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The mechanical action of proprioceptive length feedback in a model of cat hindlimb.

Postural regulation is an important part of a variety of motor tasks, including quiet standing and locomotion. Muscle length feedback, both the autogenic length feedback arising from a muscle's own spindles, and heterogenic length feedback, arising from its agonists and antagonists, is a strong modulator of muscle force and well suited to postural maintenance. The effects of this reflex feedback on 3-D force generation and limb mechanics are not known. In this paper, we present a mechanical model for relating 3-D changes in cat hindlimb posture to changes in muscle lengths. These changes in muscle length are used to estimate changes in both intrinsic muscle force generation and muscle activation by length feedback pathways. Few muscles are found to have directly agonist mechanical actions, and most differ by more than 20 degrees. Endpoint force fields are largely uniform across the space investigated. Both autogenic and heterogenic feedback contribute to whole limb resistance to perturbation, autogenic pathways being most dramatic. Length feedback strongly reinforced a restoring force in response to endpoint displacement.

Animals↗

Adaptation to displaced vision: visual, motor, or proprioceptive change?

After seeing his hand through wedge prisms, a subject points incorrectly with that hand at auditory as well as visual targets. The other hand is virtually unaffected. Thus the change cannot be solely visuo-motor or visual. Other evidence suggests that it is a change in felt hand location, rather than motor learning. When the subject's adapted hand feels as if it is pointing straight ahead, for example, it is actually pointing off to one side.

Acclimatization↗

Effect of warming up on knee proprioception before sporting activity.

BACKGROUND: It is now generally accepted that the ligamentous structures of the knee not only act as mechanical restraints but also have a neurophysiological role in joint function and protection. A report that knee joint laxity increases with exercise raised the question as to whether there is any compensatory change in joint position appreciation. OBJECTIVE: To test whether there is a compensatory mechanism for increased ligamentous laxity during normal levels of activity. METHODS: Joint position appreciation was measured, using a previously reported technique, in the knees of sportsmen at rest and after warm up. RESULTS: Joint position appreciation was found to be significantly more sensitive after warm up (p = 0.005). CONCLUSIONS: These findings indicate that joint position appreciation within the knee accommodates physiological changes within the ligaments and muscles after exercise.

Adolescent↗

Preservation of complete anal sphincteric proprioception in restorative proctocolectomy: the inhibitory reflex and fine control of continence need not be impaired.

This study evaluates whether reflux function of the anal sphincter remains unchanged after restorative proctocolectomy, provided that the sphincter remaining is kept intact, without mucosal stripping or endo-anal anastomosis. Paired tests of anorectal function were performed before, and a median of 6 (range 2-12) months after restorative proctocolectomy with stapled, end to end pouch-anal anastomosis. Beforehand, distension of the rectum with 50 ml of air produced a median (interquartile range) increase in pressure within the rectum of 22 (15-29) cm H2O and reflex inhibition of the anal sphincter from a pressure of 76 (62-106) cm H2O to a pressure of 34 (15-52) cm H2O. After the procedure, distension of the ileal pouch with 50 ml of air produced an increase in pressure within the pouch of only 5 (4-8) cm H2O (p < 0.001 compared with beforehand) and reflex inhibition of the anal sphincter from a pressure of 62 (25-79) cm H2O to 37 (17-68) cm H2O. Maximal reflex inhibition of the upper third if the anal sphincter to a pressure of 26 (15-48) cm H2O was observed when pressure within the pouch increased by 16 (11-22) cm H2O. After restorative proctocolectomy, all patients were continent (two experienced minor nocturnal leakage of mucus) and 25 could discriminate between flatus and faeces. Thus, reflux function was preserved in response to changes in pressure, ensuring that the subtler aspects of anal continence were preserved.

Adolescent↗

Proprioceptive pathways of the spinal cord.

