Changes in the direction of vestibulomotor response in the course of adaptation to protracted static head turning in man

Vestibulomotor response during the course of adaptation to prolonged (10 min) static head turning to the furthest limit was investigated in healthy subjects standing upright with the eyes closed. The head was either actively or passively maintained in this position. The sensation of a decline in the angle of head turning was experienced during adaptation to the position by five of the 12 subjects tested. Error in appreciating this angle ranged up to 70–80°. Matching changes occurred in the direction of vestibulomotor response to electrical stimulation of the vestibular apparatus. When true and perceived head position conflict, direction of vestibulomotor response thus matches spatial perception rather than actual orientation of the head.Institute for Research into Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 21, No. 2, pp. 210–217, March–April, 1989.     

Effect of movement and illusion of movement on human vestibulomotor response

Lateral stabilographic response to galvanic labyrinth stimulation was investigated in healthy subjects in the standing position. Vestibulomotor response increased during forwards volitional body tilt as well as involuntary tilt occurring in response to stimulating (by vibration) the proprioceptors of the anterior tibial muscles. An illusion of the forward body tilt induced by stimulating (vibrating) the proprioceptors of the triceps surae muscles with the trunk fastened in a fixed position was accompanied by practically the same intensification of vestibulomotor response as during actual body movement. It was concluded that reinforcement of vestibulomotor response during volitional movements is brought about by the spatial perception system.Institute for Research into Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 20, No. 2, pp. 250–255, March–April, 1988.     

Postural responses to vibrostimulation of the neck muscle proprioceptors in man

Postural responses to vibrostimulation (50–100 Hz, 0.5 mm, 4–8 sec) of muscles of the back surface of the neck were studied in healthy subjects. In the sitting position, vibrostimulation evoked local displacements (backward head deflection), but global postural responses (forward inclination of the whole body) developed in the standing position. The amplitude of the evoked body inclination was dependent upon the site of the vibrostimuli application along the vertebral column. Asymmetrical application of vibrostimuli to the muscles of the right or left neck side was accompanied by development of a lateral component in the postural response. Changes in the spatial orientation of the head led to the changes in postural response direction: head turning to the right resulted in right-side body deviation during vibration, and vice versa. Illusions of head bend caused by habituation to its static turning were accompanied by precisely the same changes in the direction of body deviation. It is assumed that neck-evoked motor events are mediated via central mechanisms that are involved in perception of the head and body position in space.Translated from Neirofiziologiya, Vol. 25, No. 2, pp. 101–108, March–April, 1993.     

Changes in vestibular postural response determined by information content of visual feedback

Electrical unipolar monoaural stimulation of the labyrinth led to body sway mainly on a frontal plane in normal human subjects in a standing position. Early and late stages of response with latencies of 120–200 and 200–500 msec respectively changing in size in accordance with conditions of visual control were distinguished in the stabilographic response. Maximum response was recorded when the eyes were closed. Response declined upon opening the eyes, fixing the gaze on a static target, and with visual feedback according to stabilograms. The early and late components declined by 10–20 and 50–70% respectively in all cases. Fixing the gaze, in darkness, on an illuminated light spot stationary in relation to the head had no effect on level of response. Once the expected direction of body sway had been imparted, a significant and almost identical decrease of 70–80% in both components took place with the gaze fixed, however. Early and late components of vestibulomotor response are thought to be mediated by regulatory mechasisms with differing time courses and functional connections.Institute of Research into Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vo. 22, No. 1, pp. 80–87, January–February, 1990.     

Eye Movements Induced by Changes in the Internal Representation of Body Posture

Oculomotor responses to body rotation were investigated in subjects standing with the eyes closed. A rotatable platform was used to provide body rotation relative to the space-stationary head or upper part of the body (fixation of the head; the head and the shoulders; and the head, the shoulders, and the pelvis). A slow rotation of the body about the longitudinal axis by ±6.5° within 10–150 s evoked an illusion of the upper part of the body turning in space, while the moving footplate was perceived as stationary in space. This illusion was accompanied by marked eye movements in the direction of the illusory rotation. In subjects grasping a rigid ground-based handle, the perception of body movements corresponded to the actual rotation of body parts. In this case, the amplitude of eye movements was substantially lower. It was concluded that the eye movement pattern depends not only on the actual relative movement of the body segments but also on the perception of this movement relative to the extrapersonal space.     

