Corollary discharge enables proprioception from lateral line sensory feedback

Animals modulate sensory processing in concert with motor actions. Parallel copies of motor signals, called corollary discharge (CD), prepare the nervous system to process the mixture of externally and self-generated (reafferent) feedback that arises during locomotion. Commonly, CD in the peripheral nervous system cancels reafference to protect sensors and the central nervous system from being fatigued and overwhelmed by self-generated feedback. However, cancellation also limits the feedback that contributes to an animal’s awareness of its body position and motion within the environment, the sense of proprioception. We propose that, rather than cancellation, CD to the fish lateral line organ restructures reafference to maximize proprioceptive information content. Fishes’ undulatory body motions induce reafferent feedback that can encode the body’s instantaneous configuration with respect to fluid flows. We combined experimental and computational analyses of swimming biomechanics and hair cell physiology to develop a neuromechanical model of how fish can track peak body curvature, a key signature of axial undulatory locomotion. Without CD, this computation would be challenged by sensory adaptation, typified by decaying sensitivity and phase distortions with respect to an input stimulus. We find that CD interacts synergistically with sensor polarization to sharpen sensitivity along sensors’ preferred axes. The sharpening of sensitivity regulates spiking to a narrow interval coinciding with peak reafferent stimulation, which prevents adaptation and homogenizes the otherwise variable sensor output. Our integrative model reveals a vital role of CD for ensuring precise proprioceptive feedback during undulatory locomotion, which we term external proprioception.

Animals modulate sensory processing in concert with motor actions. A study of the corollary discharge in zebrafish reveals that it modulates the sensitivity of the lateral line during swimming to prevent sensor adaptation and maintain the high-quality feedback necessary for kinematic control.     

Effect of sitting postures and shoulder position on the cervicocephalic kinesthesia in healthy young males

Information about head orientation, position, and movement with respect to the trunk relies on the visual, vestibular, extensive muscular, and articular proprioceptive system of the neck. Various factors can affect proprioception since it is the function of afferent integration, and tuning of muscular and articular receptors. Pain, muscle fatigue, and joint position have been shown to affect proprioceptive capacity. Thus, it can be speculated that changes in body posture can alter the neck proprioception. This study was undertaken to investigate the effect of body posture on cervicocephalic kinesthetic sense in healthy subjects. Cervicocephalic kinesthetic sensibility was measured by the kinesthetic sensibility test in healthy young adults while in (a) habitual slouched sitting position with arms hanging by the side (SS), (b) habitual slouched sitting position with arms unloaded (supported) (SS-AS), and (c) upright sitting position with arms hanging by the side (US) during maximum and 30 degree right, left rotations, flexion, and extension. Thirty healthy male adults (mean age 27.83; SD 3.41) volunteered for this study. The least mean error was found for the SS-AS position (0.48; SD 0.24), followed by SS (0.60; SD 0.43) and US (0.96; SD 0.71), respectively. For all test conditions, there was significant difference in mean absolute error while head repositioning from maximum and 30 degree rotation during SS and SS-AS positions (p?相似文献   

Load dependence of simulated central tremor

Previous studies have argued that tremors of central versus peripheral origin can be distinguished based on their load dependence: the frequency of peripheral tremor decreases when a weight is added to the tremulous limb, while the frequency of central tremors remains unchanged. The present study scrutinizes the latter statement. We simulated central tremor using a simple network of coupled neural oscillators, which receives proprioceptive feedback from the motor periphery. The network produced a self-sustained, stable oscillation. When the gain of proprioceptive feedback was high, oscillation frequency decreased in the presence of an inertial load. When the gain was low, the oscillation frequency was load independent. We conclude that load dependence is not an exclusive property of peripheral tremors but may be found in tremors of central origin as well. Therefore, the load test is not sufficient to reject a central tremor origin. Received: 1 October 1998 / Accepted in revised form: 28 January 1999     

