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Fusimotor control of proprioceptive feedback during locomotion and balancing: can simple lessons be learned for artificial control of gait?

The possibilities for central control of primary spindle afferents through fusimotor efferents for gain control in motor control mechanisms are briefly reviewed. While the existence of separate pathways for independent control of static and dynamic gamma-motoneurones is well established, it proved more difficult to demonstrate that gain control of spindle feedback, attributable to alterations in static and dynamic fusimotor drive, indeed took place in voluntary movements. However, earlier qualitative indications, that Ia sensitivity (and hence the balance of static over dynamic drive) was adjusted differently in different motor tasks, have recently been confirmed in experimental simulation studies, in which the fusimotor activation profiles, that were required to reproduce chronically recorded spindle Ia discharge patterns, were reconstructed. These studies indicated that Ia sensitivity and dynamic gamma-drive were low in routine movements (walking), but that they could be dramatically increased in motor tasks which were either difficult or unfamiliar (landing from falls, balancing on narrow walk beams, adjustment to imposed disturbances). This suggested that sensitization of spindle feedback could play a significant role in motor adaptation. In line with this, studies in patients with large fibre sensory (including proprioceptive) neuropathies indicated that long-term motor deficits (affecting motor adaptation and learning) could be at least as serious as short-term motor dysfunction (due to loss of reflex control). It is suggested that spindle Ia feedback may play a dual role: in addition to its contribution to short-term reflex control of posture and movement, it may also be used for optimization or maintenance of motor programs, especially if its gain is increased by significant dynamic fusimotor drive.(ABSTRACT TRUNCATED AT 250 WORDS)

Afferent Pathways↗

Full-body gaze control mechanisms elicited during locomotion: effects of VOR adaptation.

We have previously shown that multiple, interdependent, full- body sensorimotor subsystems aid gaze stabilization during locomotion. In the present study we investigated how the full-body gaze control system responds following exposure to visual-vestibular conflict known to adaptively modify vestibulo-ocular reflex (VOR) function. Subjects (n = 14) walked (6.4 km/h) on a motorized treadmill before and after they were exposed to 0.5X minifying lenses worn for 30 minutes during self-generated sinusoidal vertical head rotations performed while seated. Results indicate that, following the exposure the major changes that subjects showed were to: 1) decrease the amplitude of head pitch and vertical translation of the torso movement with respect to space; 2) increase the amount of knee and ankle flexion during the initial stance phase of the gait cycle. A correlation analysis showed that: 1) changes in the head pitch significantly co-varied with that of the vertical torso translation 2) changes in the knee flexion significantly co-varied with that of the ankle flexion during the initial stance phase of the gait cycle 3) changes in the vertical torso translation significantly co-varied with that of the ankle flexion during the initial stance phase of the gait cycle. Thus we infer that the changes in the magnitude after VOR adaptation in comparison to their pre adaptation responses serve to aid gaze stabilization during locomotion. The significant covariation of the changes between subsystems provides further evidence that the full body contributes to gaze stabilization during locomotion, and its different functional elements are subject to adaptive reorganization following exposure to visual-vestibular conflict.

Adaptation, Physiological↗

Horizontal vestibulo-ocular reflex and head stability in response to torso perturbations during visual search.

