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Effect of a tone-inhibiting dynamic ankle-foot orthosis on stride characteristics of an adult with hemiparesis.

A single-subject, alternating-treatment design was used to investigate differences in stride characteristics of a hemiparetic subject under three conditions: 1) barefoot, 2) using a prefabricated plastic molded ankle-foot orthosis (AFQ), and 3) using a tone-inhibiting dynamic ankle-foot orthosis (TIAFO). Five barefoot baseline sessions were conducted. Following these baseline sessions, the three conditions were randomly varied during each of 12 alternating-treatment sessions. Data analysis revealed a significant improvement in walking velocity, step length, and stance time on the hemiparetic limb and a significant decrease in cadence when either the AFO or the TIAFO condition was compared with the barefoot condition. The TIAFO was associated with a significant increase in walking velocity and step length when evaluated against the prefabricated AFO. The subject reported that use of the TIAFO increased his ability to ambulate distances and that the TIAFO was more comfortable and less restrictive than the prefabricated AFO. The results suggest that the TIAFO may be a promising alternative to conventional orthotic management of adults with hemiparesis.

Adult↗

Subjective straight-ahead during neck muscle vibration: effects of ageing.

The subjective visual straight-ahead (SVA) was measured psychophysically before and during unilateral vibration of the posterior neck muscles in order to determine the particular contribution of neck muscle spindles to the perception of body orientation during ageing. We found a symmetrical increase of the vibration-induced displacement of the SVA with advancing age (R = +0.73, p < 0.01, n = 60) in 30 healthy subjects of different ages (range 20-81 years). These data indicate the cervical proprioceptive contribution to multisensory body orientation increases with ageing. One possible interpretation is that the sensorial weight of proprioception increases as the peripheral vestibular input decreases with advancing age, thereby substituting for the lack of vestibular function.

Adult↗

The functions of the proprioceptors of the eye muscles.

This article sets out to present a fairly comprehensive review of our knowledge about the functions of the receptors that have been found in the extraocular muscles--the six muscles that move each eye of vertebrates in its orbit--of all the animals in which they have been sought, including Man. Since their discovery at the beginning of the 20th century these receptors have, at various times, been credited with important roles in the control of eye movement and the construction of extrapersonal space and have also been denied any function whatsoever. Experiments intended to study the actions of eye muscle receptors and, even more so, opinions (and indeed polemic) derived from these observations have been influenced by the changing fashions and beliefs about the more general question of how limb position and movement is detected by the brain and which signals contribute to those aspects of this that are perceived (kinaesthesis). But the conclusions drawn from studies on the eye have also influenced beliefs about the mechanisms of kinaesthesis and, arguably, this influence has been even larger than that in the converse direction. Experimental evidence accumulated over rather more than a century is set out and discussed. It supports the view that, at the beginning of the 21st century, there are excellent grounds for believing that the receptors in the extraocular muscles are indeed proprioceptors, that is to say that the signals that they send into the brain are used to provide information about the position and movement of the eye in the orbit. It seems that this information is important in the control of eye movements of at least some types, and in the determination by the brain of the direction of gaze and the relationship of the organism to its environment. In addition, signals from these receptors in the eye muscles are seen to be necessary for the development of normal mechanisms of visual analysis in the mammalian visual cortex and for both the development and maintenance of normal visuomotor behaviour. Man is among those vertebrates to whose brains eye muscle proprioceptive signals provide information apparently used in normal sensorimotor functions; these include various aspects of perception, and of the control of eye movement. It is possible that abnormalities of the eye muscle proprioceptors and their signals may play a part in the genesis of some types of human squint (strabismus); conversely studies of patients with squint in the course of their surgical or pharmacological treatment have yielded much interesting evidence about the central actions of the proprioceptive signals from the extraocular muscles. The results of experiments on the eye have played a large part in the historical controversy, now in at least its third century, about the origin of signals that inform the brain about movement of parts of the body. Some of these results, and more of the interpretations of them, now need to be critically re-examined. The re-examination in the light of recent experiments that is presented here does not support many of the conclusions confidently drawn in the past and leads to both new insights and fresh questions about the roles of information from motor signals flowing out of the brain and that from signals from the peripheral receptors flowing into it. There remain many lacunae in our knowledge and filling some of these will, it is contended, be essential to advance our understanding further. It is argued that such understanding of eye muscle proprioception is a necessary part of the understanding of the physiology and pathophysiology of eye movement control and that it is also essential to an account of how organisms, including Man, build and maintain knowledge of their relationship to the external visual world. The eye would seem to provide a uniquely favourable system in which to study the way in which information derived within the brain about motor actions may interact with signals flowing in from peripheral receptors. The review is constructed in relatively independent sections that deal with particular topics. It ends with a fairly brief piece in which the author sets out some personal views about what has been achieved recently and what most immediately needs to be done. It also suggests some lines of study that appear to the author to be important for the future.

