Head Yaw Research Articles

The fine structure of honeybee head and body yaw movements in a homing task.

Honeybees turn their thorax and thus their flight motor to change direction or to fly sideways. If the bee's head were fixed to its thorax, such movements would have great impact on vision. Head movements independent of thorax orientation can stabilize gaze and thus play an important and active role in shaping the structure of the visual input the animal receives. Here, we investigate how gaze and flight control interact in a homing task. We use high-speed video equipment to record the head and body movements of honeybees approaching and departing from a food source that was located between three landmarks in an indoor flight arena. During these flights, the bees' trajectories consist of straight flight segments combined with rapid turns. These short and fast yaw turns ('saccades') are in most cases accompanied by even faster head yaw turns that start about 8 ms earlier than the body saccades. Between saccades, gaze stabilization leads to a behavioural elimination of rotational components from the optical flow pattern, which facilitates depth perception from motion parallax.

Enhanced visual fields in hammerhead sharks

Several factors that influence the evolution of the unusual head morphology of hammerhead sharks (family Sphyrnidae) are proposed but few are empirically tested. In this study we tested the 'enhanced binocular field' hypothesis (that proposes enhanced frontal binocularity) by comparison of the visual fields of three hammerhead species: the bonnethead shark, Sphyrna tiburo, the scalloped hammerhead shark, Sphyrna lewini, and the winghead shark, Eusphyra blochii, with that of two carcharhinid species: the lemon shark, Negaprion brevirostris, and the blacknose shark, Carcharhinus acronotus. Additionally, eye rotation and head yaw were quantified to determine if species compensate for large blind areas anterior to the head. The winghead shark possessed the largest anterior binocular overlap (48 deg.) and was nearly four times larger than that of the lemon (10 deg.) and blacknose (11 deg.) sharks. The binocular overlap in the scalloped hammerhead sharks (34 deg.) was greater than the bonnethead sharks (13 deg.) and carcharhinid species; however, the bonnethead shark did not differ from the carcharhinids. These results indicate that binocular overlap has increased with lateral head expansion in hammerhead sharks. The hammerhead species did not demonstrate greater eye rotation in the anterior or posterior direction. However, both the scalloped hammerhead and bonnethead sharks exhibited greater head yaw during swimming (16.9 deg. and 15.6 deg., respectively) than the lemon (15.1 deg.) and blacknose (15.0 deg.) sharks, indicating a behavioral compensation for the anterior blind area. This study illustrates the larger binocular overlap in hammerhead species relative to their carcharhinid sister taxa and is consistent with the 'enhanced binocular field' hypothesis.

Control and function of arm swing in human walking and running

We investigated the control and function of arm swing in human walking and running to test the hypothesis that the arms act as passive mass dampers powered by movement of the lower body, rather than being actively driven by the shoulder muscles. We measured locomotor cost, deltoid muscle activity and kinematics in 10 healthy adult subjects while walking and running on a treadmill in three experimental conditions: control; no arms (arms folded across the chest); and arm weights (weights worn at the elbow). Decreasing and increasing the moment of inertia of the upper body in no arms and arm weights conditions, respectively, had corresponding effects on head yaw and on the phase differences between shoulder and pelvis rotation, consistent with the view of arms as mass dampers. Angular acceleration of the shoulders and arm increased with torsion of the trunk and shoulder, respectively, but angular acceleration of the shoulders was not inversely related to angular acceleration of the pelvis or arm. Restricting arm swing in no arms trials had no effect on locomotor cost. Anterior and posterior portions of the deltoid contracted simultaneously rather than firing alternately to drive the arm. These results support a passive arm swing hypothesis for upper body movement during human walking and running, in which the trunk and shoulders act primarily as elastic linkages between the pelvis, shoulder girdle and arms, the arms act as passive mass dampers which reduce torso and head rotation, and upper body movement is primarily powered by lower body movement.

