origins of the Parkinsonian
rigidity by examining the relationship of the rigi-
dity scores to the alpha-motoneuron excitability le-
vels in patients with a predominant right, left, or both
leg rigidity, and (iii) to elucidate any asymmetric
influence of postural changes (lying and standing) on
Alpha-motoneuron excitabilities in relation to Parkinsonian rigidity
the alpha-motoneuron excitabilities from the right
and left sides of healthy individuals and cases with
one sided and both sided Parkinsonian rigidity.
Methods
There were two categories of subjects voluntarily
participated in this study: healthy individuals and PD
patients. These were divided into four groups. The
first category (group 1) consisted of 45 healthy
individuals (15 women and 30 men) with no known
neurological signs and symptoms. The second cate-
gory comprised 50 cases with PD (25 men and 25
women). This category was divided into three groups:
patients with predominantly right leg rigidity (n522),
patients with predominantly left leg rigidity (n512),
and patients with both leg rigidity (n516). The PD
patients were newly diagnosed and not yet taking
medication for their disease.
The rigidity subscore of the unified PD rating score
(UPDRS) was used to assess the severity of the
rigidity,18 which was determined using the rigidity
scores of part III of the UPDRS subscores. All of the
PD patients had the following cardinal signs: distal
resting tremor, bradykinesia, and asymmetric onset.19
The statistical analysis was performed using the SPSS
statistical package, version 16, for the univariate
analysis of variance (ANOVA) with post hoc tests
(Bonferroni) for multiple comparisons. The statistical
tests should at least yield P,0.05 for the results to be
statistically significant.
All of the subjects in the four groups were
consistent right handers and consistent right footers.
The hand preference was assessed by the Oldfield’s
handedness questionnaire, and foot preference was
assessed by kicking a ball.14
Each subject signed a consent form prior to the
examinations, and the experimental procedure was
approved by the ethics committee of the Medical
School at Baskent University (Ankara, Turkey).
Traditional methods were used to elicit and record
the H-reflexes. That is, the skin was cleaned with
alcohol, shaved, and slightly brushed. A pair of
recording electrodes 1 cm in diameter and 5 cm apart
was placed on the midpoint of the soleus muscle. Two
stimulus electrodes, with a 1 cm diameter cathode
placed at the medial popliteal nerve and a larger plate
placed over the patella, were used to elicit the H-
reflex, using single shock stimuli of 0.5 milliseconds
duration at 10 seconds intervals. The correct stimulus
point for the largest reflex amplitudes was determined
by moving the stimulus electrodes over the popliteal
fossa until the largest reflexes with smallest muscle
responses appeared. The amplitudes of the H-reflexes
first increased then decreased and disappeared with
increasing the 61.0 threshold stimuli. The amplitudes
of the M (muscle) and H (reflex) responses were
Neurological Research 2011 VOL . 33 NO . 9 977
 Karaca et al. Alpha-motoneuron excitabilities in relation to Parkinsonian rigidity

Figure 1 Muscle (M) and H-reflex (H) responses recorded

from the right soleus muscle following single-shock stimula-
tion of the posterior tibial nerve at the popliteal fossa in a
healthy individual. Ordinate: amplitude (mV), abscissa: time
(millisecond).

measured from their lowest to highest peaks during

recording sessions. The alpha-motoneuron excitabil-
ities were assessed by taking the ratio of the highest
Published by Maney Publishing (c) W. S. Maney & Son Limited

amplitude H response to the highest amplitude M

response.15 Examining the H/M ratios with the
subjects both during prone lying and upright standing
was taken as a reliable procedure since a high test-
retest reliability has been previously reported for the
H-reflex amplitude during three different positions
including standing.16,17
To obtain the H-reflex and muscle responses, we
used a Nihon Cohden Neuropack (Tokyo, Japan)
electromyography monitor with a sampling rate of
2 kHz per channel. SPSS, version 16, was used to
make statistical analysis, for one-way ANOVA and
post hoc tests.

