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Biomedical subjects

Sergei B Yakushin

Publications and source records attributed to Sergei B Yakushin.

15 recordsLinked to original sources
PubMed2006

Dynamics of quadrupedal locomotion of monkeys: implications for central control.

We characterized the three-dimensional kinematics and dynamics of quadrupedal gait of young adult rhesus and cynomolgus monkeys while they walked with diagonal and lateral gaits at 0.4-1.0 m/s on a treadmill. Rigid bodies on the wrist, ankle, and back were monitored by an optical motion detection system (Optotrak). Kinematic data could be normalized using characteristic stride length, reducing variance due to different gait styles, to emphasize common characteristics of swing and stance parameters among animals. Mean swing phase durations fell as walking speed increased, but the swing phase durations increased at each walking velocity as a linear function of increases in amplitude, thereby following a main sequence relationship. The phase plane trajectories of the swing phases, i.e., plots of the relation of the rising and falling limb velocity to limb position in the sagittal (X-Z) plane, had unique dynamic characteristics. Trajectories were separable at each walking velocity and increases in swing amplitude were linearly related to peak swing velocities, thus comprising main sequences. We infer that the swing phase dynamics are set by central neural mechanisms at the onset of the swing phases according to the intended amplitude, which in turn is based on the walking velocity in the stance phases. From the many dynamic similarities between swing phases and rapid eye movements, we further suggest that the swing phases may be generated by neural mechanisms similar to those that produce saccades and quick phases of nystagmus from a signal related to sensed or desired walking velocity.

Animals↗
PubMed2006

Modeling gravity-dependent plasticity of the angular vestibuloocular reflex with a physiologically based neural network.

A neural network model was developed to explain the gravity-dependent properties of gain adaptation of the angular vestibuloocular reflex (aVOR). Gain changes are maximal at the head orientation where the gain is adapted and decrease as the head is tilted away from that position and can be described by the sum of gravity-independent and gravity-dependent components. The adaptation process was modeled by modifying the weights and bias values of a three-dimensional physiologically based neural network of canal-otolith-convergent neurons that drive the aVOR. Model parameters were trained using experimental vertical aVOR gain values. The learning rule aimed to reduce the error between eye velocities obtained from experimental gain values and model output in the position of adaptation. Although the model was trained only at specific head positions, the model predicted the experimental data at all head positions in three dimensions. Altering the relative learning rates of the weights and bias improved the model-data fits. Model predictions in three dimensions compared favorably with those of a double-sinusoid function, which is a fit that minimized the mean square error at every head position and served as the standard by which we compared the model predictions. The model supports the hypothesis that gravity-dependent adaptation of the aVOR is realized in three dimensions by a direct otolith input to canal-otolith neurons, whose canal sensitivities are adapted by the visual-vestibular mismatch. The adaptation is tuned by how the weights from otolith input to the canal-otolith-convergent neurons are adapted for a given head orientation.

Adaptation, Physiological↗
PubMed2005

Spatial properties of central vestibular neurons.

We studied the spatial characteristics of 45 vestibular-only (VO) and 12 vestibular-plus-saccade (VPS) neurons in two cynomolgus monkeys using angular rotation and static tilt. The purpose was to determine the contribution of canal and otolith-related inputs to central vestibular neurons whose activity is associated with the central velocity storage integrator. Lateral canal-related neurons responded maximally during vertical axis rotation when the head was tilted 25 +/- 6 and 22 +/- 3 degrees forward relative to the axis of rotation in the two animals, and vertical canal-related neurons responded maximally with the head tilted back 63+/- 5 and 57 +/- 7 degrees . The origin of the vertical canal-related input was verified by rotation about a spatial horizontal axis. Thirty-one percent of cells received input in a single canal plane. Sixty-seven percent of canal-related cells received otolith input, 31% of vertical canal neurons had lateral canal input, and 43% of lateral canal neurons had vertical canal input. Twenty percent of neurons had convergent input from the lateral canals, the vertical canals, and the otolith organs. Some VO and VPS cells had spatial-temporal convergent (STC) properties; more of these cells had STC properties at lower frequencies of rotation. Thus VO and VPS neurons associated with velocity storage receive a broad range of convergent inputs from each portion of the vestibular labyrinth. This convergence could provide the basis for gravity-dependent eye velocity orientation induced through velocity storage.

Acceleration↗
PubMed2005

Spatial properties of central vestibular neurons of monkeys after bilateral lateral canal nerve section.