Eye, head, and torso movements were recorded using magnetic search coils while six normal human subjects made unconstrained eye and head movements as they searched for targets in a panoramic visual environment. Torso movements were imposed by pseudorandom rotations of a servomotor-driver chair in which subjects were seated; body motion was partially transmitted to the head as a perturbation. Horizontal vestibulo-ocular reflex (VOR) gain (eye velocity divided by head velocity) and head gain (head velocity divided by torso velocity) were determined. Measurements were performed with unaided vision and while subjects wore x4 binocular telescopic spectacles. Since the head was free to move during the experiment, much of the perturbation delivered to the torso was compensated by head rotation on the neck. During the 50 ms immediately following chair rotation, the head corrected 98% of the torso motion. For the interval 50-80 ms after the perturbation 81-85% of the perturbation was corrected by head movement. The degree of head compensation did not significantly depend on magnification or type of visual target. The density distribution for VOR gain was calculated over the entire course of each trial and was found to be sharply centered between 0.9 and 1.0 for trials with unmagnified vision. The gain density distribution with x4 telescopes was broader and centered around 1.5, reflecting visual enhancement. Gain of the VOR was also determined during four discrete epochs covering the period from 50 ms before to 130 ms after the onset of each imposed torso rotation. The first, second, and fourth epochs were 50 ms each, while the third epoch was 30 ms. The torso began to rotate in the second epoch (0-50 ms), and the onset of head rotation was in the third epoch (50-80 ms). Gains of the VOR determined during the first three epochs were in response to self-generated head rotation and were not significantly different from each other, averaging 1.0+/-0.4 (n=1604, mean+/-SD) with unaided vision and increased significantly (P<0.05) to 1.4+/-0.6 (n=2464) with telescopic spectacles. Gain of the VOR during the fourth (80-130 ms) epoch was in response to the imposed perturbation; this averaged 0.9+/-0.3 (n=1380) with unaided vision and increased significantly to 1.1+/-0.4 (n=2185) with telescopic spectacles. The wearing of telescopic spectacles thus induced an enhancement of VOR gain, which was dependent on the context of the associated head movement. The greater enhancement of VOR gain during self-generated head movement suggests that the large enhancement may be at least partially mediated by the motor program itself. However, the smaller, but still significant gain enhancement with telescopic spectacles observed during unpredictable, externally imposed head motion had a latency too short to be mediated by visual pursuit. We propose that the smaller gain enhancement during passive rotation is due to a small, context-dependent, parametric increase in the gain of canal or proprioceptive mediated eye movements.

Acceleration↗

Injections of beta-noradrenergic substances in the cerebellar anterior vermis of cats affect adaptation of the vestibulospinal reflex gain.

In precollicular decerebrate cats, intermittent sinusoidal roll tilt of the whole animal (at 0.15 Hz, +/- 10 degrees) produced a vestibulospinal reflex (VSR), characterized by an increased EMG activity of the forelimb extensor triceps brachii during side-down and a decreased activity during side-up tilt. This reflex was first tested during and after a 3-h period of sustained animal tilt at the same parameters indicated above. An adaptive increase in gain of the VSR progressively developed in some experiments, but not in others. In a second group of experiments, however, sinusoidal roll tilt of the head (0.15 Hz, +/- 10 degrees) was associated with a synchronous roll tilt of the body (at 0.15 Hz, +/- 12.5 degrees). This additional stimulus led to 2.5 degrees of neck rotation, which was thus out of phase with respect to head rotation. In all these experiments, submitted to a 3-h period of sustained neck-vestibular stimulation, the gain of the VSR tested every 10-15 min consistently increased to reach the maximum at the end of the third hour of stimulation. This adaptive process was followed up to 1 h after stimulation. Microinjection into the hemivermal cortex of the cerebellar anterior lobe of the beta-noradrenergic antagonists propranolol or sotalol (0.25-0.50 microliter at 8 micrograms/microliter saline) produced only slight and short-lasting changes in the basic amplitude of the VSR, but always decreased or prevented the occurrence of the adaptive increase in gain of the VSR during sustained out of phase head-neck rotation. The same agents also suppressed the increase in gain of the VSR which occurred in some experiments during sustained roll tilt of the whole animal (at 0.15 Hz, +/- 10 degrees), leading to selective stimulation of labyrinth receptors. On the other hand, the beta-noradrenergic agonist isoproterenol (0.25 microliters at 8 micrograms/microliters saline) brought to the light the adaptive process in other experiments in which no adaptation occurred during sustained animal rotation. These effects occurred when the sites of injection were located within the zone B of the cerebellar anterior vermis, which projects to the lateral vestibular nucleus. In conclusion, the results indicate that the adaptive changes affecting the gain of the VSR in decerebrate cats are facilitated by the noradrenergic afferent system acting on the cerebellar vermis through beta-adrenoceptors.