Results
The subjects comprised four groups: healthy indivi-
duals (n545) aged 42–69 years (58.1¡6.5 years
with no known neurological signs or symptoms, PD
patients with predominantly right leg rigidity (n522)
aged between 44 and 72 years, PD patients with
Figure 2 Mean H/M ratios (ordinate) from right (A) and left
predominantly left leg rigidity (n512) aged between
51 and 65 years, and PD patients with rigidity in both
(B) soleus muscles, during lying (o) and standing ( ) of N
healthy subjects and PD patients with predominantly right or
legs (n516) aged 63–76 years. The right and left mean left leg rigidity (abscissa). Vertical lines above and below the
UPDRS motor subscores for rigidity were distributed mean H/M ratios: ¡95% confidence intervals.
among patients as follows: right UPDRS512.4¡2.5
and left UPDRS53.6¡3.1 for the patients with pre-
responses belonged to the M response (5.2 millise-
dominantly right leg rigidity, right UPDRS51.2¡0.9
conds) evoked only by alpha-efferent stimulation.
and left UPDRS512.6¡3.7 for the patients with
The H response had a much longer latency
predominantly left leg rigidity, right UPDRS5
(30.0 milliseconds) because of its reflex nature with
17.3¡3.5 and left UPDRS517.9¡6.6 for the patients
afferent–efferent connections and synaptic conduc-
with both leg rigidity. The difference between the mean
tion times.
ages of the normal subjects and patients was not
statistically significant (t51.4, df5103, P.0.10), Postural differences
indicating the groups participating in the present The mean right leg H/M ratios during lying and
study were age matched. standing of healthy subjects, and PD patients with
The M and H responses recorded from the right rigidity predominantly in the right and left leg were
soleus muscle of a healthy subject are depicted in compared in Fig. 2A. One-way ANOVA indicated
Fig. 1. As expected, the shortest latency of these that there were no significant differences between

978 Neurological Research 2011 VOL . 33 NO . 9

 Karaca et al. Alpha-motoneuron excitabilities in relation to Parkinsonian rigidity

groups during lying (F2, 7750.25, P.0.75), but the

mean H/M ratios between groups were significantly
different during standing (F2, 77512.44, P,0.001).
For the standing posture, post hoc tests showed that
the mean right H/M ratio of the patients with
predominantly right leg rigidity was significantly
greater than the mean right H/M ratio of the control
subjects (P,0.001) and the patients with left leg
rigidity (P,0.01). The difference between the mean
right H/M ratios of the control subjects and the
patients with left leg rigidity was not significant
(P.0.50).
Considering only the postural conditions in
Fig. 2A, the mean right H/M ratio was significantly
greater during standing than lying in healthy
individuals (F1, 43511.26, P,0.001) and in patients
with predominantly right leg rigidity (F1, 21520.24,
Published by Maney Publishing (c) W. S. Maney & Son Limited