Head Yaw Estimation From Asymmetry of Facial Appearance

This paper proposes a novel method to estimate the head yaw rotations based on the asymmetry of 2-D facial appearance. In traditional appearance-based pose estimation methods, features are typically extracted holistically by subspace analysis such as principal component analysis, linear discriminant analysis (LDA), etc., which are not designed to directly model the pose variations. In this paper, we argue and reveal that the asymmetry in the intensities of each row of the face image is closely relevant to the yaw rotation of the head and, at the same time, evidently insensitive to the identity of the input face. Specifically, to extract the asymmetry information, 1-D Gabor filters and Fourier transform are exploited. LDA is further applied to the asymmetry features to enhance the discrimination ability. By using the simple nearest centroid classifier, experimental results on two multipose databases show that the proposed features outperform other features. In particular, the generalization of the proposed asymmetry features is verified by the impressive performance when the training and the testing data sets are heterogeneous.

Static and dynamic vestibulo-cervico-ocular responses after prolonged exposure to microgravity

The vestibular function was investigated in 13 Russian crew members of the ISS missions on days 1(2), 4(5), and 8(9) after prolonged exposure to microgravity (126 to 195 days). The static torsional otolith-cervico-ocular reflex was studied, as well as the dynamic vestibulo-cervico-ocular responses, vestibular reactivity, and spontaneous oculomotor activity using videooculography (VOG) and electrooculography (EOG) for simultaneous recording of eye movements. On days 1-2 of return to the gravity (R+1-2), the cosmonauts were found to increase the spontaneous oculomotor activity (floating eye movements, both typical and atypical forms of spontaneous nystagmus, square wave jerks, gaze nystagmus) with the head held in the vertical position. The otolith function during static head inclinations to the right or left shoulder at 30 degrees was suppressed as determined by the inversion or absence, or reduction by half of the amplitude of torsional compensatory eye counter-rolling and the vestibular reactivity during head yaw movements at 0.125 Hz was increased as revealed by a lowered threshold and an increased intensity of vestibular nystagmus. The pattern, depth, dynamics, and velocity of the vestibular function recovery varied with individual participants in the investigation. However, the suppressed otolith functioning in the period of readaptation to the normal gravity was, as a rule, accompanied by an exaggerated vestibular reactivity.

Vestibular adaptation to centrifugation does not transfer across planes of head rotation

Out-of-plane head movements performed during fast rotation produce non-compensatory nystagmus, sensations of illusory motion, and often motion sickness. Adaptation to this cross-coupled Coriolis stimulus has previously been demonstrated for head turns made in the yaw (transverse) plane of motion, during supine head-on-axis rotation. An open question, however, is if adaptation to head movements in one plane of motion transfers to head movements performed in a new, unpracticed plane of motion. Evidence of transfer would imply the brain builds up a generalized model of the vestibular sensory-motor system, instead of learning a variety of individual input/output relations separately. To investigate, over two days 9 subjects performed pitch head turns (sagittal plane) while rotating, before and after a series of yaw head turns while rotating. A Control Group of 10 subjects performed only the pitch movements. The vestibulo-ocular reflex (VOR) and sensations of illusory motion were recorded in the dark for all movements. Upon comparing the two groups we failed to find any evidence of transfer from the yaw plane to the pitch plane, suggesting that adaptation to cross-coupled stimuli is specific to the particular plane of head movement. The findings have applications for the use of centrifugation as a possible countermeasure for long duration spaceflight. Adapting astronauts to unconstrained head movements while rotating will likely require exposure to head movements in all planes and directions.