Adaptation, Physiological↗

Modulation of the masseteric reflex by gastric vagal afferents.

Several investigations have shown that the vagal nerve can affect the reflex responses of the masticatory muscles acting at level either of trigeminal motoneurons or of the mesencephalic trigeminal nucleus (MTN). The present experiments have been devoted to establish the origin of the vagal afferent fibres involved in modulating the masseteric reflex. In particular, the gastric vagal afferents were taken into consideration and selective stimulations of such fibres were performed in rabbit. Conditioning electrical stimulation of truncus vagalis ventralis (TVV) reduced the excitability of the MTN cells as shown by a decrease of the antidromic response recorded from the semilunar ganglion and elicited by MTN single-shock electrical stimulation. Sympathetic and cardiovascular influences were not involved in these responses. Mechanical stimulation of gastric receptors, by means of gastric distension, clearly diminished the amplitude of twitch tension of masseteric reflex and inhibited the discharge frequency of proprioceptive MTN units. The effect was phasic and depended upon the velocity of distension. Thus the sensory volleys originating from rapid adapting receptors reach the brain stem through vagal afferents and by means of a polysynaptic connection inhibits the masseteric reflex at level of MTN cells.

Animals↗

Postural coactivation and adaptation in the sway stabilizing responses of normals and patients with bilateral vestibular deficit.

The experiments were designed to test two hypotheses and their corollaries: 1. That adaptation of EMG responses to support surface rotations is due to a decrease in the gain of proprioceptively triggered long-loop stretch reflexes (Nashner 1976), and that the adaptation is dependent on a normally functioning vestibular system (Nashner et al. 1982); 2. That EMG responses to rotations are generated primarily by vestibulo-spinal reflexes triggered by head accelerations (Allum and Pfaltz 1985) and comprise a coactivation of opposing leg muscles (Allum and Büdingen 1979). Adaptation with successive dorsi-flexive rotations of the support surface was investigated in the EMG responses of the ankle muscles, soleus (SOL) and tibialis anterior (TA), as well as the neck muscles, trapezius (TRAP) and splenius capitis (SPLEN CAP), both for normal subjects and for patients with bilateral peripheral vestibular deficit. Both normals and patients who first received the stimulus with their eyes open demonstrated decreasing activation at medium latency (ML), that is, with an onset at about 125 ms, and long latency (LL) responses with an onset ca 200 ms. This was the case for both ankle and neck muscles when the EMG response areas for the first 3 and second 7 of 10 trials were compared. Ankle muscle responses in the patients were diminished in area with respect to normals both with the eyes open and with the eyes closed. Ankle torque recordings from the patients were also smaller in amplitude, and these attenuated differently from normal torque responses. Functional coupling of the opposing ML and LL SOL and TA muscle responses was confirmed by the nearly coincident onset times and significantly correlated EMG response areas. At ML, ankle torque was highly correlated with TA activity when the influence of SOL was controlled. At LL, SOL activity was highly correlated with torque when the influence of TA was controlled. The delay of torque adaptation beyond the period of ML activity in normals, but not in the patients was attributed to the proportionally balanced coactivated muscle patterns producing a consistent force output and level of stability in normals. The results indicate that the adaptation in EMG response amplitudes during a sway stabilisation task is not dependent on a normally functioning vestibular system nor on visual inputs but rather appears to be due to a generalized habituation in the postural control system.(ABSTRACT TRUNCATED AT 400 WORDS)

Adolescent↗

Feedforward activity of the cervical flexor muscles during voluntary arm movements is delayed in chronic neck pain.