P,0.0001), there being no significant postural

difference in the right leg H/M ratios of the patients
with left leg rigidity (F1, 1151.12, P.0.30).
Figure 2B illustrates the mean left H/M ratios
during lying and standing of the healthy individuals,
and patients with right or left leg rigidity. The one-
way ANOVA indicated that the difference be-
tween groups was significant for the lying posture
(F2, 77512.82, P,0.001). Post hoc tests for multiple
comparisons showed that the mean left H/M ratio of
patients with predominantly right leg rigidity was
significantly smaller than in the healthy individuals
(P,0.001), and patients with left leg rigidity
(P,0.001). The largest left H/M ratio belonged to
Figure 3 Mean H/M ratios with ¡95% confidence intervals
the patients with predominantly left leg rigidity, being
significantly greater in these patients than those with N
(ordinate) from right (o) and left ( ) sides during lying (A) and
standing (B) of healthy subjects, and PD patients with
right leg rigidity and healthy subjects (P,0.001). For predominantly right, left, or both leg rigidity (abscissa).
the standing posture, there was a significant difference
between groups (F2, 77533.5, P,0.001). Post hoc tests
were no significant differences between the mean
for multiple comparisons showed, however, the mean
right H/M ratios during lying of patients with
left H/M ratio during standing of healthy individuals
rigidity in the left leg and both legs (P.0.90 and
was not significantly different from the patients with
0.95). With regard to the left side (Fig. 3A, closed
right leg rigidity (P.0.05), but the mean left H/M ratio
circles), the mean H/M ratios were significantly
for the patients with left leg rigidity was significantly
different between groups (F2, 92514.01, P,0.001).
greater than those for the healthy subjects and patients
Post hoc tests showed the mean left H/M ratio for
with right leg rigidity (P,0.001).
the patients with right leg rigidity was significantly
Side differences smaller than the healthy individuals (P,0.001); the
Figure 3A illustrates the mean right and left H/M mean left H/M ratio for patients with left leg rigidity
ratios during lying of the healthy subjects, and was significantly greater than the patients with right
patients with right, left, or both leg rigidity. One- leg rigidity (P,0.001), both leg rigidity (P,0.05),
way ANOVA indicated significant differences be- and healthy subjects (P,0.005).
tween the groups for the mean right H/M ratios Considering the right and left differences during
(F2, 9253.45, P,0.05). Post hoc tests showed, lying of the healthy subjects (Fig. 3A), the mean left
however, that the only significant difference between H/M ratio was significantly greater than the mean
groups was due to the mean H/M ratio of the patients right H/M ratio (F1, 43512.15, P,0.001). By contrast,
with predominantly right leg rigidity, being signifi- the mean right H/M ratio was significantly greater
cantly greater than the control group, and patients than the mean left H/M ratio in patients with right leg
with right, left, or both leg rigidity (P,0.05). There rigidity (F1, 205P,0.001). In patients with left leg