Incremental adaptation to yaw head turns during 30 RPM centrifugation

A 3-day incremental protocol was conducted with the aim of adapting human subjects to make head movements comfortably during 30 RPM centrifugation. With motion sickness as a potentially limiting factor, the protocol was designed using a quantitative motion sickness model based upon the neural mismatch sensory conflict theory. Centrifuge velocity was incremented from 14 RPM on day 1, to 23 RPM on day 2, to 30 RPM on day 3, with subjects making a total of 42 head movements on each day. Twenty-four subjects completed the experiment with average motion sickness levels below five (out of 20). Four subjects aborted due to motion sickness. Adaptation of non-compensatory vertical nystagmus was observed through an 18% decrease in the vertical aVOR time constant over the 3 days. Subjective intensity ratings for the head movements decreased by approximately 40% over the 3 days, while illusory motion duration decreased by 18%. Feasibility of head movements during 30 RPM rotation was demonstrated with only 3 days of incremental training.

Threshold-based vestibular adaptation to cross-coupled canal stimulation

Prior experiments have demonstrated that people are able to adapt to cross-coupled accelerations associated with head movements while spinning at high rotation rates (e.g., 23 rpm or 138 degrees/s). However, while adapting, subjects commonly experience serious side effects, such as motion sickness, non-compensatory eye movements, and strong and potentially disorienting illusory body tilt or tumbling sensations. In the present study, we investigated the feasibility of adaptation using a threshold-based method, which ensured that the illusory tilt sensations remained imperceptible or just barely noticeable. This was achieved by incrementally increasing the angular velocity of the horizontal centrifuge while supine subjects made repeated consistent yaw head turns. Incremental adaptation phases started at centrifugation speeds of 3 rpm. Centrifuge speed was slowly increased in steps of 1.5 rpm until a light illusory tilt was experienced. At the end of the incremental procedure, subjects were able to make head turns while rotating 14 rpm without experiencing illusory tilt. Moreover, motion sickness symptoms could be avoided and a limited carry over of the adaptive state to stronger stimulation at 23 rpm was found. The results are compared to prior studies which adapted subjects to super-threshold stimuli.

Rotation Axes of the Head During Positioning, Head Shaking, and Locomotion

Static head orientations obey Donders' law and are postulated to be rotations constrained by a Fick gimbal. Head oscillations can be voluntary or generated during natural locomotion. Whether the rotation axes of the voluntary oscillations or during locomotion are constrained by the same gimbal is unknown and is the subject of this study. Head orientation was monitored with an Optotrak (Northern Digital). Human subjects viewed visual targets wearing pin-hole goggles to achieve static head positions with the eyes centered in the orbit. Incremental rotation axes were determined for pitch and yaw by computing the velocity vectors during head oscillation and during locomotion at 1.5 m/s on a treadmill. Static head orientation could be described by a generalization of the Fick gimbal by having the axis of the second rotation rotate by a fraction, k, of the angle of the first rotation without a third rotation. We have designated this as a k-gimbal system. Incremental rotation axes for both pitch and yaw oscillations were functions of the pitch but not the yaw head positions. The pivot point for head oscillations was close to the midpoint of the interaural line. During locomotion, however, the pivot point was considerably lower. These findings are well explained by an implementation of the k-gimbal model, which has a rotation axis superimposed on a Fick-gimbal system. This could be realized physiologically by the head interface with the dens and occipital condyles during head oscillation with a contribution of the lower spine to pitch during locomotion.

Modeling sensory conflict and motion sickness in artificial gravity

It is necessary to characterize the vestibular response associated with head movements for various centrifuge rotation rates if one is to explore short-radius centrifugation as a viable form of artificial gravity for future spaceflights. An existing motion sickness model was modified to design an adaptation protocol to facilitate head movements at a centrifuge speed of 30 rpm. Modification involved addition of a quantitative sensory conflict model to serve as the input to the motion sickness model. Sensory conflict in this context was based on the dynamics of head movements during centrifugation as well as a previously developed transfer function relating angular accelerations to semicircular canal firing rates. Additionally, an adaptation parameter based on comparison between model predictions and previous experimental results was added. A 3-day incremental adaptation protocol was conducted in which 16 subjects successfully made 30 yaw head movements during rotation at 30 rpm on day 3. Motion sickness results showed good agreement with model predictions and demonstrated the feasibility of adaptation to increasingly high rotation rates.