Thirty-seven neurons were recorded in the superior vestibular nucleus (SVN) of two cynomolgus monkeys 1-2 yr after bilateral lateral canal nerve section to test whether the central neurons had spatially adapted for the loss of lateral canal input. The absence of lateral canal function was verified with eye movement recordings. The relation of unit activity to the vertical canals was determined by oscillating the animals about a horizontal axis with the head in various orientations relative to the axis of rotation. Animals were also oscillated about a vertical axis while upright or tilted in pitch. In the second test, the vertical canals are maximally activated when the animals are tilted back about -50 degrees from the spatial upright and the lateral canals when the animals are tilted forward about 30 degrees . We reasoned that if central compensation occurred, the head orientation at which the response of the vertical canal-related neurons was maximal should be shifted toward the plane of the lateral canals. No lateral canal-related units were found after nerve section, and vertical canal-related units were found only in SVN not in the rostral medial vestibular nucleus. SVN canal-related units were maximally activated when the head was tilted back at -47 +/- 17 and -50 +/- 12 degrees (means +/- SD) in the two animals, close to the predicted orientation of the vertical canals. This indicated that spatial adaptation of vertical canal-related vestibular neurons had not occurred. There were substantial neck and/or otolith-related inputs activating the vertical canal-related neurons in the nerve-sectioned animals, which could have contributed to oculomotor compensation after nerve section.

Action Potentials↗
PubMed2005

Spatial distribution of gravity-dependent gain changes in the vestibuloocular reflex.

This study determined whether dependence of angular vestibuloocular reflex (aVOR) gain adaptation on gravity is a fundamental property in three dimensions. Horizontal aVOR gains were adaptively increased or decreased in two cynomolgus monkeys in upright, side down, prone, and supine positions, and aVOR gains were tested in darkness by yaw rotation with the head in a wide variety of orientations. Horizontal aVOR gain changes peaked at the head position in which the adaptation took place and gradually decreased as the head moved away from this position in any direction. The gain changes were plotted as a function of head tilt and fit with a sinusoid plus a bias to obtain the gravity-dependent (amplitude) and gravity-independent (bias) components. Peak-to-peak gravity-dependent gain changes in planes containing the position of adaptation and the magnitude of the gravity-independent components were both approximately 25%. We assumed that gain changes over three-dimensional space could be described by a sinusoid the amplitude of which also varied sinusoidally. Using gain changes obtained from the head position in which the gains were adapted, a three-dimensional surface was generated that was qualitatively similar to a surface obtained from the experimental data. This extends previous findings on vertical aVOR gain adaptation in one plane and introduces a conceptual framework for understanding plasticity in three dimensions: aVOR gain changes are composed of two components, one of which depends on head position relative to gravity. It is likely that this gravitational dependence optimizes the stability of retinal images during movement in three-dimensional space.

Adaptation, Physiological↗
PubMed2005

The role of gravity in adaptation of the vertical angular vestibulo-ocular reflex.

The gain of the vertical angular vestibulo-ocular reflex (aVOR) was adapted in side-down and prone positions in two monkeys and tested in four planes: left-/right-side down; forward/backward; and two intermediate planes that lie approximately in the planes of the vertical semicircular canal pairs, left anterior/right posterior (LA/RP) and right anterior/left posterior (RA/LP). Gain changes, expressed as a percent of preadapted values, were plotted as a function of head orientation in the planes of tilt, and fitted with sinusoids to obtain the gravity-dependent (amplitude) and gravity-independent (bias) components of adaptation. Gravity-dependent gain changes were always maximal when tested in a plane that included the head orientation in which the aVOR gain had been adapted. Changes were minimal when the head was tilted in a plane orthogonal to the plane of adaptation, and were smaller but still significant when tested in the two intermediate planes. Gravity-independent VOR gain changes were uniform over all planes of head tilt. Thus, the gravity-dependent and gravity-independent components could be separated experimentally. The aVOR gain changes from the head tilts in different directions were utilized to reconstruct the gain changes in three dimensions. They formed a continuous surface, which peaked in and around the position of adaptation. These studies support the postulate that gain adaptation has both gravity-independent and gravity-dependent components, and further show that these gain changes have a three-dimensional structure. These results are similar to those in humans, indicating that the gravity-dependent adaptation of the aVOR is likely to be a common phenomenon across species.

Acclimatization↗
PubMed2004

Gravity-dependent and gravity-independent gain changes during vertical vestibulo-ocular reflex (VOR) adaptation.

The gain of the vertical angular vestibulo-ocular reflex (aVOR) was adaptively increased or decreased with monkeys in a side down position, and the gains were tested with the axis of rotation tilted in 10 degrees increments from left- to right-side-down. Gain changes, expressed as a percentage of the preadapted values, were plotted as a function of head tilt, and fit with a cosine function. The amplitude of the cosine was half of the gravity-dependent component of the gain change and the bias, the gravity independent component. The largest changes in the gain of both components occurred in the first 30 min and continued at a slower rate throughout adaptation. The gravity-dependent and -independent gain changes were larger for gain decreases than for gain increases, but both components had similar dynamics. We conclude that the alteration in gain of the aVOR always occurs in the context of gravity.