A biophysical approach to the Spatial Function of eye movements extraocular proprioception and the vestibulo-ocular reflex

The eyeball and the extraocular muscles are used as a paradigm to design a linear spatial model of a single joint with a redundant set of muscles. On the basis of this model relations are derived between orientation, torque, motor commands, and proprioceptive signals. These relations show that the tenet underlying the tensorial interpretation of neural signals in sensorimotor systems does not have general validity. A mechanism is proposed to show how proprioception may play a role in optimizing the coordination of muscles during spatial tasks. Further, a new concept is suggested that allows one to predict the neural connectivities mediating the redundant spatial vestibulo-ocular reflex. This concept has the advantage of minimizing both sensorial error and motor effort.     

Correlation analysis of proprioceptive acuity in ipsilateral position-matching and velocity-discrimination

In order to plan and control movements the central nervous system (CNS) needs to continuously keep track of the state of the musculoskeletal system. Therefore the CNS constantly uses sensory input from mechanoreceptors in muscles, joints and skin to update information about body configuration on different levels of the CNS. On the conscious level, such representations constitute proprioception. Different tests for assessment of proprioceptive acuity have been described. However, it is unclear if the proprioceptive acuity measurements in these tests correlate within subjects. By using both uni- and multivariate analysis we compared proprioceptive acuity in different variants of ipsilateral active and passive limb position-matching and ipsilateral passive limb movement velocity-discrimination in a group of healthy subjects. The analysis of the position-matching data revealed a higher acuity of matching for active movements in comparison to passive ones. The acuity of matching was negatively correlated to movement extent. There was a lack of correlation between proprioceptive acuity measurements in position-matching and velocity-discrimination.     

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.     

Correlation analysis of proprioceptive acuity in ipsilateral position-matching and velocity-discrimination

In order to plan and control movements the central nervous system (CNS) needs to continuously keep track of the state of the musculoskeletal system. Therefore the CNS constantly uses sensory input from mechanoreceptors in muscles, joints and skin to update information about body configuration on different levels of the CNS. On the conscious level, such representations constitute proprioception. Different tests for assessment of proprioceptive acuity have been described. However, it is unclear if the proprioceptive acuity measurements in these tests correlate within subjects. By using both uni- and multivariate analysis we compared proprioceptive acuity in different variants of ipsilateral active and passive limb position-matching and ipsilateral passive limb movement velocity-discrimination in a group of healthy subjects. The analysis of the position-matching data revealed a higher acuity of matching for active movements in comparison to passive ones. The acuity of matching was negatively correlated to movement extent. There was a lack of correlation between proprioceptive acuity measurements in position-matching and velocity-discrimination.     

Development of leg chordotonal sensory organs in normal and heat shocked embryos of the cricket Teleogryllus commodus (Walker)

This paper describes the embryonic development of some parts of the sensory peripheral nervous system in the leg anlagen of the cricket Teleogryllus commodus in normal and heat shocked embryos. The first peripheral neurons appear at the 30% stage of embryogenesis. These tibial pioneer neurons grow on a stereotyped path to the central nervous system and form a nerve which is joined by the growth cones of axons that arise later, including those from the femoral chordotonal organ, subgenual organ and tympanal organ. The development of these organs is described with respect to the increase in number of sensory receptor cells and the shape and position of the organs. At the 100% stage of embryogenesis all three organs have completed their development in terms of the number of sense cells and have achieved an adult shape. To study the function of the tibial pioneer neurons during embryogenesis a heat shock was used to prevent their development. Absence of these neurons has no effect on the development of other neurons and organs proximal to them. However, the development of distal neurons and organs guided by them is impaired. The tibial pioneer neurons grow across the segmental boundary between femur and tibia early in development, and the path they form seems to be essential for establishing the correct connections of the distal sense organs with the central nervous system.     

