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Biomedical subjects

G E Loeb

Publications and source records attributed to G E Loeb.

At least 91 records · Page 5Linked to original sources

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 unit's 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)

Action Potentials↗

Cat hindlimb motoneurons during locomotion. III. Functional segregation in sartorius.

Cat sartorius has two distinct anatomical portions, anterior (SA-a) and medial (SA-m). SA-a acts to extend the knee and also to flex the hip. SA-m acts to flex both the knee and the hip. The objective of this study was to investigate how a "single motoneuron pool" is used to control at least three separate functions mediated by the two anatomical portions of one muscle. Discharge patterns of single motoneurons projecting to the sartorius muscle were recorded using floating microelectrodes implanted in the L5 ventral root of cats. The electromyographic activity generated by the anterior and medial portions of sartorius was recorded with chronically implanted electrodes. The muscle portion innervated by each motoneuron was determined by spike-triggered averaging of the EMGs during walking on a motorized treadmill. During normal locomotion, SA-a exhibited two bursts of EMG activity per step cycle, one during the stance phase and one during the late swing phase. In contrast, every recorded motoneuron projecting to SA-a discharged a single burst of action potentials per step cycle. Some SA-a motoneurons discharged only during the stance phase, whereas other motoneurons discharged only during the late swing phase. In all cases, the instantaneous frequencygram of the motoneuron was well fit by the rectified smoothed EMG envelope generated by SA-a during the appropriate phase of the step cycle. During normal locomotion, SA-m exhibited a single burst of EMG activity per step cycle, during the swing phase. The temporal characteristics of the EMG bursts recorded from SA-m differed from the swing-phase EMG bursts generated by SA-a.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Cat hindlimb motoneurons during locomotion. IV. Participation in cutaneous reflexes.

The responses of 11 individual motoneurons, the muscle to which each projected, plus all other muscles in the anterior thigh of the cat, were recorded following single non-noxious electrical stimuli to cutaneous nerves while the intact animal walked on a treadmill. The various excitatory and/or inhibitory responses were qualitatively similar for stimuli within the range 1.1-10 times threshold for group I fibers in the stimulated nerve (usually saphenous). Monarticular knee extensor muscles in the vastus group and their motoneurons were usually inhibited in the period 10- to 25-ms poststimulus. The faster contracting vastus medialis and lateralis muscles tended to have an excitatory rebound at approximately 25- to 40-ms poststimulus that was confined to the stance phase of the step cycle when these muscles were normally active. Biarticular hip flexor muscles rectus femoris and both the anterior and medial parts of sartorius and their motoneurons all had similar bimodal excitatory responses, including an early period 3- to 18-ms poststimulus and a later period 20- to 35-ms poststimulus. The short-latency excitatory responses appeared to be proportional to the normal recruitment of the muscles in the step cycle, whereas the long-latency responses tended to be phase advanced with respect to normal recruitment. Motoneurons projecting to muscles with two excitatory peaks tended to have similar excitatory responses at both latencies and occasionally responded at both latencies to a single stimulus.

Animals↗

The distal hindlimb musculature of the cat. Cutaneous reflexes during locomotion.

In order to better understand the organization of the locomotor control system, we examined the temporal patterns of distal hindlimb muscle responses to brief electrical stimulation of cutaneous nerves during walking on a treadmill. Electromyographic recordings were made from twelve muscles; stimuli were applied individually to three nerves at random times throughout the step cycle. A new graphical technique was developed to assist detailed examination of the time course and gating of complex reflex patterns. The short latency reflexes were of two primary types: inhibition of extensors and excitation of flexors; these responses were only evident during locomotor phases in which the respective motoneuron pools were active. Longer-latency response components were gated in a similar but not identical manner, suggesting some independence from the basic locomotory influence on the motoneuronal pool. The phase-dependent gating of reflexes appeared to be consistent with a functional role for reflex responses during locomotion. The reflex responses of muscles with complex anatomical actions were often correspondingly complex.

Animals↗

Activity of spindle afferents from cat anterior thigh muscles. I. Identification and patterns during normal locomotion.

The naturally occurring activity patterns of anterior thigh muscle spindle afferents were recorded during unrestrained treadmill locomotion by means of floating microelectrodes chronically implanted in the fifth lumbar dorsal root ganglion. Conduction velocity of units from primary and secondary endings was determined by spike-triggered averaging of the signals from a chronically implanted nerve cuff. Activity from knee extensor muscle spindles generally occurred during periods of muscle lengthening, but was often greater for small stretches when the muscle was active (during stance phase of walking) than for larger stretches when the muscle was passive (swing phase), indicating fusimotor enhancement of spindle sensitivity in phase with extrafusal muscle recruitment. Activity from spindles in biarticular muscles acting across the knee and hip was more variable and complex than that seen in the pure knee extensors, and frequently included activity during rapid muscle shortening (swing phase) indicative of strong static fusimotor input. Changes in speed of gait caused changes in the range and velocity of muscle length excursions monitored by chronically implanted length gauges, but such changes were accompanied by only modest changes in spindle afferent activity, suggesting concurrent and compensatory changes in fusimotor influence on spindles. Activity from spindle secondary endings was generally lower, more regular, and less velocity dependent than that from primary endings, consistent with their lack of input from the dynamic fusimotor apparatus. The activity of all spindle afferents studied was similarly well modulated during extrafusal activity of the parent muscles, regardless of the kinematic conditions of muscle length and velocity during which this muscle work occurred. This suggests that the fusimotor apparatus is well orchestrated to regulate the static and dynamic sensitivity of primary spindle afferents at levels appropriate to the anticipated motion.