The objective of this study was to compare onset of deep and superficial cervical flexor muscle activity during rapid, unilateral arm movements between ten patients with chronic neck pain and 12 control subjects. Deep cervical flexor (DCF) electromyographic activity (EMG) was recorded with custom electrodes inserted via the nose and fixed by suction to the posterior mucosa of the oropharynx. Surface electrodes were placed over the sternocleidomastoid (SCM) and anterior scalene (AS) muscles. While standing, subjects flexed and extended the right arm in response to a visual stimulus. For the control group, activation of DCF, SCM and AS muscles occurred less than 50 ms after the onset of deltoid activity, which is consistent with feedforward control of the neck during arm flexion and extension. When subjects with a history of neck pain flexed the arm, the onsets of DCF and contralateral SCM and AS muscles were significantly delayed ( p<0.05). It is concluded that the delay in neck muscle activity associated with movement of the arm in patients with neck pain indicates a significant deficit in the automatic feedforward control of the cervical spine. As the deep cervical muscles are fundamentally important for support of the cervical lordosis and the cervical joints, change in the feedforward response may leave the cervical spine vulnerable to reactive forces from arm movement.

Adult↗

Expected and unexpected head yaw movements result in different modifications of gait and whole body coordination strategies.

During locomotion we routinely make voluntary head movements, similar to those made during steering tasks, in order to scan our environment and obtain information about objects in the environment and our proximity to these objects. Given the importance that head segment orientation during locomotion has received in the recent literature, two studies were designed to investigate responses following a voluntarily generated and an unexpected, externally applied head turn. During a voluntary head turn, an efferent copy of the head movement could cancel the sensory effects of the head turn, effectively isolating the movement response to that segment. Alternatively, if the steering synergy is a part of our motor repertoire, as has been suggested, movement of the head could automatically release a steering "synergy" of segmental control and coordination. A unique head mounted air-jet apparatus, designed and developed at the University of Waterloo, was used for both studies to ensure that auditory stimuli and the physical presence of the apparatus on the head were similar for participants of the two experiments. During certain points in the gait cycle, this device was triggered and a short burst of compressed air (350 ms) was released to cue participants to make a voluntary head turn (Experiment 1). The same device was triggered in Experiment 2; however, in this experiment compressed air was released for a longer duration (1,500 ms) which resulted in an unexpected and quick turn of the participants' head to either the left or right. In these experiments, vision was also manipulated in certain trials with liquid crystal display glasses that occluded vision for the duration of the head turn. Data from the first experiment indicates that a subset of the steering synergy previously observed is released following the voluntary head movement; however, the travel trajectory path is preserved, suggesting that sensory input resulting from the head movement is partially nullified by the central nervous system. Overall safety is ensured by maintaining the same travel path. In the second experiment, an unexpected perturbation was applied to the head during locomotion to determine how the absence of an efferent copy of the movement pattern influences the level of control over body segments during locomotion. Whole body responses similar to those observed during steering tasks were observed following application of this unexpected head perturbation. It is proposed that the CNS interprets an unexpected yaw movement of the head as a change in the frame of reference, and global modifications of the walking trajectory, similar to that observed during steering tasks, are made in the perceived new direction of travel. Collectively this work extends our understanding of how the CNS establishes a head based orientation frame for locomotion. The CNS interprets and integrates anticipated and unexpected changes in sensory information from the head segment and subsequently modifies locomotion patterns according to the perceived whole body orientation in space. The sequence of control following these head movements appears to be part of a movement repertoire that is not immutable; maintaining whole body stability during locomotion is paramount.

Acoustic Stimulation↗

Perception of angular displacement without landmarks: evidence for Bayesian fusion of vestibular, optokinetic, podokinesthetic, and cognitive information.