In the Macaque, surgical lesions were made in the dorsal funiculus, in the dorsolateral funiculus, and through half of the spinal cord. The somatosensory and motor capacity of the animal were examined neurologically and electrophysiologically. The exact lesion was then confirmed pathologically in detail. The results of these experiments indicate that limb position information from the distal limb and proximal limb are relayed to the brain in two different fashions. Distal limb position information, especially the cortical representation of the limbs' volar surface as it moves in space, is drastically impaired by dorsal funiculus or posterior white column lesions. Proximal limb position may or may not be impaired by similar lesions, for this information while initially in the dorsal or posterior white columns is sorted out (as it ascends in the spinal cord) to the dorsolateral funiculus or white columns. For example, in the lower thoracic spinal cord, both distal and proximal hind limb sensation are impaired by posterior white column damage; in the cervical cord, only distal sensation is impaired by the same lesion, and proximal information is spared. We refer to this neuroanatomic rearranging as "fibre sorting", and we believe that it is clinically significant in spinal cord disease.

Afferent Pathways↗

Proprioceptive modulation of somatosensory evoked potentials during active or passive finger movements in man.

The effects of active and passive finger movements on somatosensory potentials evoked by stimulation of the median nerve at the wrist or of finger I were investigated in 15 healthy volunteers. As compared to the resting condition, both active and passive movements of the stimulated hand fingers induced a marked reduction in the amplitude of the primary cerebral response (N20-P25 complex) as well as of the N17 SEP component, which is supposed to reflect the activity of the thalamo-cortical radiation. The following cerebral SEP components, within 100 ms after the stimulus, were also depressed during motor activity. Neither N11 nor N13 components of the cervical response, reflecting the activation of dorsal columns and dorsal column nuclei respectively, were modified. The SEP changes induced by active or passive movements were absent after ischaemic block of large group I afferent fibers from the hand, thus suggesting the relevance of the feedback generated by such peripheral afferents during movement. The results indicate that the activation of peripheral receptors (probably muscle spindle endings) during both active and passive finger movement would induce a gating effect at both cortical and subcortical (thalamic) level, which might modulate selectively the different sensory inputs to the cortex.

Adult↗

Parietal kinetic ataxia without proprioceptive deficit.

A patient with acute onset "classic" cerebellar ataxia of the right arm without clinically detectable deep sensory loss is reported, in relation to an acute posterior parietal infarct. Wild back and forth swaying of the arm, giving away, or worsening by suppression of vision were not seen. The lesion involved area 5, parts of area 7, the angular gyrus, the middle and posterior parieto-occipital gyri, and posterior parts of the superior and middle temporal gyri. The paracentral lobule, commonly thought to be responsible for parietal ataxia, was spared. Thus posterior parietal lesions can mimick cerebellar ataxia, possibly by severing specific projections to the ventrolateral thalamic nuclei. On the basis of previous studies in primates, the superior parietal gyrus may play a major part in the ataxia presented by this patient.

Ataxia↗

Proprioceptive input from the jaw muscles and its influence on lapping, chewing, and posture.

Experiments are described which define criteria for identifying fusimotor axons discharging in filaments of the masseter nerve in lightly anaesthetized cats. During reflex movements of the jaw two patterns of discharge were observed in different fusimotor fibres. One type, called "sustained," fired at a fairly constant increased rate. The other, called "modulated," fired at high frequencies during the extrafusal muscle contraction. Evidence from spindle primary and secondary recordings in similar experiments strongly suggests that the "sustained" type were dynamic fusimotor fibres and the "modulated" type were static fusimotor fibres. New spindle recordings in normal unanaesthetized cats indicate that the modulated pattern of static fusimotor discharge also occurs in these conditions. Its effect is to reduce the tendency for spindle afferents to become silent during muscle shortening. A proposal is made that the static fusimotor discharge in cyclic movements may represent a temporal "template" for the intended movement as directed by the central pattern generator.

Animals↗

Postural adjustments associated with different unloadings of the forearm: effects of proprioceptive and cutaneous afferent deprivation.