Afferent Pathways↗

The concerted activity in parallel proprioceptive pathways controls the initiation of co-activation in the locust kick motor programme.

To jump and kick the locust uses a catapult mechanism implemented by a three-stage motor programme: initial flexion of the hind tibiae, co-activation of the antagonist flexor and extensor tibiae motor neurons and trigger inhibition of the flexor motorneurons. The transition from stage 1 to stage 2 thus involves a switch from the normal alternate activation to co-activation of the antagonist tibrae motorneurons. However, co-activation has never been observed when the central nervous system has been isolated from the leg. This led us to investigate the possibility that the transition from stage 1 to stage 2 is controlled by a proprioceptive signal. In the first set of experiments intracellular recordings were made in the flexor and extensor motorneurons while the position of the tendon of the femoral chordotonal organ (FCO), which signals tibial position and movement, was experimentally controlled. In these heavily dissected preparations, stretch of the FCO tendon (signalling tibial flexion) was a necessary condition for co-activation. However, in minimally dissected preparations (in which merely EMG recordings were made), we found that co-activation occurred even when the FCO was signalling tibial extension, suggesting the involvement of other proprioceptors. A series of experiments were then conducted on minimally dissected preparations to determine the relative contributions of each of the three main hind leg proprioceptors which might signal tibial flexion: the FCO, the lump receptor and Brünners organ. When all three proprioceptors were intact the chance of evoking co-activation was largest, when all three were eliminated co-activation could no longer be evoked, irrespective of the level of arousal. Various combinations of partial de-afferentation showed that the FCO plays the major role, with the lump receptor and Bünners organ playing significant, but progressively less important, roles. We conclude that the three receptors act together as a permissive proprioceptive gate for the kick and jump motor programme, but with a hierarchy of the strengths of their effectiveness.

Animals↗

Neuronal generation of the leech swimming movement.

The swimming movement of the leech is produced by an ensemble of bilaterally symmetric, rhythmically active pairs of motor neurons present in each segmental ganglion of the ventral nerve cord. These motor neurons innervate the longitudinal muscles in dorsal or ventral sectors of the segmental body wall. Their duty cycles are phase-locked in a manner such that the dorsal and ventral body wall sectors of any given segment undergo an antiphasic contractile rhythm and that the contractile rhythms of different segments form a rostrocaudal phase progression. This activity rhythm is imposed on the motor neurons by a central swim oscillator, of which four bilaterally symmetric pairs of interneurons present in each segmental ganglion appear to constitute the major component. These interneurons are linked intra- and intersegmentally via inhibitory connections to form a segmentally iterated and inter-segmentally concatenated cyclic neuronal network. The network appears to owe its oscillatory activity pattern to the mechanism of recurrent cyclic inhibition.

Action Potentials↗

Absence of enhanced physiological tremor in patients without muscle or cutaneous afferents.

Patients with proprioceptive loss due to a neuropathy affecting large myelinated sensory nerve fibres were studied to determine the role of somaesthetic sensory inputs in enhanced physiological tremor. Involuntary movements in patients and controls attempting to hold the outstretched arm immobile were recorded during prolonged arm extension. Fatigue led to increased movements in both controls and patients but only the controls developed a rhythmic tremor. These data indicate that enhanced physiological tremor is dependent on somaesthetic afferent input.

Aged↗

Proprioception during voluntary movement.