Adaptation, Physiological↗
PubMed2003

Dependence of adaptation of the human vertical angular vestibulo-ocular reflex on gravity.

We determined the spatial dependence of adaptive gain changes of the vertical angular vestibulo-ocular reflex (aVOR) on gravity in five human subjects. The gain was decreased for 1 h by sinusoidal oscillation in pitch about a spatial vertical axis in a subject-stationary surround with the head oriented left-side down. Gains were tested by sinusoidal oscillation about a spatial vertical axis while subjects were tilted in 15 degrees increments from left- to right-side down positions through the upright. Changes in gain of the vertical component of the induced eye movements were expressed as a percentage of the preadapted values for the final analysis. Vertical aVOR gain changes were maximal in the position in which the gain had been adapted and declined progressively as subjects were moved from this position. Gain changes were plotted as a function of head orientation and fit with a sine function. The bias level of the fitted sines, i.e., the gravity-independent gain change, was -29+/-10% (SD). The gains varied around this bias as a function of head position by +/-18+/-6%, which were the gravity-dependent gain changes. The gravity-dependent gain changes induced by only 1 h of adaptation persisted, gradually declining over several days. We conclude that there is a component of the vertical aVOR gain change in humans that is dependent on the head orientation in which the gain was adapted, and that this dependence can persist for substantial periods.

Adaptation, Physiological↗
PubMed2003

Gravity-specific adaptation of the angular vestibuloocular reflex: dependence on head orientation with regard to gravity.

The gain of the vertical angular vestibuloocular reflex (aVOR) was adaptively altered by visual-vestibular mismatch during rotation about an interaural axis, using steps of velocity in three head orientations: upright, left-side down, and right-side down. Gains were decreased by rotating the animal and visual surround in the same direction and increased by visual and surround rotation in opposite directions. Gains were adapted in one head position (single-state adaptation) or decreased with one side down and increased with the other side down (dual-state adaptation). Animals were tested in darkness using sinusoidal rotation at 0.5 Hz about an interaural axis that was tilted from horizontal to vertical. They were also sinusoidally oscillated from 0.5 to 4 Hz about a spatial vertical axis in static tilt positions from yaw to pitch. After both single- and dual-state adaptation, gain changes were maximal when the monkeys were in the position in which the gain had been adapted, and the gain changes progressively declined as the head was tilted away from that position. We call this gravity-specific aVOR gain adaptation. The spatial distribution of the specific aVOR gain changes could be represented by a cosine function that was superimposed on a bias level, which we called gravity-independent gain adaptation. Maximal gravity-specific gain changes were produced by 2-4 h of adaptation for both single- and dual-state adaptations, and changes in gain were similar at all test frequencies. When adapted while upright, the magnitude and distribution of the gravity-specific adaptation was comparable to that when animals were adapted in side-down positions. Single-state adaptation also produced gain changes that were independent of head position re gravity particularly in association with gain reduction. There was no bias after dual-state adaptation. With this difference, fits to data obtained by altering the gain in separate sessions predicted the modulations in gain obtained from dual-state adaptations. These data show that the vertical aVOR gain changes dependent on head position with regard to gravity are continuous functions of head tilt, whose spatial phase depends on the position in which the gain was adapted. From their different characteristics, it is likely that gravity-specific and gravity-independent adaptive changes in gain are produced by separate neural processes. These data demonstrate that head orientation to gravity plays an important role in both orienting and tuning the gain of the vertical aVOR.

Adaptation, Physiological↗
PubMed2003

Adaptive changes in the angular VOR: duration of gain changes and lack of effect of nodulo-uvulectomy.

Alterations in the gain of the vertical angular vestibulo-ocular reflex (VOR) are dependent on the head position in which the gain changes were produced. We determined how long gravity-dependent gain changes last in monkeys after four hours of adaptation, and whether the adaptation is mediated through the nodulus and uvula of the vestibulocerebellum. Vertical VOR gains were adaptively modified by rotation about an interaural axis, in phase or out of phase with the visual surround. Vertical VOR gains were modified with the animals in one of three orientations: upright, left-side down, or right-side down. Monkeys were tested in darkness for up to four days after adaptation using sinusoidal rotation about an interaural axis that was incrementally tilted in 10 degrees steps from vertical to side down positions. Animals were unrestrained in their cages in normal light conditions between tests. Gravity-dependent gain changes lasted for a day or less after adaptation while upright, but persisted for two days or more after on-side adaptation. These data show that gravity-dependent gain changes can last for prolonged periods after only four hours of adaptation in monkeys, as in humans. They also demonstrate that natural head movements made while upright do not provide an adequate stimulus for rapid recovery of vertical VOR gains that were induced on side. In two animals, the nodulus and uvula were surgically ablated. Vertical gravity-dependent gain changes were not significantly different before and after surgery, indicating that the nodulus and uvula do not have a critical role in producing them.