Postural adjustments to imposed (passive) and voluntary (active) unloading conditions of the forearm were studied in normal subjects and a deafferented patient. The latency of the postural behaviour (deactivation of the biceps supporting the weight) was linearly related to the displacement amplitude of the unloaded forearm, independent of the unloading conditions. The postural behaviour consisted of an anticipatory postural adjustment (APA) occurring prior to active unloading (in both normals and the patient) and conversely in an unloading reflex response following passive unloading (only in the normals). In both the deafferented and the normal subjects, the amplitudes of the displacement during active unloadings were much smaller (3x) than in the passive conditions and an APA was present in both the deafferented and the normal subjects. The APA could not be triggered by some types of active movement and was absent when the movement was not directly producing the unloading. The EMG latencies of the APA and of the contralateral muscles used to unload were tightly coupled. However, the latency would sometimes be decoupled, particularly when a temporal delay was introduced between the active movement and the unloading in normal subjects. In contrast to the normal subjects, who were able to adapt quickly to an unusual unloading condition (produced by voluntary knee flexion), the deafferented patient did not show an APA in this task. It was concluded that, although the APA is of central origin, it cannot be generated only on the basis of internal timing cues and must rely on afferent information for its generation during unfamiliar unloading conditions.

Adult↗

Control variables and proprioceptive feedback in fast single-joint movement.

Sensorimotor mechanisms were studied on the basis of kinematic and electromyographic data as well as the static torque developed by the muscles as a function of joint angle. The latter relationship is known as the torque/angle characteristic. Fast single-joint movement may result from a shift in this characteristic and a change in its slope. Such movements were studied at the wrist in 9 normal and 1 deafferented subject. After training to flex the wrist to a target, subjects repeated the same movements but in random test trials movements were opposed by the load generated by linear position feedback to a torque motor. At the end of the loaded trials, the load was suddenly removed. In the second experiment, subjects made wrist movements to the target that were opposed by the load and, on random test trials, the movements were not loaded. In these test trials, the wrist arrived in a static position outside the target zone. In both experiments, subjects were instructed not to correct errors. The final torque/angle characteristics specified in the movements were reconstructed on the basis of the static wrist positions and torques before and after unloading. Normal subjects made movements by shifting the position of the torque/angle characteristic and by increasing its slope. If subjects indeed maintained the same pattern of control variables (descending commands), the same final position of the characteristic would be reproduced from trial to trial regardless of load perturbations. This assumption of equifinality was tested by comparing the final position of the wrist in nonloaded movements with that after removal of the load in loaded movements. Equifinality was observed in normal subjects. Movements in the deafferented subject were also associated with a shift of the torque/angle characteristic and a change in its slope. However, she was unable to consistently reproduce its final position. In spite of muscle coactivation, her maximal stiffness was lower than in normal subjects. In the absence of vision, the subject made movements with the load by increasing the slope of the characteristic instead of by shifting its position far enough. Load perturbation affected her final wrist position (inequifinality), which may reflect the presence of a significant hysteresis of the characteristic as a result of the absence of stretch reflexes. The deficits following deafferentation presumably result from the destruction of biomechanical and sensorimotor mechanisms including the ability of control variables to specify the positional frame of reference for afferent and descending systems.

Adult↗

Activity-dependent sensitivity of proprioceptive sensory neurons in the stick insect femoral chordotonal organ.

Mechanosensory neurons exhibit a wide range of dynamic changes in response, including rapid and slow adaptation. In addition to mechanical factors, electrical processes may also contribute to sensory adaptation. We have investigated adaptation of afferent neurons in the stick insect femoral chordotonal organ (fCO). The fCO contains sensory neurons that respond to position, velocity, and acceleration of the tibia. We describe the influence of random mechanical stimulation of the fCO on the response of fCO afferent neurons. The activity of individual sensory neurons was recorded intracellularly from their axons in the main leg nerve. Most fCO afferents (93%) exhibited a marked decrease in response to trapezoidal stimuli following sustained white noise stimulation (bandwidth = 60 Hz, amplitudes from +/-5 to +/-30 degrees ). Concurrent decreases in the synaptic drive to leg motoneurons and interneurons were also observed. Electrical stimulation of spike activity in individual fCO afferents in the absence of mechanical stimulation also led to a dramatic decrease in response in 15 of 19 afferents tested. This indicated that electrical processes are involved in the regulation of the generator potential or encoding of action potentials and partially responsible for the decreased response of the afferents. Replacing Ca(2+) with Ba(2+) in the saline surrounding the fCO greatly reduced or blocked the decrease in response elicited by electrically induced activity or mechanical stimulation when compared with control responses. Our results indicate that activity of fCO sensory neurons strongly affects their sensitivity, most likely via Ca(2+)-dependent processes.