Afferent Pathways↗

Activity of spindle afferents from cat anterior thigh muscles. II. Effects of fusimotor blockade.

Chronically implanted electrodes and nerve cuff catheters were used to record the activity of individual muscle spindle afferents during treadmill walking as low doses of lidocaine were infused around the femoral nerve to progressively block gamma motoneuron activity. Both primary and secondary endings from both the monarticular knee extensors and the biarticular hip/knee muscles of the anterior thigh showed large decreases in afferent activity, usually well before changes in the electromyographic activity, force output, or length and velocity were seen in the parent muscles. This decline in the proprioceptive signal feeding back onto the spinal cord, which we presume to have involved most of the spindles supplied by the femoral nerve, did not cause noticeable irregularities or instability of the walking gait. At the peak of the fusimotor blockade, spindle afferents from knee extensor muscles lost about half of their usually brisk activity during the near-isometric contraction of the stance phase. Significant decreases in the response to passive stretch during the flexion phase also occurred. At the peak of the fusimotor blockade, spindle afferents from the biarticular muscles lost all of their activity during the rapidly shortening swing phase and about half of their activity during the rapidly lengthening stance phase. For both monarticular and biarticular muscle spindles, the activity decreases in stance and swing phase often occurred at distinctly different stages of the progressive fusimotor blockade, indicating several different sources of fusimotor control. From these data, we infer that the sensitivity of most spindle afferents is substantially influenced by fusimotor activity during phases of both extrafusal activity and extrafusal silence. At least some of this influence appears to come from fusimotor neurons whose recruitment is independent of the extrafusal recruitment. The extent and type of fusimotor effects on spindle afferent sensitivity (dynamic or static) appear to be specialized for the mechanical events that tend to occur during those phases.

Afferent Pathways↗

Activity of spindle afferents from cat anterior thigh muscles. III. Effects of external stimuli.

Chronically implanted electrodes were used to record the activity of identified single muscle spindle afferents in awake cats during responses to various types of manual and electrical stimulation. During vigorous cyclical responses such as shaking and scratching, spindle afferents generally maintained at least some activity during both lengthening and shortening of the parent muscle, indicating that the programs for these movements include both extra- and intrafusal recruitment. During noncyclical responses such as ipsilateral limb withdrawal and crossed-extension, spindle activity was modest and poorly correlated with extrafusal activity. Weak cutaneous nerve shocks during walking elicited complex excitatory and inhibitory phase-dependent reflexes in the various muscles studied but caused relatively little change in spindle afferent activity, indicating a lack of correlation between alpha and gamma motoneuron activity. A primary and a secondary afferent from sartorius muscle were recorded simultaneously during walking cycles that were perturbed by electrically induced twitches of the antagonist hamstring muscles; both demonstrated highly sensitive, short latency responses to the resulting skeletal motion, consistent with their previously suggested roles in detecting small brief mechanical perturbations. The degree to which fusimotor responses were correlated with extrafusal responses to somatosensory perturbations was highly dependent on the specific nature of the stimulus and the response. Fusimotor reprogramming of the spindle sensitivity appears to be a feature of cyclical movements that are presumably under proprioceptive control, whereas brief perturbations within the context of a particular motor program may be ignored by the fusimotor system.

Afferent Pathways↗

Motoneurone task groups: coping with kinematic heterogeneity.

Physiological principles of motor control have generally been based on linear servocontrol theories in which transducers of force, length and velocity are used to provide feedback to the motor actuators. Within the past few years, recordings of the activity of motoneurones and proprioceptors during normal motor behaviour have indicated a diversity that is not consistent with any one theory of motor control. This paper examines the heterogeneity of kinematic conditions under which muscles are called on to perform, and attempts to correlate this with the effects of various fusimotor states on the activity of the muscle spindle afferents, the major sensory signal source in most feedback control schemes. The concept of 'task groups' is proposed, in which functional groups of alpha and gamma motoneurones and spindle afferents are programmed to achieve optimal control over relatively restricted but more linear parts of their operating curves. Such a functional compartmentalization of the motor apparatus is thus consistent with several different theories of servocontrol, although it remains unclear whether such conceptual mechanisms are actually embodied in the highly complex neural circuitry present in the spinal cord and higher motor centres.