The perception of angular displacement during self turning is generally based on a combination of redundant signals from different sources. For example, during active turning in a visually structured environment devoid of landmarks, podokinesthetic, vestibular, and optokinetic velocity signals are fused and integrated over time to yield a unitary percept of the ongoing change in angular position ('podokinesthetic' refers to proprioceptive and corollary signals related to leg and foot movement). Previously we have shown that the fusion of two of these afferents improves perceptual accuracy and reliability in comparison to when only one is available. For example, with only a single modality available, slow rotations are perceived to be significantly larger than fast ones, whereas the combination of two modalities greatly reduces this difference. These observations spurred the hypothesis that displacement perception results from a weighted average of bottom-up (sensory) signals and top-down signals (a priori knowledge or expectation), with the weight of the latter decreasing the more sensory information is available. We now ask (1) whether the accuracy of angular displacement estimation can be further improved if it can draw on all three sensory modalities instead of only two, and (2) whether bottom-up sensory and top-down a priori information is combined for displacement estimation in a statistically optimal way. To this end 12 healthy subjects (Ss) standing on a turning platform surrounded by a rotatable optokinetic pattern were exposed to 6 different sensory conditions: pure podokinesthetic (P), vestibular (V), or optokinetic (O) stimulation, and combined podokinesthetic-vestibular (PV), vestibular-optokinetic (VO), or podokinesthetic-vestibular-optokinetic (PVO) stimulation. Stimuli had constant angular velocities of either 15, 30, or 60 degrees /s. Subjects were to press a signal button when they felt that angular displacement had reached a previously instructed magnitude (150-900 degrees ). In agreement with earlier observations, the combination of two sensory signals improved the accuracy of displacement perception by reducing both the variance of subjects' displacement estimates and their dependence on turning velocity. Adding a third sensory signal (condition PVO) led to a further reduction of variance and almost eliminated the effect of velocity. We show that these experimental results are compatible with a probabilistic fusion mechanism based on Bayes' law. This mechanism would operate on logarithmic representations of turning velocity and proceed in two stages. A first stage fuses all available bottom-up information to create a unitary representation of the velocity signalled by the different sensory modalities. A second stage then fuses this sensory information with top-down a priori information; the latter creates a bias in favour of a 'default velocity' that grows as the uncertainty of the sensory information increases. Our experimental data agree with the relation between (1) the variance of displacement estimates and (2) their modulation by velocity predicted by this scheme.

Adolescent↗

Neck proprioceptive inputs to primate vestibular nucleus neurons.

The contribution of neck proprioceptive signals to signal processing in the vestibular nucleus was studied by recording responses of secondary horizontal canal-related neurons to neck rotation in the squirrel monkey. Responses evoked by passive neck rotation while the head was held stationary in space were compared with responses evoked by passive whole body rotation and by forced rotation of the head on the trunk. Most neurons (76%; 45/59) were sensitive to neck rotation. The nature and strength of neck proprioceptive inputs varied and usually combined linearly with vestibular inputs. In most cases (94%), the direction of the neck proprioceptive input was "antagonistic" or "reciprocal" with respect to vestibular sensitivity and, consequently, reduced the vestibular response during head-on-trunk rotation. Different types of vestibular neurons received different types of proprioceptive input. Neurons whose firing behavior was related to eye position (position-vestibular-pause neurons and position-vestibular neurons) were often sensitive to the position of the head with respect to the trunk. The sensitivity to head position was usually in the same direction as the neuron's eye position sensitivity. Non-eye-movement related neurons and eye-head-velocity neurons exhibited the strongest sensitivity to passive neck rotation and had signals that were best related to neck velocity. The results suggest that neck proprioceptive inputs play an important role in shaping the output of the primate vestibular nucleus and its contribution to posture, gaze and perception.

Animals↗

The influence of vision on the automatic postural muscle responses of newly standing and newly walking infants.

In adults, visual inputs do not appear to contribute significantly to automatic postural muscle responses (90-100 ms latency) activated by transient support surface displacements causing threats to standing balance, but are activated through slow pathways with latencies of more than 200 ms. However, it has been shown that the postural sway behavior of early walking infants is strongly influenced by visual flow cues that falsely signal self-movement. To determine whether there also are significant contributions of vision to automatic postural muscle responses in this age group, two groups of infants were tested on a moveable platform; pre-walkers (n=6) and early walkers (n=6). Pre-walkers did not show any measurable effect of visual condition (vision vs no vision) on muscle response characteristics. However, the integrated gastrocnemius activity of early walkers increased significantly in vision versus no vision conditions (P<0.05). These results show that visual cues contribute to, or modulate, the automatic postural responses in children who are in the developmental transition to independent walking.