Eye movements are commutative

Purpose: To provide evidence that all eye movements are commutative and to use that evidence to establish a coherent theory of ocular kinematics. Methods: Fixation targets and two miniature video cameras, one to record eye movements and the other to record head movements, were suspended from a head band. The apparatus (VOG) keeps the eyes stable in the orbits during head yaw, pitch, and tilt while allowing the head to move freely. It was also used for tracking eye movements with the head held stationary. Ocular orientation was determined from downloaded video clips by superimposing serial frames. Results: Video frame analysis reveals that all eye movements are commutative. The eye has one fixed axis within its substance, which serves eye movements to and from the midsagittal orbital plane. Brow-up and chin-down eye movements occur around an axis fixed in the orbit. Oblique movements consist of alternating arcs of great and small circles. The vertical extraocular muscles act synergistically when the head is held stationary producing slope and reciprocally during head tilt producing transient torsional movements. Conclusions: The globe rotates around two axes when the eye is stationary and one axis when the head tilts. The orientation of the eye can be calculated using the formula for the derivative of an ellipse. The results suggest a coherent theory of ocular movement: The function of the extraocular muscles is to maintain the fixation plane of each eye in equilibrium with the brain and coplanar with the fixation plane of the other eye in all gaze directions and head positions.

Locomotor Skills and Balance Strategies in Adolescents Idiopathic Scoliosis

Locomotor balance control assessment was performed to study the effect of idiopathic scoliosis on head-trunk coordination in 17 patients with adolescent idiopathic scoliosis (AIS) and 16 control subjects. The aim of this study was to explore the functional effects of structural spinal deformations like idiopathic scoliosis on the balance strategies used during locomotion. Up to now, the repercussion of the idiopathic scoliosis on head-trunk coordination and balance strategies during locomotion is relatively unknown. Seventeen patients with AIS (mean age 14 years 3 months, 10 degrees < Cobb angle > 30 degrees) and 16 control subjects (mean age 14 years 1 month) were tested during various locomotor tasks: walking on the ground, walking on a line, and walking on a beam. Balance control was examined in terms of rotation about the vertical axis (yaw) and on a frontal plane (roll). Kinematics of foot, pelvis, trunk, shoulder, and head rotations were measured with an automatic optical TV image processor in order to calculate angular dispersions and segmental stabilizations. Decreasing the walking speed is the main adaptive strategy used in response to balance problems in control subjects as well as patients with AIS. However, patients with AIS performed walking tasks more slowly than normal subjects (around 15%). Moreover, the pelvic stabilization is preserved, despite the structural changes affecting the spine. Lastly, the biomechanical defect resulting from idiopathic scoliosis mainly affects the yaw head stabilization during locomotion. Patients with AIS show substantial similarities with control subjects in adaptive strategies relative to locomotor velocity as well as balance control based on segmental stabilization. In contrast, the loss of the yaw head stabilization strategies, mainly based on the use of vestibular information, probably reflects the presence of vestibular deficits in the patients with AIS.

Binaural transformation coding with simulated head tracking

The binaural transformation codec synthesizes generic binaural audio signals and generates accompanying side information. In the decoder the side information is used to personalize the generic signal. The performance of the basic codec is described in another paper; this paper describes the estimated effect on quality of incorporating limited head tracking. Generic HRTFs were used to spatialize two concurrent sound sources spatialized at ±30 or ±80 deg. The generic binaural signal was personalized in the BX decoder and at the same time the individual sound sources were rotated. These induced rotations are equivalent to compensating for head yaw rotations of −80, −40, −20, −10, and −5 deg, and for head pitch rotations of 22.5, 45, and 90 deg. Listening tests based on Recommendation ITU‐R BS.1116‐1 were used to evaluate the processed sounds. The tests were conducted using speech, vocals, guitars, and percussion source materials. It was found that the quality of the processed sound tended to degrade as the head yaw rotation increased. However, except for the head yaw condition −80 deg, the quality of the processed sound remained greater than 4.0 on the mean opinion scale. No deterioration of the processed sound was observed for the applied pitch rotations.