Humans↗

The distal hindlimb musculature of the cat. Patterns of normal use.

Chronic recordings were made of electromyographic (EMG) activity, tension, and length of distal hindlimb muscles in six cats performing a variety of normal motor tasks. Muscles studied thoroughly or in part were medial gastrocnemius, lateral gastrocnemius, plantaris, soleus, flexor digitorum brevis, flexor digitorum longus, flexor hallucis longus, tibialis posterior, tibialis anterior, extensor digitorum longus, peroneus longus, and peroneus brevis. Postural and locomotor activities were examined, as well as jumping, landing, scratching, and paw shaking. In general, muscles could be assigned to traditional groupings (e.g. extensor, flexor) related to the demands of the motor task. Patterns of muscle activity were most often consistent with current understanding of muscle mechanics and neural coordination. However, purely functional distinctions between flexor digitorum longus and flexor hallucis longus ("anatomical synergists") were made on the basis of activity patterns. Likewise, the activity of plantaris and flexor digitorum brevis, which are attached in series, was differentiated in certain tasks. The rhythmical oscillatory patterns of scratching and paw shaking were found to differ temporally in a manner consistent with the limb mechanics. In several cases, mechanical explanations of specific muscle activity required length and force records, as well as EMG patterns. Future efforts to study motor patterns should incorporate information about the relationships between muscle activation, tension, length and velocity.

Animals↗

Conduction velocity changes along lumbar primary afferent fibers in cats.

Though peripheral conduction velocity is widely used to characterize afferent fibers according to somatosensory modality, disagreement exists as to whether or not conduction velocity varies along such an axon's length. Therefore, in this experiment, conduction velocities were measured over very short axonal segments (7.5 to 15 mm) within the posterior tibial nerve, sciatic nerve, and L7 dorsal root, using the method of spike-triggered averaging of neurograms recorded from tripolar electrodes. The conduction velocities for several units were also determined using electrical stimulation, so that the accuracy of the two techniques could be compared. For most units, dorsal root and sciatic nerve conduction velocities were not significantly different; however, they were not tightly correlated. Tibial nerve conduction velocity averaged 86% of that within the sciatic nerve. Variations in sciatic nerve conduction velocity within adjacent axonal segments (9 mm in length) rarely exceeded experimental error. It appears that spike-triggered averaging of signals from tripolar electrodes separated 15 mm or more apart is a more accurate method for measuring conduction velocity than electrical stimulation, which was subject to several large errors.

Afferent Pathways↗

Spatial cross-correlation. A proposed mechanism for acoustic pitch perception.

We propose in this paper a new class of model processes for the extraction of spectral information from the neural representation of acoustic signals in mammals. We are concerned particularly with mechanisms for detecting the phase-locked activity of auditory neurons in response to frequencies and intensities of sound associated with speech perception. Recent psychophysical tests on deaf human subjects implanted with intracochlear stimulating electrodes as an auditory prosthesis have produced results which are in conflict with the predictions of the classical place-pitch and periodicity-pitch theories. In our model, the detection of synchronicity between two phase-locked signals derived from sources spaced a finite distance apart on the basilar membrane can be used to extract spectral information from the spatiotemporal pattern of basilar membrane motion. Computer simulations of this process suggest an optimal spacing of about 0.3-0.4 of the wavelength of the frequency to be detected. This interval is consistent with a number of psychophysical, neurophysiological, and anatomical observations, including the results of high resolution frequency-mapping of the anteroventral cochlear nucleus which are presented here. One particular version of this model, invoking the binaurally sensitive cells of the medial superior olive as the critical detecting elements, has properties which are useful in accounting for certain complex binaural psychophysical observations.

Auditory Pathways↗

Physiological characterization of motor unit properties in intact cats.

Single motor units were isolated in intact cats, by microstimulation through chronically implanted microwires in the L5 ventral roots. Motor unit axonal and mechanical properties were obtained by stimulus-triggered averaging the signals from an implanted femoral nerve recording cuff and patellar tendon force transducer. All unit types were sampled with this technique, and it was also possible to stimulate in isolation an axon whose ventral root spike was recorded during treadmill locomotion. A new technique was described, spike-triggered microstimulation, for verifying the identity of a stimulated and a recorded axon.

Animals↗

Discharge patterns of hindlimb motoneurons during normal cat locomotion.

Long-term recording from single lumbar motoneurons of intact cats revealed activation patterns fundamentally different from those seen in decerebrate preparations. In intact cats, motoneuron bursts showed marked rate modulation without initial doublets. Each unit's frequencygram generally resembled the envelope of the gross electromyogram simultaneously recorded from the corresponding muscle. Average and peak discharge rates increased for faster gaits. These findings suggest that, in cat locomotion, rate modulation is a more important contributor to force regulation than was previously thought.

Animals↗