In the last decade, a number of laboratories have accumulated data on the firing of single afferent fibres from muscle and skin during movement in awake cats, monkeys and human subjects. While there is general agreement on the firing behaviour of skin afferents and tendon organ (Ib) afferents during movement, there remains a significant divergence of opinion regarding the way in which the response of muscle spindle afferents (Ia and II) to length changes is modified by fusimotor action (e.g., alpha-gamma linkage versus "fusimotor set"). The controversies surrounding the fusimotor system have tended to overshadow the emergence of several important characteristics of proprioceptive behaviour, corroborated in separate laboratories. (i) Mean Ia firing rates during active movements are nearly always higher than at rest. Thus, activation of the fusimotor system is reserved for the control of, or preparation for, movement. In animals, there is now strong evidence that there is usually a tonic component of fusimotor action during rhythmical movements. (ii) During fast, unloaded movements (peak muscle speeds, 0.2 resting lengths/s or more), the firing of both Ia and II afferents usually increases during lengthening and decreases during shortening. Ib afferents fire during even the most rapid active shortening of their parent muscles. (iii) During powerful shortening contractions performed against significant loads, Ia firing is often appreciable, suggesting that there is at least some underlying alpha-gamma coactivation. (iv) During fast imposed muscle stretches, Ia afferents respond with segmented bursts of firing (threshold speed for segmentation, 0.5-1.0 resting length/s). Ib afferents show far less segmentation of discharge under similar circumstances.(ABSTRACT TRUNCATED AT 250 WORDS)

Afferent Pathways↗

Vestibular, proprioceptive, and haptic contributions to spatial orientation.

The control and perception of body orientation and motion are subserved by multiple sensory and motor mechanisms ranging from relatively simple, peripheral mechanisms to complex ones involving the highest levels of cognitive function and sensory-motor integration. Vestibular contributions to body orientation and to spatial localization of auditory and visual stimuli have long been recognized. These contributions are reviewed here along with new insights relating to sensory-motor calibration of the body gained from space flight, parabolic flight, and artificial gravity environments. Recently recognized contributions of proprioceptive and somatosensory signals to the appreciation of body orientation and configuration are described. New techniques for stabilizing posture by means of haptic touch and for studying and modeling postural mechanisms are reviewed. Path integration, place cells, and head direction cells are described along with implications for using immersive virtual environments for training geographic spatial knowledge of real environments.

Cognition↗

Cutaneous receptors contribute to kinesthesia at the index finger, elbow, and knee.

The neural mechanisms underlying the sense of joint position and movement remain controversial. While cutaneous receptors are known to contribute to kinesthesia for the fingers, the present experiments test the hypothesis that they contribute at other major joints. Illusory movements were evoked at the interphalangeal (IP) joints of the index finger, the elbow, and the knee by stimulation of populations of cutaneous and muscle spindle receptors, both separately and together. Subjects matched perceived movements with voluntary movements of homologous joints on the contralateral side. Cutaneous receptors were activated by stretch of the skin (using 2 intensities of stretch) and vibration activated muscle spindle receptors. Stimuli were designed to activate receptors that discharge during joint flexion. For the index finger, vibration was applied over the extensor tendons on the dorsum of the hand, to evoke illusory metacarpophalangeal (MCP) joint flexion, and skin stretch was delivered around the IP joints. The strong skin stretch evoked the illusion of flexion of the proximal IP joint in 6/8 subjects (12 +/- 5 degrees, mean +/- SE). For the group, strong skin stretch delivered during vibration increased the perceived flexion of the proximal IP joint by eight times with a concomitant decrease in perceived flexion of the MCP joint compared with vibration alone (P < 0.05). For the elbow, vibration was applied over the distal tendon of triceps brachii and skin stretch over the dorsal forearm. When delivered alone, strong skin stretch evoked illusory elbow flexion in 5/10 subjects (9 +/- 4 degrees). Simultaneous strong skin stretch and vibration increased the illusory elbow flexion for the group by 1.5 times compared with vibration (P < 0.05). For the knee, vibration was applied over the patellar tendon and skin stretch over the thigh. Skin stretch alone evoked illusory knee flexion in 3/10 subjects (8 +/- 4 degrees) and when delivered during vibration, perceived knee flexion increased for the group by 1.4 times compared with vibration (P < 0.05). Hence inputs from cutaneous receptors, muscle receptors, and combined inputs from both receptors likely subserve kinesthesia at joints throughout the body.

Adult↗

Slow dynamics of postural sway are in the feedback loop.