Action Potentials↗

Critical points in the forelimb fictive locomotor cycle and motor coordination: evidence from the effects of tonic proprioceptive perturbations in the cat.

During locomotion, the limbs of one girdle must remain coordinated in different conditions. To understand the neural mechanisms underlying such coordination, tonic protraction/ retraction of one shoulder or tonic flexion/extension of one elbow was applied during fictive locomotion in high decerebrate and paralyzed cats. We studied bilateral changes in the timing and amplitude characteristics of electroneurographic (ENG) muscle nerve bursts of cleidobrachialis (ClB, elbow flexor and shoulder protractor) and the two heads of triceps (long, TriLo, elbow extensor and shoulder retractor and lateral, TriLa, elbow extensor). Perturbations induced bilateral changes in amplitude and timing of ENG bursts that were anchored on certain critical points in the cycle. These critical points could correspond to morphological characteristics within the bursts or to bilateral onsets or offsets of ENG bursts. For instance, in response to shoulder and elbow perturbations, burst changes occur in relation to a fixed point, labeled point C, occurring at about mid-extensor burst and corresponding to a simultaneous abrupt increase in TriLa amplitude and a decrease in amplitude of contralateral ClB. At a point labeled B, corresponding to about mid-flexor burst, ClB amplitude increases above control with elbow extension or starts decreasing with shoulder protraction. Although cycle reorganization is specific for each type of tonic perturbation, a common feature is that the changes in burst duration are achieved through discrete shifts between consecutive critical points. It is postulated that coordination may be based on a discrete temporal cycle structure along which critical points delimiting burst components are shifted.

Animals↗

Stance- and locomotion-dependent processing of vibration-induced proprioceptive inflow from multiple muscles in humans.

We performed a whole-body mapping study of the effect of unilateral muscle vibration, eliciting spindle Ia firing, on the control of standing and walking in humans. During quiet stance, vibration applied to various muscles of the trunk-neck system and of the lower limb elicited a significant tilt in whole body postural orientation. The direction of vibration-induced postural tilt was consistent with a response compensatory for the illusory lengthening of the stimulated muscles. During walking, trunk-neck muscle vibration induced ample deviations of the locomotor trajectory toward the side opposite to the stimulation site. In contrast, no significant modifications of the locomotor trajectory could be detected when vibrating various muscles of the lower as well as upper limb. The absence of correlation between the effects of muscle vibration during walking and standing dismisses the possibility that vibration-induced postural changes can account for the observed deviations of the locomotor trajectory during walking. We conclude that the dissimilar effects of trunk-neck and lower limb muscle vibration during walking and standing reflect a general sensory-motor plan, whereby muscle Ia input is processed according to both the performed task and the body segment from which the sensory inflow arises.

Action Potentials↗

Positive proprioceptive feedback elicited by isometric contractions of ankle flexors on pretibial motoneurons in cats.

Pretibial flexor motoneurons were recorded intracellularly in anesthetized cats during unfused isometric contractions of a subpopulation of motor units from either tibialis anterior (TA) or extensor digitorum longus (EDL) muscles. The contractions elicited excitatory postsynaptic potentials in 23 of 28 pretibial flexor motoneurons. No effect was observed in the remaining motoneurons. In control experiments, the effects of electrical stimulation of afferents within the TA nerve were investigated to help identify afferents responsible for the contraction-induced positive feedback. This feedback was ascribed to actions of Ia fibers because the pattern of the contraction-induced excitatory potentials was consistent with the known pattern of Ia discharge; in control experiments, electrical stimulation of group I fibers elicited only monosynaptic excitatory potentials; and the distribution of both the contraction-induced positive feedback among motor nuclei as well as the electrically evoked Ia excitatory monosynaptic potentials were restricted to homonymous and synergic motoneurons. Observation of the Ia contraction-induced positive feedback was facilitated by the absence of Ib autogenic inhibition. This contraction-induced Ia excitatory feedback in ankle flexors might either reinforce Ia-induced reflexes when these muscles are lengthened or help to lift the leg over an obstacle.

Anesthesia↗