Brain function in antarctic fish: frequency response analysis of central vestibular units

Summary Extracellular single-unit recordings were made from central vestibular units responding to horizontal head rotation in antarctic fish (Pagothenia borchgrevinki) at temperatures below 0 °C. The frequency of head rotation was varied between 0.05–16 Hz and the results were analysed in terms of the phase and gain of the response with respect to the stimulus. The model of the peripheral vestibular system derived by Hartmann and Klinke (1980) was fitted to the data from antarctic fish in order to obtain a quantitative comparison of vestibular function at two different temperatures. Despite the 20 °C temperature difference, and the different recording sites (primary afferents in the goldfish, and central vestibular units in the antarctic fish) vestibular function in antarctic fish is comparable to that previously reported in goldfish.     

Artificial vestibular feedback in conditions of modified body scheme

Subjects standing in darkness on the rigid support, kept a vertical posture which was destabilized by vibration of the Achilles tendons. To create a feedback on the vestibular input, transmastoidal bipolar galvanic stimulation was used. Changes of current in the feedback contour looked as linear function considering amplitude and velocity of the subject's head displacements in reference to the vertical. To change the body scheme we used some posture configurations: turning of the head in relation to the trunk; turning of the trunk with the head fixed; joint turning of the head and trunk. As a result of these configurations, the head could be turned approximately at right angle in relation to the feet. In addition turning of one foot at right angle in relation to the other foot was used. Artificial feedback reduces body fluctuations caused by vibration only in the vertical plane which passes through interaural axis of the head. The authors assume that directional changes of vestibulo-motor responses and results of application of artificial feedback during changes of orientation of the head in relation to the feet can be connected to change of ensembles of vestibular hair cells, which signals dominate in responses of vestibulo-spinal neurones.     

The effects of lesioning both the superior colliculus and the substantia nigra of cats on turning behavior

In twenty two adult cats, distributed in four groups, stainless steel electrodes were implanted in the superior colliculus and the substantia nigra of both sides in order: 1) to find the current intensity threshold values necessary to evoke turning behavior, and record their variations after lesion of the cited structures; 2) to study the effects of lesioning two of these structures, specifically related to the direction of turning behavior, and 3) to assess the time-course of recovery from postural asymmetry after damaging two structures involved in rotation behavior, located either in the same or in the opposite side, as well as the importance of performing these lesions simultaneously or at different periods. Three main results were observed: 1) a large proportion of lesioned cats showed an increase in threshold values necessary to evoke rotation of the implanted structures located either in the same or in the opposite side; 2) the lesions induced in a significant number of cats a transient postural asymmetry. After lesioning the superior colliculus, the direction of turning was towards the damaged hemisphere. Apomorphine injected fourteen days later demonstrated the existence of an occult asymmetry, and the direction of turning was maintained. In the substantia nigra lesioned animals, the direction of turning, was towards the non-lesioned side. Apomorphine reversed the direction of turning; 3) the cats showed a remarkable capacity to recover from the postural asymmetry produced by the lesion. This experimental series further support the hypothesis of a close functional relationship between structures of both cerebral hemispheres related to turning behavior.     

Lumbar and cervical erector spinae fatigue elicit compensatory postural responses to assist in maintaining head stability during walking.

The purpose of this study was to examine how inducing fatigue of the 1) lumbar erector spinae and 2) cervical erector spinae (CES) muscles affected the ability to maintain head stability during walking. Triaxial accelerometers were attached to the head, upper trunk, and lower trunk to measure accelerations in the vertical, anterior-posterior, and mediolateral directions during walking. Using three accelerometers enabled two adjacent upper body segments to be defined: the neck segment and trunk segment. A transfer function was applied to root mean square acceleration, peak power, and harmonic data derived from spectral analysis of accelerations to quantify segmental gain. The structure of upper body accelerations were examined using measures of signal regularity and smoothness. The main findings were that head stability was only affected in the anterior-posterior direction, as accelerations of the head were less regular following CES fatigue. Furthermore, following CES fatigue, the central nervous system altered the attenuation properties of the trunk segment in the anterior-posterior direction, presumably to enhance head stability. Following lumbar erector spinae fatigue, the trunk segment had greater gain and increased regularity and smoothness of accelerations in the mediolateral direction. Overall, the results of this study suggest that erector spinae fatigue differentially altered segmental attenuation during walking, according to the level of the upper body that was fatigued and the direction that oscillations were attenuated. A compensatory postural response was not only elicited in the sagittal plane, where greater segmental attenuation occurred, but also in the frontal plane, where greater segmental gain occurred.     

