Modulation of forelimb and hindlimb muscle activity during quadrupedal tied-belt and split-belt locomotion in intact cats.

The modulation of the neural output to forelimb and hindlimb muscles when the left and right sides step at different speeds from one another in quadrupeds was assessed by obtaining electromyography (EMG) in seven intact adult cats during split-belt locomotion. To determine if changes in EMG during split-belt locomotion were modulated according to the speed of the belt the limb was stepping on, values were compared to those obtained during tied-belt locomotion (equal left-right speeds) at matched speeds. Cats were chronically implanted for EMG, which was obtained from six muscles: biceps brachii, triceps brachii, flexor carpi ulnaris, sartorius, vastus lateralis and medial gastrocnemius. During tied-belt locomotion, cats stepped from 0.4 to 1.0m/s in 0.1m/s increments whereas during split-belt locomotion, cats stepped with left-right speed differences of 0.1 to 0.4m/s in 0.1m/s increments. During tied-belt locomotion, EMG burst durations and mean EMG amplitudes of all muscles respectively decreased and increased with increasing speed. During split-belt locomotion, there was a clear differential modulation of the EMG patterns between flexors and extensors and between the slow and fast sides. Changes in the EMG pattern of some muscles could be explained by the speed of the belt the limb was stepping on, while in other muscles there were clear dissociations from tied-belt values at matched speeds. Therefore, results show that EMG patterns during split-belt locomotion are modulated to meet task requirements partly via signals related to the stepping speed of the homonymous limb and from the other limbs.

Coordination between the fore- and hindlimbs is bidirectional, asymmetrically organized, and flexible during quadrupedal locomotion in the intact adult cat.

Despite the obvious importance of inter-girdle coordination for quadrupedal locomotion in terrestrial mammals, its organization remains poorly understood. Here, we evaluated cycle and phase durations, as well as footfall patterns of four intact adult cats trained to walk on a transverse split-belt treadmill that could independently control fore- and hindlimb speed. When the hindlimbs walked at faster speeds than the forelimbs, an equal rhythm was always maintained between the fore- and hindlimbs, even at the highest fore-hindlimb speed ratio of 1:3 (0.4:1.2 m/s). The locomotor pattern adjusted through changes in both hindlimb stance and swing phase durations, whereas only the forelimb stance phase was affected. In such conditions, when fore- and hindlimb values were compared to those obtained at matched speeds during tied-belt walking (i.e. predicted values based on treadmill speed), hindlimb cycle, stance and swing durations were consistently longer than predicted. On the other hand, forelimb cycle and stance durations were shorter than predicted but only at the highest split-belt speed ratios. Forelimb swing durations were as predicted based on front-belt speed. The sequence of footfall pattern when hindlimb speed was faster was identical to tied-belt walking. In stark contrast, when the forelimbs walked at slightly faster speeds than the hindlimbs, the rhythm between the fore- and hindlimbs broke down. In such conditions, the locomotor pattern was adjusted through changes in stance and swing phase durations in both the fore- and hindlimbs. When the rhythm between the fore- and hindlimbs broke down, hindlimb cycle and phase durations were similar to predicted values, whereas forelimb values were shorter than predicted. Moreover, several additional sequences of footfall patterns were observed. Therefore, the results clearly demonstrate the existence of a bidirectional, asymmetric, and flexible control of inter-girdle coordination during quadrupedal locomotion in the intact adult cat.

Re-expression of locomotor function after partial spinal cord injury.

After a complete spinal section, quadruped mammals (cats, rats, and mice) can generally regain hindlimb locomotion on a treadmill because the spinal cord below the lesion can express locomotion through a neural circuitry termed the central pattern generator (CPG). In this review, we propose that the spinal CPG also plays a crucial role in the locomotor recovery after incomplete spinal cord injury.

Partial denervation of ankle extensors prior to spinalization in cats impacts the expression of locomotion and the phasic modulation of reflexes.

Following peripheral nerve sections some locomotor deficits appear which are gradually compensated for by spinal and supraspinal mechanisms. The present work is aimed at identifying contributions of both types of mechanisms. We performed a denervation of the left lateral gastrocnemius-soleus (LGS) muscles in three cats which was followed by a spinalization at the 13th thoracic segment. Three other cats were not denervated prior to spinalization (i.e. intact) and served as controls. Over the years, in our laboratory, there have been no instances in which cats did not express spinal locomotion with treadmill training and/or clonidine administration. After spinalization, cats were trained on a treadmill to express spinal locomotion. Reflexes, evoked by stimulating the left tibial nerve at the ankle, the electromyography of several hindlimb muscles, and kinematics were recorded during locomotion before and after denervation, during recovery, and following complete spinalization. Denervating the left LGS before spinalization induced considerable variability in the expression of spinal locomotion from one cat to another, which was not observed in the three controls. Variability ranged from a greater ankle yield in the denervated limb in one cat to inability to recover locomotion after spinalization in another. In the two denervated cats that recovered locomotion after spinalization, some reflex changes differed from "normal" spinal cats (i.e. intact at the time of spinalization), suggesting that reorganization of spinal circuits after spinalization is dissimilar to what normally takes place if denervation is performed before spinalization. First, we conclude that the transient locomotor deficits initially incurred following the LGS denervation in cats with an intact spinal cord reappear after complete spinalization indicating that supraspinal mechanisms were involved in maintaining the adapted locomotion. Second, the reappearance of locomotor deficits and/or impairment in expressing spinal locomotion suggests that spinal mechanisms were profoundly altered to compensate for the initial denervation partly because the reflex modulation is abnormal. If the same denervation is performed after spinalization only transient deficits are observed and spinal locomotion is not compromised.