Postural sway is considered to have two fundamental stochastic components, a slow nonoscillatory component and a faster damped-oscillatory component. The slow component has been shown to account for the majority of sway variance during quiet stance. Postural control is generally viewed as a feedback loop in which sway is detected by sensory systems and appropriate motor commands are generated to stabilize the body's orientation. Whereas the mechanistic source for the damped-oscillatory sway component is most likely feedback control of an inverted pendulum, the underlying basis for the slow component is less clear. We investigated whether the slow process was inside or outside the feedback loop by providing standing subjects with sum-of-sines visual motion. Linear stochastic models were fit to the experimental sway trajectories to determine the stochastic structure of sway as well as the transfer function from visual motion to sway. The results supported a fifth-order stochastic model, consisting of a slow process and two damped-oscillatory components. Importantly, the slow process was determined to be inside the feedback loop. This supports the hypothesis that the slow component is due to errors in state estimation because state estimation is inside the feedback loop rather than a moving reference point or an exploratory process outside the feedback loop.

Adult↗

Reversal of the influence of group Ib afferents from plantaris on activity in medial gastrocnemius muscle during locomotor activity.

1. Rhythmic locomotor activity was evoked in clonidine-treated acute and chronic spinal cats, and the effect of stimulating group I afferents from the plantaris muscle on the timing and magnitude of bursts in medial gastrocnemius (MG) motoneurons was examined. 2. The locomotor rhythm was entrained when group I afferents in the plantaris nerve were electrically stimulated with trains of stimuli presented at rates above and below the intrinsic frequency of the rhythmic activity. During entrainment at rates higher than the intrinsic frequency, a burst of activity in ipsilateral MG motoneurons was initiated approximately 40 ms after the onset of each stimulus train. At lower rates of entrainment the onset of MG bursts preceded the onset of the stimulus trains, and each stimulus train had an excitatory effect on the MG burst with a latency in the range of 30-50 ms. A similar excitatory effect was observed when the stimulus trains were triggered at a preset delay after the endogenous generation of the MG bursts. 3. The excitatory action of plantaris group I afferents on the MG motoneurons was only seen during periods of locomotor activity. In the absence of rhythmic activity, the same stimulus trains reduced any ongoing tonic activity in MG motoneurons. 4. Vibration of the plantaris muscle to preferentially activate group Ia afferents neither entrained the locomotor rhythm nor increased the magnitude of the MG bursts. 5. We conclude that during locomotor activity, input from group Ib afferents of the plantaris muscle has an excitatory action on the system of interneurons generating the extensor bursts, i.e., on the extensor half-center of the central rhythm generator.(ABSTRACT TRUNCATED AT 400 WORDS)

Afferent Pathways↗

Intersegmental and local interneurons in the metathorax of the stick insect Carausius morosus that monitor middle leg position.

1. In the stick insect, proprioceptive information from the middle leg is used to define the target for the swing movement of the adjacent rear leg ("targeting behavior"). To investigate the underlying neural circuits, intracellular recordings were made in the ganglion controlling the rear leg, the metathoracic ganglion, while systematically moving the tarsus of the middle leg. 2. Several intersegmental interneurons and one local interneuron were identified as possible contributors to the targeting behavior. The intersegmental interneurons code the position of the middle leg tarsus in a highly simplified manner: test movements of the middle leg in the dorsal, lateral, and caudal directions from the standard starting position at right angles to the thorax elicit phasic-tonic responses in three different intersegmental neurons. The response in each interneuron actually reflects the movement and position at only one joint of the middle leg: for the neurons responding primarily to movement in the caudal, dorsal, and lateral test directions, the adequate stimulus is movement at the subcoxal joint, the coxa-trochanter joint, and the femur-tibia joint, respectively. 3. The metathoracic local interneuron integrates information from ipsilateral middle and rear legs in such a way as to provide an approximate measure of the distance between the two tarsi in the longitudinal direction. It is depolarized in a phasic-tonic manner both by caudal movements of the ipsilateral middle leg and by rostral movements of the ipsilateral rear leg. The adequate stimulus in each case is the change in the angle at the subcoxal joint of the leg moved. Depolarization of this neuron activates retractor motoneurons, which is consistent with a role in terminating the swing movement. 4. Altogether the results indicate first, that the targeting behavior could be controlled by very few intersegmental channels and, second, that the nervous system encodes the position of the middle leg tarsus in terms of joint angles rather than in abstract, body-centered coordinates.

Animals↗

Properties of sensory neurons innervating synovial joints.

While it is clear that the TMJ joint has a sensory innervation like other synovial joints, there is little specific information about the function of the innervation of the TMJ. In order to provide a template upon which future studies might build, this article briefly reviews the function of sensory neurons in other synovial joints.

Afferent Pathways↗