Neurological Research 2011 VOL . 33 NO . 9 979

 Karaca et al. Alpha-motoneuron excitabilities in relation to Parkinsonian rigidity
rigidity the mean left H/M ratio was significantly
greater than the mean right H/M ratio (F1, 10526.84,
P,0.001). In patients with both leg rigidity, there was
no significant difference between the mean right and
left H/M ratios (F1, 1450.48, P.0.45).
Figure 3B illustrates the mean right and left H/M
ratios during standing of healthy subjects and
patients with right, left, or both leg rigidity.
ANOVA yielded significant differences between
groups for the right side (F3, 91518.71, P,0.001).
Post hoc tests for multiple comparisons indicated the
highest right H/M ratio for the patients with right leg
rigidity, being higher than controls (P,0.001),
patients with left leg rigidity (P,0.001), and both
leg rigidity (P,0.001). For the left H/M ratios,
ANOVA yielded significant differences between
groups (F3, 91559.43, P,0.001). According to post
Published by Maney Publishing (c) W. S. Maney & Son Limited
hoc tests for multiple comparisons, the mean left H/M
ratio was significantly greatest for the patients with
left leg rigidity, being greater than patients with right
leg rigidity, both leg rigidity, and controls (P,0.001).
With regard to right left differences during standing
upright (Fig. 3B), the mean right H/M ratio was
significantly greater than the left H/M ratio in healthy
subjects (F1, 43528.82, P,0.001), and patients with
right leg rigidity (F1, 20526.56, P,0.001), being
significantly smaller in patients with left leg rigidity
(F1, 10549.12, P,0.001). There was no significant
right–left difference in patients with both leg rigidity
(F1, 1450.56, P.0.45).
Discussion
The quintessence of this study was the side and
postural differences in the alpha-motoneuron excit-
ability in healthy subjects and patients with PD in
relation to one sided and both sided rigidities.
In healthy individuals the mean right and left H/M
ratios, as an index for the alpha-motoneuron excit-
ability, exhibited significant side differences that were
modified by postural changes: higher motoneuron
excitability during standing than lying on the right
side, but lower excitability during standing than lying
on the left side. This reversal in spinal motor
asymmetry depending upon posture – reported for
the first time in the literature – may be segmental as
well as supraspinal in origin, suggesting the ‘mod-
ulatory effects of descending and segmental inputs on
the excitability of the H-reflex circuit’.20 However, the
H-reflex amplitudes have been shown to decrease
during standing in patients with a complete spinal
cord injury,21 suggesting a primary contribution of
peripheral sensory inputs to suppression of the so-
leus H-reflex while standing. Thus, the spinal motor
asymmetry2 modified by postural conditions may
mainly have peripheral origins independent of sup-
raspinal influences.
980 Neurological Research 2011 VOL . 33 NO .
Considering the side differences in relation to
postural conditions, in the control subjects the alpha-
motoneuron excitability was significantly higher on
the left than the right side during lying, but higher on
the right than the left side during standing. These
results are partly consistent with previous reports, with
higher motoneuron excitability on the left than the
right side during lying of the right handed healthy
subjects.2 Thus, the asymmetric spinal motor excit-
ability in the lower extremities opposite to hand and
foot preferences seems to be well established, but only
when subjects are lying down.
It was previously argued2 that the discrepancy
between hand and foot preference and motoneuron
excitability in the lower extremities may be explained
by a predominantly inhibitory action of the cerebral
cortex on the contralateral extensor motoneurons.20
However, the subjects in these studies were lying
down and no comparison was made for various
postural conditions other than lying prone. The
above results now demonstrate that the opposite
could be observed when the subjects changed their
posture from lying to standing, with higher moto-
neuron excitability on the right than the left. The
mechanisms of this change in asymmetric motoenur-
onal excitability may be explained by supraspinal
influences on the extensor motoneurones, but, as
mentioned above, peripheral-segmental mechanisms
may be essential in the reverse change of spinal motor
asymmetry, i.e., following transition from lying to
standing posture.22
Lateralization in alpha-motoneuron excitability
depending upon the postural conditions was also
observed in the Parkinson’s patients in relation to
rigidity. The motoneuron excitabilities of the right
and left sides were significantly greater during
standing than lying in patients with predominantly
right leg rigidity. However, this was not the case for
the patients with predominantly left leg rigidity:
the motoneuron excitabilities were not significantly
different from each other on the right and left sides
during lying and standing in these cases.
These results, which are novel in the literature, may
be due to exclusively right handedness and right
footedness of the subjects who participated in the
study, but there is no supporting information in the
literature for this hypothesis. On the other hand, an
increase in the alpha-motoneuron excitability in PD
was previously reported,23 but without considering
the side differences and modulations by posture or
their combined actions, and indirectly measuring
the alpha-motoneuron excitability using the H-reflex
amplitudes. We have also found that the mean right
leg H/M ratios during lying and standing of the
patients with left leg rigidity were not significantly
different from those of the control subjects. This is
9
 Karaca et al.
not consistent with reports about the increase of the