Proprioceptive encoding of head position in the black soldier fly,Hermetia illucens(L.) (Stratiomyidae)

Because the eyes of insects cannot be moved independently of the head, information about head posture is essential for stabilizing the visual world or providing information about the direction of gaze. We examined the external anatomy and physiological capabilities of a head posture proprioceptor, the prosternal organ (PO), located at the base of the neck in the black soldier fly, Hermetia illucens (L.) (Family: Stratiomyidae). The PO is sexually isomorphic and is composed of two fused plates of about 130 mechanosensory hairs set in asymmetrical sockets whose orientation varies across the organ. A multi-joint mechanical coupling between the head, neck membrane, and contact sclerites deflects the hairs more or less to increase or decrease their level of excitation. The PO sensory afferents project to the central nervous system (CNS) via a pair of bilateral prosternal nerves (PN) to the fused thoracic ganglia. Simultaneous recording of spiking activity in the PN and videotaping of wind-induced and voluntary head movements around all three axes of head rotation reveal that a few PN afferents are active at rest, but activity increases tonically in response to head deflections. Activity is significantly modulated by change in head angles around the pitch (+/-40 degrees ), yaw (+/-30 degrees ) and roll (more than +/-90 degrees ) axes, although the dynamic range of spiking activity differs for each axis of rotation. Prosternal nerve afferents are bilaterally excited (inhibited) by pitch down (up); excited (inhibited) by head yaw toward the ipsilateral (contralateral) side; excited by roll down toward the ipsilateral side, but little inhibited by roll toward the opposite side. Although bilateral comparison of activity in PN afferents reliably encodes head posture around a given rotational axis, from the point of view of the CNS, the problem of encoding head posture is ill-posed with three axes of rotation and only two streams of afferent information. Furthermore, when the head is rotated around more than one axis simultaneously, mechanical interactions in the neck modify the responses to postural changes around the three rotational axes, which adds further ambiguity to reliable encoding of head posture. The properties of the PO in this relatively basal fly species are compared to those of higher flies and possible mechanisms of disambiguation are discussed.

Three-Dimensional Computation during Off-Vertical Axis Rotation (OVAR) in Monkeys

In order to study how dorso-ventral and naso-occipital linear accelerations affect the ocular responses induced by inter-aural axis linear accelerations, three-dimensional cyclic modulation of eye positions and velocities during off-vertical axis rotation (OVAR) were analyzed in four rhesus monkeys. The animals were rotated in darkness at 60°/s about an earth horizontal axis while placed in a pitched position relative to the rotation axis, so that the plane in which the gravitational vector rotated around the head was varied while the acceleration applied along the inter-aural axis remained identical across different pitch conditions. The amplitude of the compensatory bias slow phase eye velocity was the largest when the gravitational vector rotated in a plane between the animals' stereotaxic horizontal and presumed utricular planes. The phase of the modulation in yaw (horizontal) slow phase velocity, but not in eye position, tended to lag as the stimulus axis deviated from the normal to the presumed utricular plane. The modulation of roll (torsional) eye position was a function of the interaural acceleration, but the amplitude of modulation of roll slow phase velocity decreased with an increase in the roll bias velocity. Modulation of pitch (vertical) eye position was the largest when the axis of rotation corresponded to the head yaw axis, but modulation of pitch slow phase velocity was the largest during rotation about an axis close to the roll axis. Thus, the time differentiation of the positional modulation did not predict the velocity modulation. It is hypothesized that there are separate pathways for processing cyclic modulations of the eye position and velocity, and that the velocity storage utilizes both utricular and saccular information.