Adaptive Staircase Measurement of Hand Proprioception

Clinicians and researchers often need to measure proprioception (position sense), for example to monitor the progress of disease, to identify the cause of movement or balance problems, or to ascertain the effects of an intervention. While researchers can use sophisticated equipment to estimate proprioceptive acuity with good precision, clinicians lack this option and must rely on the subjective and imprecise methods currently available in the clinic. Here we describe a novel technique that applies psychometric adaptive staircase procedures to hand proprioception with a simple tablet-style apparatus that could easily be adapted for the clinic. We report test-retest reliability, inter-rater reliability, and construct validity of the adaptive staircase method vs. two other methods that are commonly used in clinical settings: passive motion direction discrimination (PMDD) and matching. As a first step, we focus on healthy adults. Subjects ages 18–82 had their proprioception measured with each of the three techniques, at the metacarpophalangeal joint in the second finger of the right hand. A subset completed a second session in which the measures were repeated, to assess test-retest reliability. Another subset had the measurements done by two different testers to assess inter-rater reliability. Construct validity was assessed using stepwise regression on age and activity level, and correlations calculated across the three methods. Results suggest that of the three methods, the adaptive staircase method yields the best test-retest reliability, inter-rater reliability, and construct validity. The adaptive staircase method may prove to be a valuable clinical tool where more accurate assessment of proprioception is needed.     

An elasticity-curvature illusion decouples cutaneous and proprioceptive cues in active exploration of soft objects

Our sense of touch helps us encounter the richness of our natural world. Across a myriad of contexts and repetitions, we have learned to deploy certain exploratory movements in order to elicit perceptual cues that are salient and efficient. The task of identifying optimal exploration strategies and somatosensory cues that underlie our softness perception remains relevant and incomplete. Leveraging psychophysical evaluations combined with computational finite element modeling of skin contact mechanics, we investigate an illusion phenomenon in exploring softness; where small-compliant and large-stiff spheres are indiscriminable. By modulating contact interactions at the finger pad, we find this elasticity-curvature illusion is observable in passive touch, when the finger is constrained to be stationary and only cutaneous responses from mechanosensitive afferents are perceptible. However, these spheres become readily discriminable when explored volitionally with musculoskeletal proprioception available. We subsequently exploit this phenomenon to dissociate relative contributions from cutaneous and proprioceptive signals in encoding our percept of material softness. Our findings shed light on how we volitionally explore soft objects, i.e., by controlling surface contact force to optimally elicit and integrate proprioceptive inputs amidst indiscriminable cutaneous contact cues. Moreover, in passive touch, e.g., for touch-enabled displays grounded to the finger, we find those spheres are discriminable when rates of change in cutaneous contact are varied between the stimuli, to supplant proprioceptive feedback.     

Control of functional electrical stimulation with extended physiological proprioception

The use of functional electrical stimulation (FES) of muscle for paraplegic locomotion, or grasp augmentation in tetraplegia, is limited by the variability in muscle response to stimulation as a result of several external and internal factors. Previous approaches to this problem have used position-servo controllers, which have been shown to function satisfactorily in the laboratory. However, such systems will fail should obstacles be encountered or should the stimulation hardware develop a fault. To prevent such potentially dangerous failures some form of sensory feedback is required. This paper describes the first application of a technique known as extended physiological proprioception (EPP) to the control of FES to compensate for muscle response variability and provide proprioceptive feedback via the appropriate sensory pathways. In the experimental system described, a paraplegic subject controlled the extension of his paralysed knee by shoulder protraction. A Bowden cable linked the two joints, and a dynamometer in this cable was used to derive the control signal for a computer-controlled stimulator which delivered surface stimulation to the quadriceps muscle group. Modelling and parameter identification were performed by analysis of the step response, and the controller was designed from consideration of the root locus. The advantages of the system, in terms of improved proprioceptive feedback and reduced limb-positioning error were assessed in a test of joint positioning accuracy with vision occluded. The EPP system showed improvements over both open and closed-loop position-servo controllers.     

