C. A. Pratt

Functionally complex muscles of the cat hindlimb

SummaryThe biceps femoris (BF) muscle is divided into three neuromuscular compartments defined by the innervation patterns of the main nerve branches (English and Weeks 1987). The goals of this study were i) to determine how different regions of the biceps femoris muscle are activated in the intact cat during a broad range of limb movements evoked by perturbations of stance posture, and ii) to determine the relationship between the anatomical compartments of biceps femoris and the functional units as defined in this task. Cats were trained to stand on a moveable platform with each paw on a triaxial force plate. The animals stance was perturbed by linear translation of the platform in each of sixteen different directions in the horizontal plane. EMG activity was recorded from eight sites across the width of the left biceps femoris muscle. During quiet stance only the anterior compartment was tonically active, presumably contributing to hip extensor torque in the maintenance of stance. During platform translation, evoked EMG activity was recorded from each electrode pair for a wide range of directions of perturbation; as direction changed progressively, the amplitude of evoked activity from any electrode pair increased to a maximum and then decreased. When the EMG amplitude was plotted in polar coordinates as a function of translation direction, the region of response formed a petal shaped area in the horizontal plane, termed the EMG tuning curve. The compartments of the BF muscle were not activated homogeneously. The tuning curve of the anterior BF compartment was similar to that of other hip extensors, and coincided with the region of postero-lateral force production by the hindlimb against the support. The tuning curve of the middle BF compartment was shifted in a counterclockwise direction from that of the anterior compartment, but overlapped extensively with it; the middle BF tuning curve was similar to that of anterior gracilis. The tuning curve of the posterior biceps compartment was rotated further counterclockwise and overlapped very little with that of the middle BF compartment. The posterior BF was activated in a pattern similar to that of other knee flexors. The functional units of BF activation were not identical with the neuromuscular compartments defined by the main nerve branches. As direction of the perturbation changed, the region of BF that was activated moved progressively across the muscle. This progression of the active region was continuous across BFa and BFm, whereas there was a jump, or discontinuity at the border between BFm and BFp. Thus, differences in activation were observed not only across compartments, but also within compartments, and different regions of the BF muscle were activated independently during responses to postural perturbations.

Cat hindlimb motoneurons during locomotion. II. Normal activity patterns.

Activity patterns were recorded from 51 motoneurons in the fifth lumbar ventral root of cats walking on a motorized treadmill at a range of speeds between 0.1 and 1.3 m/s. The muscle of destination of recorded motoneurons was identified by spike-triggered averaging of EMG recordings from each of the anterior thigh muscles. Forty-three motoneurons projected to one of the quadriceps (vastus medialis, vastus lateralis, vastus intermedius, or rectus femoris) or sartorius (anterior or medial) muscles of the anterior thigh. Anterior thigh motoneurons always discharged a single burst of action potentials per step cycle, even in multifunctional muscles (e.g., sartorius anterior) that exhibited more than one burst of EMG activity per step cycle. The instantaneous firing rates of most motoneurons were lowest upon recruitment and increased progressively during a burst, as long as the EMG was still increasing. Firing rates peaked midway through each burst and tended to decline toward the end of the burst. The initial, mean, and peak firing rates of single motoneurons typically increased for faster walking speeds. At any given walking speed, early recruited motoneurons typically reached higher firing rates than late recruited motoneurons. In contrast to decerebrated cats, initial doublets at the beginning of bursts were seen only rarely. In the 4/51 motoneurons that showed initial doublets, both the instantaneous frequency of the doublet and the probability of starting a burst with a doublet decreased for faster walking speeds. The modulations in firing rate of every motoneuron were found to be closely correlated to the smoothed electromyogram of its target muscle. For 32 identified motoneurons, the units instantaneous frequencygram was scaled linearly by computer to the rectified smoothed EMG recorded from each of the anterior thigh muscles. The covariance between unitary frequencygram and muscle EMG was computed for each muscle. Typically, the EMG profile of the target muscle accounted for 0.88-0.96 of the variance in unitary firing rate. The EMG profiles of the other anterior thigh muscles, when tested in the same way, usually accounted only for a significantly smaller fraction of the variance. Brief amplitude fluctuations observed in the EMG envelopes were usually also reflected in the individual motoneuron frequencygrams. To further demonstrate the relationship between unitary frequencygrams and EMG, EMG envelopes recorded during walking were used as templates to generate depolarizing currents that were applied intracellularly to lumbar motoneurons in an acute spinal preparation.(ABSTRACT TRUNCATED AT 400 WORDS)

SINGLE UNIT CONDUCTION VELOCITIES FROM AVERAGED NERVE CUFF ELECTRODE RECORDS IN FREELY MOVING CATS

The conduction velocity of peripheral neurons recorded by wire microelectrodes implanted in intact, freely moving cats was determined on-line using the technique of spike-triggered averaging of nerve cuff electrode records described here. Axonal velocity was estimated from the conduction latency between two adjacent sets of tripolar recording electrodes inside a cuff, thereby avoiding uncertainties that could arise from differences in spike shape, variable conduction distance, or unknown stimulus utilization time. This method rendered conduction velocity values for individual afferent and efferent myelinated fibers ranging from 27 to 120 m/sec, estimated with an uncertainty of +/-5%. In addition, predictions from theoretical models relating extracellular potential amplitude, wavelength, and conduction velocity were confirmed experimentally for en passant records obtained from intact myelinated fibers.

A multiple-contact EMG recording array for mapping single muscle unit territories

The glycogen-depletion technique has become a well-established method for determining histologically the cross-sectional distribution of a single muscle unit. A major drawback of this method is its low yield of one depleted unit per experiment. Furthermore, this technique is particularly unsuited for determining the longitudinal distribution of single muscle units in long, broad muscles because of the formidable serial sectioning job that would be required. Our alternative, electrophysiological method utilizes a multiple-contact, two-dimensional EMG recording array to map efficiently the cross-sectional and longitudinal distributions of numerous single muscle units in anatomically diverse muscles. Additionally, architectural information on muscle fiber lengths, end-plate locations, motor subunit (MSU) arrangements and muscle conduction velocities can be determined.

Activity Patterns of Identified Alpha Motoneurons to Cat Anterior Thigh Muscles during Normal Walking and Flexor Reflexes

The organization of the motor apparatus into anatomically and functionally defined pools of regularly recruited motor units derives from some of the earliest and most enduring observations of neurophysiology. In recent years, there has been much productive research concentrated on discovering the anatomical bases of this organization in the spinal cord circuitry and the properties of the final common pathway, the alpha motoneurons themselves (for review, see Burke, 1981a). However, almost all of the direct evidence for the function of this system has been derived from experiments on reduced or anesthetized animals and on human subjects performing highly constrained and artificial motor tasks.