Cues↗

Antagonist motor responses correlate with kinesthetic illusions induced by tendon vibration.

In humans, vibration applied to muscle tendons evokes illusory sensations of movement that are usually associated with an excitatory tonic response in muscles antagonistic to those vibrated (antagonist vibratory response or AVR). The aim of the present study was to investigate the neurophysiological mechanisms underlying such a motor response. For that purpose, we analyzed the relationships between the parameters of the tendon vibration (anatomical site and frequency) and those of the illusory movement perceived (direction and velocity), as well as the temporal, spatial, and quantitative characteristics of the corresponding AVRs (i.e., surface EMG, motor unit firing rates and activation latencies). Analogies were supposed between the characteristics of AVRs and voluntary contractions. The parameters of the AVR were thus compared with those of a voluntary contraction with similar temporal and mechanical characteristics, involving the same muscle groups as those activated by vibration. Wrist flexor muscles were vibrated either separately or simultaneously with wrist extensor muscles at frequencies between 30 and 80 Hz. The illusory movement sensations were quantified through contralateral hand-tracking movements. Electromyographic activity from the extensor carpi radialis muscles was recorded with surface and intramuscular microelectrodes. The results showed that vibration of the wrist flexor muscle group induced both a kinesthetic illusion of wrist extension and a motor response in the extensor carpi radialis muscles. Combined vibration of the two antagonistic muscle groups at the same frequency evoked neither kinesthetic illusion nor motor activity. In addition, vibrating the same two antagonistic muscle groups at different frequencies induced both a kinesthetic illusion and a motor response in the muscle vibrated at the lowest frequency. The surface EMG amplitude of the extensor carpi radialis as well as the motor unit activation latency and discharge frequency were clearly correlated to the parameters of the illusory movement evoked by the vibration. Indeed, the faster the illusory sensation of movement, the greater the surface EMG in these muscles during the AVRs and the sooner and the more intense the activation of the motor units of the wrist extensor muscles. Moreover, comparison of the AVR with voluntary contraction showed that all parameters were highly similar. Mainly slow motor units were recruited during the AVR and during its voluntary reproduction. That the AVR is observed only when a kinesthetic illusion is evoked, together with the similarities between voluntary contractions and AVRs, suggests that this vibration-induced motor response may result from a perceptual-to-motor transformation of proprioceptive information, rather than from spinal reflex mechanisms.

Adult↗

Vestibular influences on human postural control in combinations of pitch and roll planes reveal differences in spatiotemporal processing.