Analysis of surface wave direction by the lateral line system of Xenopus: Source localization before and after inactivation of different parts of the lateral line

The turning responses of clawed toads (Xenopus laevis) to surface waves were examined in animals with an intact lateral line or with different combinations of lateral lines reversibly inactivated by CoCl₂. The responses were characterized with respect to response frequency, turning accuracy, turning side, response time, and swim distance. After the inactivation most animals still responded to surface waves but the responses were different from those of animals with an intact lateral line. They also differed according to the combination of inactivated lines. In all experiments the responses for stimuli in some sectors of the surface did not differ from controls. The location of these sectors co-varied with the position of the intact lines, i.e., normal responses were found for frontal stimulus directions when head lines were intact and for caudolateral stimulus directions when trunk lines were intact. Their size was larger when lines on both sides of the body were intact and smaller when only lines on one side were intact. When the number of functional lines was reduced to one or two on one side of the body the turning angles shown within the sector of normal responses were maintained for stimulus directions outside these sectors. These results can be interpreted as indicating that head and trunk lines represent different position values. When only a single line was functional the toads still turned towards the stimulus source more often than by chance.It is hypothesized that Xenopus uses two mechanisms to determine the direction of surface waves. One uses the position values of head and trunk lines; this mechanism is comparable to the place value postulated for individual head neuromasts of surface feeding fish. The other uses the information encoded in the activity pattern that is elicited in one line when the surface wave travels over the line. This second mechanism yields information about stimulus side but not about stimulus angle.     

Behavioral analysis of polarization vision in tethered flying locusts

For spatial navigation many insects rely on compass information derived from the polarization pattern of the sky. We demonstrate that tethered flying desert locusts (Schistocerca gregaria) show e-vector-dependent yaw-torque responses to polarized light presented from above. A slowly rotating polarizer (5.3° s^–1) induced periodic changes in yaw torque corresponding to the 180° periodicity of the stimulus. Control experiments with a rotating diffuser, a weak intensity pattern, and a stationary polarizer showed that the response is not induced by intensity gradients in the stimulus. Polarotaxis was abolished after painting the dorsal rim areas of the compound eyes black, but remained unchanged after painting the eyes except the dorsal rim areas. During rotation of the polarizer, two e-vectors (preferred and avoided e-vector) induced no turning responses: they were broadly distributed from 0 to 180° but, for a given animal, were perpendicular to each other. The data demonstrate polarization vision in the desert locust, as shown previously for bees, flies, crickets, and ants. Polarized light is perceived through the dorsal rim area of the compound eye, suggesting that polarization vision plays a role in compass navigation of the locust.     

Chlorophylls and carotenoids states in vivo I - A linear dichroism study of pigments orientation in spinach chloroplasts

Using a spectropolarimeter for measuring the linear dichroism of pigments in oriented spinach chloroplasts we found a composite signal that could be interpreted neither as textural dichroism nor as a sample birefringence. We detected a fairly good orientation of pigments with respect to the normal at the plane of chloroplast lamellae. We failed to show any orientation axe in this plane. We found that all the Ca 683 is oriented, its Y direction being parallel to, or lying in the lamellae plane. Ca 673 is either unoriented or is oriented with its Y direction making an angle of 55° with the normal. If Ca 673 is unoriented, then the X direction of Ca 683 could be space positionned at about 45° of the lamellae plane. Carotenoids are oriented in the lamellar plane or close to it. Cb is equally oriented.     

Flight-phase and visual-field related optomotor yaw responses in gregarious desert locusts during tethered flight

Tethered flying desert locusts, Schistocerca gregaria, generate yaw-torque in response to rotation of a radial grating located beneath them. By screening parts of the pattern, rotation of the unscreened grating turned out to induce a compensatory steering (by pattern motion within transversally oriented 90° wide sectors) as well as an upwind/downwind turning response (by pattern motion within the anterior ventral 90° wide sector). The strength and polarity of responses upon the unscreened grating results from a linear superposition of these two response components. The results are discussed with regard to a functional specialization of eye regions.In a typical experiment, 3 consecutive flight-phases, assumed to mirror start, long-range flight, and landing of a free-flying locust, were distinguished. They may result from a time dependent variation of the polarity and relative strength of upwind/downwind turning and compensatory steering responses. Starting and landing phases were under strong optomotor control and were dominated by the high-gain compensatory steering. In contrast, the phase of long-range flight was under weak optomotor control resulting from a low gain in both of the two response components. The biological significance of this variable strength of optomotor control on free flight orientation of swarming locusts is discussed.     