Physiology of vibration-sensitive afferents in the femoral chordotonal organ of the stick insect

The femoral chordotonal organ in orthopterans signals proprioceptive sensory information concerning the femur-tibia joint to the central nervous system. In the stick insect, 80 out of 500 afferents sense tibial position, velocity, or acceleration. It has been assumed that the other sensory cells in the chordotonal organ would serve as vibration detectors. Extracellular recordings from the femoral chordotonal organ nerve in fact revealed a sensitivity of the sense organ for vibrations with frequencies ranging from 10 Hz to 4 kHz, with a maximum sensitivity between 200 and 800 Hz. Single vibration-sensitive afferents responded to the same range of frequencies. Their spike activity depended on acceleration amplitude and displacement amplitude of the vibration stimulus. Additionally, 80% of the vibration-sensitive afferents received indirect presynaptic inputs from themselves or from other afferents of the femoral chordotonal organ, the amplitude of which depended on stimulus frequency and displacement amplitude. They were associated with a decrease of input resistance in the afferent terminal. From the present investigation we conclude that the femoral chordotonal organ of the stick insect is a bifunctional sensory organ that, on the one hand, measures position and movement of the tibia and, on the other hand, detects vibration of the tibia. Accepted: 6 November 1998     

Proprioception, gait kinematics, and rate of loading during walking: are they related?

The cyclic nature of walking can lead to repetitive stress and associated complications due to the rate of loading (ROL) experienced by the body at the initial contact of the foot with the ground. An individual's gait kinematics at initial contact has been suggested to give rise to the ROL, and a repetitive, high ROL may lead to several disorders, including osteoarthritis. Additionally, proprioception, the feedback signaling of limb position and movement, may play a role in how the foot strikes the ground and thus, the ROL. Our goal was to explore the relationship between proprioception, gait kinematics and ROL. Thirty-eight women were recruited for gait analysis, and the gait characteristics 50 ms prior to and at initial contact were examined. Two proprioception tests, joint angle reproduction and threshold to detect passive motion were used to examine the subject's proprioceptive acuity. Our results indicate that individuals with a larger knee angle (i.e., greater extension) 50 ms prior to initial contact (IC) experience a higher ROL during gait and have poorer proprioceptive scores. However, it remains unclear whether poor proprioception causes a high ROL or if a high ROL damages the mechanoreceptors involved in proprioception, but the apparent relationship is significant and warrants further investigation.     

Asymmetries and relationships between dynamic loading,muscle strength,and proprioceptive acuity at the knees in symptomatic unilateral hip osteoarthritis

Introduction

High joint loading, knee muscle weakness, and poor proprioceptive acuity are important factors that have been linked to knee osteoarthritis (OA). We previously reported that those with unilateral hip OA and bilateral asymptomatic knees are more predisposed to develop progressive OA in the contralateral knee relative to the ipsilateral knee. In the present study, we evaluate asymmetries in muscle strength and proprioception between the limbs and also evaluate relationships between these factors and joint loading that may be associated with the asymmetric evolution of OA in this group.

Methods

Sixty-two participants with symptomatic unilateral hip OA and asymptomatic knees were evaluated for muscle strength, joint position sense and dynamic joint loads at the knees. Muscle strength and proprioception were compared between limbs and correlations between these factors and dynamic joint loading were evaluated. Subgroup analyses were also performed in only those participants that fulfilled criteria for severe hip OA.

Results

Quadriceps muscle strength was 15% greater, and in the severe subgroup, proprioceptive acuity was 25% worse at the contralateral compared to ipsilateral knee of participants with unilateral hip OA (P <0.05). In addition, at the affected limb, there was an association between decreased proprioceptive acuity and higher knee loading (Spearman’s rho = 0.377, P = 0.007) and between decreased proprioceptive acuity and decreased muscle strength (Spearman’s rho = −0.328, P = 0.016).