alpha-motoneuronal excitability in PD. Accordingly,
the increase in the alpha-motoneuron excitability in
PD may not be considered as a rule, provided that the
affected sides during lying and standing are investi-
gated separately. Moreover, the mean right leg H/M
ratios of both legs of the patients exhibiting rigidity
predominantly in the left leg and both legs not
significantly different from healthy controls during
lying and standing conditions. This finding is also not
consistent with the view of an increased motoneuron
excitability in PD patients.
The results suggest that lateralization of the alpha-
motoneuron excitability may be associated with
lateralization of Parkinsonian rigidity. Namely, the
mean H/M ratios from right and left sides were not
significantly different from each other in patients with
Published by Maney Publishing (c) W. S. Maney & Son Limited
rigidity in both legs, indicating association of sym-
metric rigidity with symmetric motoneuron excitabil-
ity. In patients with predominantly left leg rigidity,
the mean left leg H/M ratios during lying and
standing were significantly greater than the mean
right leg H/M ratios during lying and standing; the
mean left leg H/M ratios in both postural conditions
were significantly greater than in the controls, but the
mean right leg H/M ratios were not significantly
different from the controls. These results indicate
association of the increased left leg rigidity in patients
with predominantly left leg rigidity with a predomi-
nantly left sided increase in the alpha-motoneuron
excitability, without any significant change in the
right sided motoneuron excitability. With regard to
the patients with predominantly right leg rigidity, the
mean right leg H/M ratios during lying and standing
were significantly greater than for the controls, while
the mean left leg H/M ratio during standing was not
statistically different from the controls, but was
significantly smaller than in the controls during lying.
These results show an association of the strongly
right sided rigidity with an increase in the purely right
sided alpha-motoneuron excitability.
These never before reported results present direct
evidence for the contribution of asymmetric levels of
the alpha-motoneuron excitabilities to the level of
the asymmetric Parkinsonian rigidity. This contra-
dicts the notion of an overall motoneuron excit-
ability increase in PD found in studies that did not
consider the sides and the postural conditions, and
which used an unreliable method such as the H-
reflex amplitudes to measure the motoneuron excit-
ability changes in patients with different levels of
Parkinsonian rigidity.
This is the first study presenting the asymmetric
occurrence of the alpha-motoneuron excitabilities in
relation to the asymmetric distribution of Parkin-
sonian rigidity in relation to postural changes. The
Alpha-motoneuron excitabilities in relation to Parkinsonian rigidity
results of the present work are also of predictive value:
by simply examining the rigidity of legs (right, left or
both) one can predict the degree and lateralization of
the spinal alpha-motoenruon excitabilities, and this
may in turn be beneficial for the follow-up studies and
may have therapeutical implications.
The increased alpha-motoneuron excitability, play-
ing a role in the mechanisms of the Parkin-
sonian rigidity, may result from a decrease in the
autogenetic Ib inhibition in PD, while the role of the
presynaptic inhibition in Parkinsonian rigidity is
controversial.24,25 These studies, however, did not
report any side differences modulated by postural
changes. On the other hand, contrary to our results,
in other studies no statistically significant difference
was found between motoneuron excitabilities of the
healthy subjects and Parkinson’s patients,25,26 but
these studies also did not consider the side differences
in various postural conditions.
Conclusions
The excitability of the alpha-motoneurons innervat-
ing the postural soleus muscle was assessed by taking
the mean H/M ratios in normal subjects, and PD
patients rigidity predominantly in the right, left or
both legs, to gain an insight into a possible
contribution of the alpha-motoneuron excitability
to the Parkinsonian rigidity. This is the first study
measuring the motoneuron excitability using the H/
M ratio from right and left soleus muscles during
lying and standing of healthy individuals and patients
with PD. The current literature did not report the side
differences in motoneuron excitabilities during pos-
tural changes, and methods such as the H-reflex
amplitudes were used to measure the motoneuron
excitability changes. A more appropriate method (H/
M ratio) was used in the present work to directly
measure the motoneuron excitability, and more
detailed information was obtained on the contribu-
tion of the alpha-motoneuron excitability to the
origins of Parkinsonian rigidity by considering the
side differences in various postural conditions. The
strong association of the symmetric or asymmetric
Parkinsonian rigidity with the symmetric or asym-
metric motoneuron excitability during lying and
standing compared to healthy controls provided
direct evidence for the spinal motor origins of the
rigidity in PD patients.
The results are also of predictive value for the side
differences in alpha-motoneuron excitabilities by a
simple examination of the rigidities on the right and
left sides of the PD patients. These original, never
before reported results also accentuate the impor-
tance of side differences under various postural
conditions, and may have implications for future
research and follow-up studies with or without drugs.
Neurological Research 2011 VOL . 33 NO . 9 981
 Karaca et al. Alpha-motoneuron excitabilities in relation to Parkinsonian rigidity
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