On the encoding of movement vectors by honeybees. Are distance and direction represented independently?

Honeybees flying repeatedly over the same trajectory link it to an associated visual stimulus such that on viewing the stimulus they perform a trajectory in the habitual direction. To test if trajectory length can also be linked to a visual stimulus, bees were trained to fly through a multi-comparmented maze. Bees flew through a multi-compartmented maze. In one compartment a short trajectory could be linked to a stripe pattern oriented at 45° to the horizontal. In another compartment a longer trajectory could be linked to 135° stripes. Bees made both associations: their trajectories were short when viewing 45° stripes and longer when viewing 135° stripes. 90° stripes evoked trajectories of intermediate length.To test if distance and direction are linked independently to stripe orientation, a bee's trajectory was linked to 135° stripes in one compartment and to 45° stripes in another. These trajectories were the same length but differed in their horizontal direction by 60° or by 120°. 90° stripes evoked trajectories of intermediate direction which were shorter than those elicited by either training pattern. Bees were also trained to generate one long and one short trajectory with directions 120° apart. The trajectories elicited by 90° stripes were then biased towards the direction of the long training vector. Length and direction are not treated separately. The rules for combining trajectories resemble those of vector averaging.     

A TEST OF TWO POSSIBLE MECHANISMS FOR PHOTOTACTIC STEERING IN VOLVOX CARTERI (CHLOROPHYCEAE)

We tested two competing models that could explain how differential flagellar activity leads to phototactic turning in spheroids of Volvox carteri f. weismannia (Powers) Iyengar. In one model, turning results from the flagella of anterior cells in the lighted and shadowed hemispheres beating at different frequencies. In a competing model, turning results from a change in beat direction in these flagella. Both models successfully explain phototactic steering under constant illumination, but they make different predictions when colonies are exposed to abrupt changes in light intensity. If turning is due to control of flagellar beat frequency, both progression and rotation rates will change in the same direction and with similar magnitudes. If spheroid turning is due to a change in flagellar beat direction, a decreased rate of progression will accompany an increased rate of rotation and vice versa. We used video-microscopy to observe the behavior of positively phototactic V. carteri spheroids exposed to 10× step-up and step-down stimuli. After a step-up stimulus, spheroids slow their progression and rotation by equal amounts. No significant changes are reported in these parameters after the reciprocal step-down response. These observations are consistent with the variable flagellar frequency model and inconsistent with the variable flagellar direction model for phototactic turning. Switching the direction of light stimulus by 180° results in reorientation of positively phototactic spheroids. The kinetics of this reorientation did not precisely match the predictions of either model.     

Direction-dependent neck and trunk postural reactions during sitting

Postural reactions in healthy individuals in the seated position have previously been described and have been shown to depend on the direction of the perturbation; however the neck response following forward and backward translations has not been compared. The overall objective of the present study was to compare neck and trunk kinematic, kinetic and electromyographic (EMG) stabilization patterns of seated healthy individuals to forward and backward translations. Ten healthy individuals, seated on a chair fixed onto a movable platform, were exposed to forward and backward translations (distance = 0.15 m, peak acceleration = 1.2 m/s²). The head and trunk kinematics as well as the EMG activity of 16 neck and trunk muscles were recorded. Neck and trunk angular displacements were computed in the sagittal plane. The centers of mass (COMs) of the head (HEAD), upper thorax (UPTX), lower thorax (LOWTX) and abdomen (ABDO) segments were also computed. Moments of force at the C7-T1 and L5-S1 levels were calculated using a top-down, inverse dynamics approach. Forward translations provoked greater overall COM peak displacements. The first peak of moment of force was also reached earlier following forward translations which may have played a role in preventing the trunk from leaning backwards. These responses can be explained by the higher postural threat imposed by a forward translation.     