Cerebellar Disease Alters the Axis of the High-Acceleration Vestibuloocular Reflex

L. W. Schultheis and D. A. Robinson showed that the axis of the rotational vestibuloocular reflex (RVOR) cannot be altered by visual-vestibular mismatch ("cross-axis adaptation") when the vestibulocerebellum is lesioned. This suggests that the cerebellum may calibrate the axis of eye velocity of the RVOR under natural conditions. Thus we asked whether patients with cerebellar disease have alterations in the RVOR axis and, if so, what might be the mechanism. We used three-axis scleral coils to record head and eye movements during yaw, pitch, and roll head impulses in 18 patients with cerebellar disease and in a comparison group of eight subjects without neurologic disease. We found distinct shifts of the eye-velocity axis in patients. The characteristic finding was a disconjugate upward eye velocity during yaw. Measured at 70 ms after the onset of head rotation, the median upward gaze velocity was 15% of yaw head velocity for patients and <1% for normal subjects (P < 0.001). Upward eye velocity was greater in the contralateral (abducting) eye during yaw and in the ipsilateral eye during roll. Patients had a higher gain (eye speed/head speed) for downward than for upward pitch (median ratio of downward to upward gain: 1.3). In patients, upward gaze velocities during both yaw and roll correlated with the difference in anterior (AC) and posterior canal excitations, scaled by the respective pitch gains. Our findings support the hypothesis that upward eye velocity during yaw results from AC excitation, which must normally be suppressed by the intact cerebellum.

Adaptation of VOR to Coriolis Stimulation

The vestibulo-ocular reflex (VOR) is normally characterized by the gain and phase of slow-phase velocity (SPV) relative to the stimulus velocity. Although this is perfectly satisfactory for steady-state sinusoidal oscillations about a single axis, it is less useful when applied to transient responses. The well-known decay of nystagmus following a step change of head velocity approximately follows a double exponential, with an initial amplitude (A), a long time constant (tau), and an adaptation time constant (tau(a)). We have developed a means of representing the transient response for a complex head velocity stimulus as experienced during high-speed artificial gravity (AG) experiments. When a subject, lying supine on a rotating horizontal platform, makes a yaw head movement of amplitude theta, the vertical semicircular canals experience a step in angular velocity. The pitch stimulus is equal to the change in the component of the centrifuge angular velocity (omega(c)) aligned with the interaural axis, and gives rise to a vertical VOR. The magnitude of the step change is omega(c) sin theta. The SPV is approximated by an exponential decay of amplitude A and single time constant tau, and then normalized relative to this stimulus step. MATLAB scripts filter the raw eye position data to remove noise, blinks, and saccades, differentiate the signal, and remove fast phases to obtain SPV. The amplitude of the fitted SPV exponential is divided by omega(c) sin theta to obtain the normalized SPV. A and tau are shown to behave differently as subjects adapt to repeated head movements of different amplitudes.

Effects of Chronic Infusion of a GABAA Receptor Agonist or Antagonist into the Vestibular Nuclear Complex on Vestibular Compensation in the Guinea Pig