Conclusions

This study demonstrated asymmetries in muscle strength and proprioception between the limbs in a unilateral hip OA population. Early alterations in these factors suggest their possible role in the future development of OA at the contralateral ‘OA-predisposed knee’ in this group. Furthermore, the significant association observed between proprioception, loading, and muscle strength at the affected hip limb suggests that these factors may be interrelated.     

Joint position reproduction and joint position discrimination at the ankle are not related

Abstract

Purpose: Limited data in current literature can be found on the relation between the two commonly-used active proprioception assessment methods –active joint position reproduction (JPR) and active movement extent discrimination assessment (AMEDA). The current study compared the two active methods, JPR and AMEDA, to investigate their interrelationship over two studies that differed in task difficulty, using active ankle inversion movements made in weight-bearing to maximise ecological validity.

Methods: 50 participants volunteered in this research, 20 of whom on a harder protocol and the other 30 on an easier protocol, were tested by both methods, JPR and AMEDA. Proprioceptive acuity was represented by absolute error (AE) and variable error (VE) for JPR and by AE and the area under the curve (AUC) for AMEDA.

Results: Proprioceptive acuity scores are found to be significantly correlated within test methods but not between methods for either hard or easy tasks, where JPR AE and VE scores were not correlated with either AMEDA AE or AUC. Further, proprioceptive acuity scores were significantly higher on the easy task when tested with the AMEDA method, but not with JPR method.

Conclusion: Scores obtained from the two active movement proprioception tests, movement extent discrimination and joint position reproduction, were not significantly correlated. Taken together with previous findings, these results show that for proprioception, scores from the three classical psychophysical methods for measuring sensitivity (adjustment, limits and constant stimuli) are not correlated with each other. This suggests that each proprioception measurement system assesses a different aspect of proprioception.     

Proprioceptive Interaction between the Two Arms in a Single-Arm Pointing Task

Proprioceptive signals coming from both arms are used to determine the perceived position of one arm in a two-arm matching task. Here, we examined whether the perceived position of one arm is affected by proprioceptive signals from the other arm in a one-arm pointing task in which participants specified the perceived position of an unseen reference arm with an indicator paddle. Both arms were hidden from the participant’s view throughout the study. In Experiment 1, with both arms placed in front of the body, the participants received 70–80 Hz vibration to the elbow flexors of the reference arm (= right arm) to induce the illusion of elbow extension. This extension illusion was compared with that when the left arm elbow flexors were vibrated or not. The degree of the vibration-induced extension illusion of the right arm was reduced in the presence of left arm vibration. In Experiment 2, we found that this kinesthetic interaction between the two arms did not occur when the left arm was vibrated in an abducted position. In Experiment 3, the vibration-induced extension illusion of one arm was fully developed when this arm was placed at an abducted position, indicating that the brain receives increased proprioceptive input from a vibrated arm even if the arm was abducted. Our results suggest that proprioceptive interaction between the two arms occurs in a one-arm pointing task when the two arms are aligned with one another. The position sense of one arm measured using a pointer appears to include the influences of incoming information from the other arm when both arms were placed in front of the body and parallel to one another.     

Mapping Proprioception across a 2D Horizontal Workspace

Relatively few studies have been reported that document how proprioception varies across the workspace of the human arm. Here we examined proprioceptive function across a horizontal planar workspace, using a new method that avoids active movement and interactions with other sensory modalities. We systematically mapped both proprioceptive acuity (sensitivity to hand position change) and bias (perceived location of the hand), across a horizontal-plane 2D workspace. Proprioception of both the left and right arms was tested at nine workspace locations and in 2 orthogonal directions (left-right and forwards-backwards). Subjects made repeated judgments about the position of their hand with respect to a remembered proprioceptive reference position, while grasping the handle of a robotic linkage that passively moved their hand to each judgement location. To rule out the possibility that the memory component of the proprioceptive testing procedure may have influenced our results, we repeated the procedure in a second experiment using a persistent visual reference position. Both methods resulted in qualitatively similar findings. Proprioception is not uniform across the workspace. Acuity was greater for limb configurations in which the hand was closer to the body, and was greater in a forward-backward direction than in a left-right direction. A robust difference in proprioceptive bias was observed across both experiments. At all workspace locations, the left hand was perceived to be to the left of its actual position, and the right hand was perceived to be to the right of its actual position. Finally, bias was smaller for hand positions closer to the body. The results of this study provide a systematic map of proprioceptive acuity and bias across the workspace of the limb that may be used to augment computational models of sensory-motor control, and to inform clinical assessment of sensory function in patients with sensory-motor deficits.     