The present study examined the influence of bilateral peripheral vestibular loss (BVL) in humans on postural responses to multidirectional surface rotations in the pitch and roll planes. Specifically, we examined the effects of vestibular loss on the directional sensitivity, timing, and amplitude of early stretch, balance correcting, and stabilizing reactions in postural leg and trunk muscles as well as changes in ankle torque and trunk angular velocity following multidirectional rotational perturbations of the support surface. Fourteen normal healthy adults and five BVL patients stood on a dual axis rotating platform which rotated 7.5 degrees at 50 degrees/s through eight different directions of pitch and roll combinations separated by 45 degrees. Directions were randomized within a series of 44 perturbation trials which were presented first with eyes open, followed by a second series of trials with eyes closed. Vestibular loss did not influence the range of activation or direction of maximum sensitivity for balance correcting responses (120-220 ms). Response onsets at approximately 120 ms were normal in tibialis anterior (TA), soleus (SOL), paraspinals (PARAS), or quadriceps muscles. Only SOL muscle activity demonstrated a 38- to 45-ms delay for combinations of forward (toe-down) and roll perturbations in BVL patients. The amplitude of balance correcting responses in leg muscles between 120 and 220 ms was, with one exception, severely reduced in BVL patients for eyes open and eyes closed conditions. SOL responses were decreased bilaterally for toe-up and toe-down perturbations, but more significantly reduced in the downhill (load-bearing) leg for combined roll and pitch perturbations. TA was significantly reduced bilaterally for toe-up perturbations, and in the downhill leg for backward roll perturbations. Forward perturbations, however, elicited significantly larger TA activity in BVL between 120 and 220 ms compared to normals, which would act to further destabilize the body. As a result of these changes in response amplitudes, BVL patients had reduced balance correcting ankle torque between 160 and 260 ms and increased torque between 280 and 380 ms compared to normals. There were no differences in the orientation of the resultant ankle torque vectors between BVL and normals, both of which were oriented primarily along the pitch plane. For combinations of backward (toe-up) and roll perturbations BVL patients had larger balance correcting and stabilizing reactions (between 350 and 700 ms) in PARAS than normals and these corresponded to excessive trunk pitch and roll velocities. During roll perturbations, trunk velocities in BVL subjects after 200 ms were directed along directions different from those of normals. Furthermore, roll instabilities appeared later than those of pitch particularly for backward roll perturbations. The results of the study show that combinations of roll and pitch surface rotations yield important spatiotemporal information, especially with respect to trunk response strategies changed by BVL which are not revealed by pitch plane perturbations alone. Our results indicate that vestibular influences are earlier for the pitch plane and are directed to leg muscles, whereas roll control is later and focused on trunk muscles.

Adult↗

Oculomotor function in the rhesus monkey after deafferentation of the extraocular muscles.

The function of extraocular muscle proprioception in the control of eye movements remains uncertain. In this study, we examined the effect of bilateral proprioceptive deafferentation of the extraocular muscles on eye movements in two rhesus monkeys. Before and after deafferentation, we analyzed baseline ocular alignment, saccades, pursuit, and vestibular eye movements. We also examined visually mediated adaptation of ocular alignment, saccades, and pursuit. Deafferentation of the eye muscles did not affect baseline ocular motor control, either acutely or over a 5-week period of study. Furthermore, visually mediated adaptation of the eye movement subtypes was also unaffected by deafferentation. These results suggest that ocular proprioception in primates is not used in the immediate, on-line control of eye movements and does not interact with visual cues in the adaptive modification of ocular motor function. We conclude that the efferent command (efference copy) provides sufficient information about eye kinematics to the brain for accurate eye movement control in normal monkeys, and that this information is modified by visual feedback independently of proprioception. We hypothesize that proprioception may be used to calibrate the efference copy during development and in response to perturbations by signaling potential mismatches between eye movement information derived from the efferent command and the actual motion of the eye transduced by the proprioceptive organs.

Adaptation, Physiological↗

Identification of the head-neck complex in response to trunk horizontal vibration.

A method is proposed for identifying the head-neck complex (HNC) in the seated human body when it is exposed to the trunk horizontal (fore-and-aft) vibration. It is assumed that the HNC only has the anteroposterior (flexion/extension) motion in the sagittal plane. An electrohydraulic vibrator is used as a source of vibration. To generate the trunk horizontal vibration, the trunk of the seated subject is fixed to the seatback. The subjects are exposed to the random vibration at a magnitude of 1.60 ms(-2) rms (root-mean-square) for 50 s. The coherence and frequency response function are then obtained in the frequency range 0.5-3 Hz. The results show that the HNC behavior is quasilinear with a resonance frequency between 1 and 1.4 Hz. Accordingly, a two-dimensional single-inverted pendulum is considered as a model for the HNC. The frequency domain identification method is then used to estimate the unknown parameters, including the HNC viscoelastic and inertia parameters. The model is examined in a time domain using the random vibration. Good agreement is obtained between experimental and simulation results, indicating the reliability of the proposed method.

Acceleration↗