Modulation of Visually Evoked Postural Responses by Contextual Visual,Haptic and Auditory Information: A ‘Virtual Reality Check’

Externally generated visual motion signals can cause the illusion of self-motion in space (vection) and corresponding visually evoked postural responses (VEPR). These VEPRs are not simple responses to optokinetic stimulation, but are modulated by the configuration of the environment. The aim of this paper is to explore what factors modulate VEPRs in a high quality virtual reality (VR) environment where real and virtual foreground objects served as static visual, auditory and haptic reference points. Data from four experiments on visually evoked postural responses show that: 1) visually evoked postural sway in the lateral direction is modulated by the presence of static anchor points that can be haptic, visual and auditory reference signals; 2) real objects and their matching virtual reality representations as visual anchors have different effects on postural sway; 3) visual motion in the anterior-posterior plane induces robust postural responses that are not modulated by the presence of reference signals or the reality of objects that can serve as visual anchors in the scene. We conclude that automatic postural responses for laterally moving visual stimuli are strongly influenced by the configuration and interpretation of the environment and draw on multisensory representations. Different postural responses were observed for real and virtual visual reference objects. On the basis that automatic visually evoked postural responses in high fidelity virtual environments should mimic those seen in real situations we propose to use the observed effect as a robust objective test for presence and fidelity in VR.     

Saccadic head and thorax movements in freely walking blowflies

Visual information processing is adapted to the statistics of natural visual stimuli, and these statistics depend to a large extent on the movements of an animal itself. To investigate such movements in freely walking blowflies, we measured the orientation and position of their head and thorax, with high spatial and temporal accuracy. Experiments were performed on Calliphora vicina, Lucilia cuprina and L. caesar. We found that thorax and head orientation of walking flies is typically different from the direction of walking, with differences of 45° common. During walking, the head and the thorax turn abruptly, with a frequency of 5–10 Hz and angular velocities in the order of 1,000°/s. These saccades are stereotyped: head and thorax start simultaneously, with the head turning faster, and finishing its turn before the thorax. The changes in position during walking are saccade-like as well, occurring synchronously, but on average slightly after the orientation saccades. Between orientation saccades the angular velocities are low and the head is held more stable than the thorax. We argue that the strategy of turning by saccades improves the performance of the visual system of blowflies.     

Anticipatory postural adjustments to arm movement reveal complex control of paraspinal muscles in the thorax

Anatomical and empirical data suggest that deep and superficial muscles may have different functions for thoracic spine control. This study investigated thoracic paraspinal muscle activity during anticipatory postural adjustments associated with arm movement. Electromyographic (EMG) recordings were made from the right deep (multifidus/rotatores) and superficial (longissimus) muscles at T5, T8, and T11 levels using fine-wire electrodes. Ten healthy participants performed fast unilateral and bilateral flexion and extension arm movements in response to a light. EMG amplitude was measured during 25 ms epochs for 150 ms before and 400 ms after deltoid EMG onset. During arm flexion movements, multifidus and longissimus had two bursts of activity, one burst prior to deltoid and a late burst. With arm extension both muscles were active in a single burst after deltoid onset. There was differential activity with respect to direction of trunk rotation induced by arm movement. Right longissimus was most active with left arm movements and right multifidus was most active with right arm movements. All levels of the thorax responded similarly. We suggest that although thoracic multifidus and longissimus function similarly to control sagittal plane perturbations, these muscles are differentially active with rotational forces on the trunk.     

Factors influencing codling moth larval response to α-farnesene

Behavioral responses of newly-emerged codling moth (Cydia pomonella L.) larvae to α-farnesene were compared for a laboratory-reared strain (‘lab’, 160 generations inbred) and a recently collected strain (‘wild’, 3 generations). Video recordings of single larvae placed 20 mm from Bond papers releasing a range of seven α-farnesene concentrations were scored for head turning, head lifting, head direction, and movement across a Petri dish. The laboratory strain was significantly less successful at finding the treated papers than the wild strain. The lab strain was also significantly less active, shown by more random orientation, slower walking speed, less head turning and less head lifting. Both strains showed lower response to α-farnesene concentrations less than 10⁻⁶ (w/v) α-farnesene. Orientation, walking speed, and head turning rate were significantly higher in the presence of α-farnesene concentrations >10⁻⁷ w/v, for both strains. Head lifting did not show any effect from α-farnesene. Head turning was associated with both head lifting and walking speed and this association improved with the presence of α-farnesene. Increased attraction to odour source was correlated with improved direction finding at concentrations above 10⁻⁷ w/v. Activity factors such as head turning and walking speed influenced time to locate odour source more than orientation factors. Odour-guided orientation to α-farnesene in codling moth larvae was composed of both locomotory, and to a lesser extent, orientation responses, which increased in a directed fashion to the stimulus.