The aim of this study was to determine the effects of chronic infusion of a GABA(A) receptor agonist/antagonist into the ipsilateral or contralateral vestibular nuclear complex (VNC) on vestibular compensation, the process of behavioral recovery that occurs after unilateral vestibular deafferentation (UVD). This was achieved by a mini-osmotic pump that infused, over 30 h, muscimol or gabazine into the ipsilateral or contralateral VNC. Spontaneous nystagmus (SN), yaw head tilt (YHT), and roll head tilt (RHT) were measured. Infusion of muscimol or gabazine into either the ipsilateral or the contralateral VNC had little effect on SN compensation. In contrast, infusion of muscimol (250, 500, and 750 ng) into the contralateral VNC and gabazine (31.25, 62.5, and 125 ng) into the ipsilateral VNC significantly affected YHT and RHT (p < 0.05), but not their rate of compensation (p > 0.05). Interestingly, the effects of muscimol and gabazine on YHT and RHT were consistent throughout the first 30 h post-UVD. Infusion of muscimol (62.5, 125, and 250 ng) into the ipsilateral VNC and gabazine (125, 375, and 750 ng) into the contralateral VNC had little effect on YHT and RHT or their rate of compensation. These results suggest that the ipsilateral gabazine and contralateral muscimol infusions are modifying the expression of the symptoms without altering the mechanism of compensation. Furthermore, the neurochemical mechanism responsible for vestibular compensation can cope with the both the GABA(A) receptor-mediated and the UVD-induced decrease in resting activity.

Stabilization and mobility of the head and trunk in vervet monkeys(Cercopithecus aethiops) during treadmill walks and gallops

The brain requires internal or external reference frames to determine body orientation in space. These frames may change, however, to meet changing conditions. During quadrupedal overground locomotion by monkeys, the head rotates on a stabilized trunk during walking, but the trunk rotates on a stabilized head during galloping. Do the same movement patterns occur during in-place locomotion? Head and trunk pitch rotations were measured, and yaw and roll rotations estimated from cine films of three adult vervet monkeys (Cercopithecus aethiops L. 1758) walking and galloping quadrupedally on a treadmill. Head and trunk rotational patterns during treadmill walks were comparable to the patterns found during overground walks. The rotational velocities of these segments during both treadmill walks and gallops were also comparable to the velocities found during natural locomotion. By contrast, whereas head and trunk rotational patterns during treadmill gallops did occur that were comparable to the patterns practiced during overground gallops, a significantly different pattern involving large and simultaneous head and trunk rotations was more commonly observed. Simultaneous head and trunk rotations may be possible during treadmill gallops because the fixed visual surround is providing an adequate spatial reference frame. Alternatively, or in addition to this visual information, a re-weighting in other sensory modalities may be occurring. Specifically, the vestibular inputs used during overground locomotion to reference gravity or a gravity-derived vector may become less important than proprioceptive inputs that are using the treadmill belt surface as a reference. Regardless, the spatial reference frame being used, blinks that occur at specific times during the largest head yaw rotations may be necessary to avoid the initiation of unwanted and potentially destabilizing lateral sway brought on by sudden increases in optic flow velocity.

Neck proprioception and spatial orientation in cervical dystonia.

Neck muscle vibration is known to influence body orientation and posture during locomotion and stance in normal subjects. To verify the hypothesis that neck proprioceptive input can be misinterpreted in patients with cervical dystonia (CD), lateral continuous vibration was applied to the sternocleidomastoid muscle during both stepping-in-place and quiet stance, with eyes closed. The orienting responses of CD patients were compared with those of normal subjects. Vibration effects on body orientation during stepping and stance were apparently different from normal, since no effects were seen when all patients' data collapsed were analysed. However, while some patients did not respond to vibratory stimuli regardless of the vibrated side, others had a 'good' side, the stimulation of which produced effects on body orientation similar to those observed in normal subjects. Homogeneous groups within the patient population were identified, based on the vibration-induced responses under stepping conditions. The different orienting or postural responses observed in CD patients were correlated with disease-related features such as spontaneous head position, maximum range of voluntary head yaw, presence or absence of a botulinum toxin treatment and disease duration. Our data suggest that, in CD patients, the reference system used in the control of body orientation in space is either refractory to the lateralized proprioceptive neck input or modified such that the input from both sides produces an orientation shift in the same sense. This would depend on the pathogenesis of the disease or on an adaptive process connected to the head abnormal posture. It seems that this refractoriness spreads to both sides of the neck with the advancement of the disease, thereby possibly entraining a progressive shift from a reference system based on the head to a more reliable egocentric reference.