Peripheral somatosensation: a touch of genetics

The somatosensory system processes information that organisms 'feel': joint position, muscle stretch, pain, pressure, temperature, and touch. The system is composed of a diverse array of peripheral nerve endings specialized to detect these sensory modalities. Several recent discoveries have shed light on the genetic pathways that control specification and differentiation of these neurons, how they accurately innervate their central and peripheral targets, and the molecules that enable them to detect mechanical stimuli. Here, we review the cadre of genes that control these processes, focusing on mechanosensitive neurons and support cells of the skin that mediate different aspects of the sense of touch.     

Relationship between postural stability and fall risk in elderly people with lumbar spondylosis during local vibratory stimulation for proprioception: a retrospective study

Abstract

Purpose: Reduced proprioception affects fall risks in elderly people with lumbar spondylosis. The decrease in proprioception in the trunk or lower legs may contribute to a decline in postural stability. We aimed to investigate the association between proprioceptive postural stability and fall risks in elderly individuals with lumbar spondylosis.

Materials and Methods: In this retrospective study, the centre-of-pressure displacement was determined in elderly individuals with lumbar spondylosis during upright stance while standing on a Wii Balance Board with their eyes closed (fall-risk group, n?=?55; non-fall-risk group, n?=?60). Vibratory stimulations at 30?Hz were applied to the lumbar multifidus and gastrocnemius to evaluate the relative contributions of proprioceptive signals used in postural control (relative proprioceptive weighting ratio).

Results: Compared with the non-fall-risk group, the fall-risk group displayed a high relative proprioceptive weighting ratio (p?=?0.024). Relative proprioceptive weighting ratio (odds ratio, 1.1; 95% confidence interval: 1.004–1.109) was independently associated with fall risks after adjusting for confounding factors. Among variables related to fall risk, the relative proprioceptive weighting ratio was a significant factor (p?<?0.035).

Conclusion: The fall-risk group of elderly individuals with lumbar spondylosis was dependent on the ankle strategy. The fall risk in elderly people with lumbar spondylosis could be due to over-dependence on the input from muscle spindles in the gastrocnemius.     

An investigation of the mechanisms of eye movement control

Summary Feedback mechanisms exist in all the periferal sense organs including the eye, which acts as a highly efficient position control servo system. Histological studies so far have not revealed the precise circuitry of the eye movement control system but some information about it can be obtained by a study of the sources of feedback. Existing theories have considered three types of feedback originating in the oculomotor tract, in the proprioceptive fibres of the extrinsic eye muscles and from retinal image displacement. In the present experiments an optical arrangement has been used to vary or eliminate the amount of information available from retinal image motion, and the response of the eye to simple harmonic displacement of a target has been recorded. The response curves of gain (eyeball movement divided by target motion) against frequency indicate that the system is lion linear when the image falls in the retinal region which is insensitive to position. Outside this area, retinal image position is used as negative feedback but the information from the oculomotor tract must be regenerative. There is also evidence for feedback proportional to the first derivative of eyeball position and this function is ascribed to the proprioceptive signals; this form of feedback appears to saturate for large amplitude movements, thus avoiding heavy damping of the flick movements.A schematic eye movement control system having the same characteristics as the eye is proposed. The transfer function of this system indicates that it should be unstable if the sign of the retinal image feedback loop is reversed. Experiments with this form of feedback show that steady fixation is impossible and the eye performs a pendular nystagmus.