202
BRES 16835
Brain Research, 555 (1991) 202-214
© 1991 Elsevier Science Publishers B.V. 0006-8993/91/$03.50
A DONIS 0006899391168355
Coordination of motor pools controlling the ankle musculature in
adult spinal cats during treadmill walking
Camille P. de Guzman 2, Roland R. Roy 1, John A. Hodgson I and V. Reggie Edgerton 1'2
1Brain Research Institute and 2Neuromuscular Research Laboratory, Department of Kinesiology, Los Angeles, CA 90024 (U.S.A.)
(Accepted 5 March 1991)
Key words: Adult spinal cat; Treadmill locomotion; Electromyography; Agonist; Antagonist; Motor control
The coordination of the motor pools of two ankle plantarflexors, i.e. the soleus (Sol) and medial gastrocnemius (MG), and an ankle
dorsiflexor, i.e. the tibialis anterior (TA) was quantified by comparing the EMG amplitude relationships in muscle pairs in normal and trained
adult spinalized cats during treadmill walking across a range of relatively slow speeds (0.1 to 1.0 m/s). The effects of increased tactile stimulation
or loading on locomotor performance were also studied in the spinal cats. Joint probability density distributions in the spinalized cats showed
a low level of MG activation relative to Sol which did not change as speed increased. In general, the coordination between Sol and MG was
similar in normal and spinal cats. However, towards the final phase of the extensor burst, the MG EMG amplitude decayed prematurely in
spinal cats, particularly at higher speeds. Preferential elevation of MG relative to Sol activity was seen as a result of tactile stimulation. An
elevated load resulted in a higher level of MG activation relative to Sol, prolonged MG activity at the end of the extensor burst, and the
reduction in the clonic pattern of EMG typical of spinal cats. Spinalized cats showed an increased incidence of SoI-TA coactivation, especially
at the higher speeds, due in part to the tonic activity in the TA. However, the overall reciprocal relationship between these antagonists was
maintained. This reciprocity was preserved, but the high level of coactivation was unaffected by tactile stimulation. An elevated load, however,
resulted in less SoI-TA coactivation. These results suggest that the coordination between synergists (SoI-MG) and between antagonists (SoI-TA
and MG-TA), as well as the level of activation are modulated in the adult spinal cat similar to that observed in the normal cat. Further, there
are specific types of proprioceptive-cutaneous information that can affect selected phases of the step cycle such that full weight-supporting
stepping is significantly improved.
INTRODUCTION
It is generally believed that the recruitment of motor
units within a motor pool occurs in a relatively constant
order related to motoneuron size or closely associated
parameters 9'1°'1s. However, the recruitment pattern
among agonist and antagonist motor pools has been
characterized only qualitatively. For example, it is known
that the onset and termination of activation of the soleus
(Sol) and medial gastrocnemius (MG) occur at about the
same time and that the relative level of activation of the
Sol motor pool exceeds the MG during slow loco-
motion 24'3°. The MG becomes more highly activated only
in movements that have high tension or power re-
quirements 19'2°'24'3°'32. Also it is well recognized that the
activation sequence of antagonists such as the tibialis
anterior (TA) and the MG are generally out of phase,
although some overlap in the initiation of TA activity and
the termination of MG activity (i.e. co-contraction)
usually occurs 14'15'24. The regulation of the relative
activation of agonist and antagonist motor pools has been
described more quantitatively. For example, the proba-
bility of occurrence of any given combination of EMG
amplitudes in normal rats during locomotion at varying
speeds and grades and during swimming, when the
demands of weight support are eliminated, have been
determined 2°. The relationship between the Sol and
gastrocnemius in normal cats during various motor tasks
has been described quantitatively in a similar manner 19.
Previous studies have detailed the kinematics and
EMG characteristics of hindlimb stepping on a treadmill
of cats spinalized at a low thoracic level at 1-2 weeks of
age 12'13, at 2-12 weeks of age 8'12'28'29 or as an adult 2'
7,11,22,23,25,26 From these data, it is clear that the
hindlimbs of spinalized cats trained daily to step on a
treadmill can recover a level of coordination that in many
respects resembles normal locomotion. In the present
study, the coordination of motor pools during locomotion
was quantified by comparing the rectified and smoothed
EMG amplitudes in muscle pairs in normal cats and in
cats spinalized as adults. The probability and sequence of
occurrence of pairs of amplitudes during the activation of
Correspondence: R.R. Roy, Brain Research Institute, Center for the Health Sciences, University of California, Los Angeles, 10833 Le Conte
Avenue, Los Angeles, CA 90024-1761, U.S.A. Fax: (1) (213) 825-6616.