Adaptation, Physiological↗
PubMed2002

Spatial orientation of caloric nystagmus.

The spatial orientation of the slow-phase eye velocity of caloric nystagmus was investigated in cynomolgus monkeys after all six semicircular canals had been plugged. Normal animals generate responses that have dominant convective components produced by movement of the endolymph in the lateral canal toward or away from gravity. As a result, the direction of horizontal slow-phase velocity induced by cold-water irrigation changes direction with changes in head position with regard to gravity. Plugging produced a dense overgrowth of bone that blocked the flow of endolymph, but the end organs were intact. Robust caloric nystagmus was elicited after recovery, but the horizontal (yaw) component was now always toward the stimulated (ipsilateral) side, regardless of head position re gravity. The induced caloric nystagmus had strong spatial orientation properties after canal plugging. With animals upright, the three-dimensional velocity vector of the caloric nystagmus was close to the yaw axis with small vertical and roll components. Roll components became stronger in supine and prone positions and vertical components were enhanced in the right- and left-side down positions. In each instance, the addition of the roll and vertical components moved the velocity vector of the nystagmus closer to the spatial vertical. Modeling supported the postulate that the caloric nystagmus after canal plugging is influenced by three factors: (1) a reduction in neural activity in the ampullary nerves on the stimulated side due to cooling of the nerves; (2) contraction of the endolymph in the closed space between the cupula and the plug due to cooling, which resulted in deflection of the cupula and hair cells toward the plug (ampullofugal deflection); and (3) alignment of eye velocity to gravity due to the orientation properties of velocity storage. Although convection is the most prominent factor in producing caloric responses in the normal state, our results suggest that alteration of nerve activity due to thermal effects, endolymph contraction or expansion, and velocity storage are also likely to contribute to the total response.

Animals↗
PubMed2002

The nodulus and uvula: source of cerebellar control of spatial orientation of the angular vestibulo-ocular reflex.

The nodulus and rostral-ventral uvula of the vestibulo-cerebellum play a critical role in orienting eye velocity of the slow component of the angular vestibulo-ocular reflex (aVOR) to gravito-inertial acceleration (GIA). This is done by altering the time constants of "velocity storage" in the vestibular system and by generating "cross-coupled" eye velocities that shift the eye velocity vector from along the body yaw axis to the yaw axis in a spatial frame. In this report, we show that eye velocity generated through the aVOR by constant velocity centrifugation in the monkey orients to the GIA in space, regardless of the position of the head with respect to the axis of rotation. We also show that, after removal of the nodulus and rostral-ventral uvula, the spatial orientation of eye velocity to the GIA is lost and that eye velocity is then purely driven by the semicircular canals in a body frame of reference. These findings are further confirmation that these regions of the vestibulo-cerebellum control spatial orientation of the aVOR.

Animals↗
PubMed2002

Compensatory and orienting eye movements induced by off-vertical axis rotation (OVAR) in monkeys.

Nystagmus induced by off-vertical axis rotation (OVAR) about a head yaw axis is composed of a yaw bias velocity and modulations in eye position and velocity as the head changes orientation relative to gravity. The bias velocity is dependent on the tilt of the rotational axis relative to gravity and angular head velocity. For axis tilts <15 degrees, bias velocities increased monotonically with increases in the magnitude of the projected gravity vector onto the horizontal plane of the head. For tilts of 15-90 degrees, bias velocity was independent of tilt angle, increasing linearly as a function of head velocity with gains of 0.7-0.8, up to the saturation level of velocity storage. Asymmetries in OVAR bias velocity and asymmetries in the dominant time constant of the angular vestibuloocular reflex (aVOR) covaried and both were reduced by administration of baclofen, a GABA(B) agonist. Modulations in pitch and roll eye positions were in phase with nose-down and side-down head positions, respectively. Changes in roll eye position were produced mainly by slow movements, whereas vertical eye position changes were characterized by slow eye movements and saccades. Oscillations in vertical and roll eye velocities led their respective position changes by approximately 90 degrees, close to an ideal differentiation, suggesting that these modulations were due to activation of the orienting component of the linear vestibuloocular reflex (lVOR). The beating field of the horizontal nystagmus shifted the eyes 6.3 degrees /g toward gravity in side down position, similar to the deviations observed during static roll tilt (7.0 degrees /g). This demonstrates that the eyes also orient to gravity in yaw. Phases of horizontal eye velocity clustered ~180 degrees relative to the modulation in beating field and were not simply differentiations of changes in eye position. Contributions of orientating and compensatory components of the lVOR to the modulation of eye position and velocity were modeled using three components: a novel direct otolith-oculomotor orientation, orientation-based velocity modulation, and changes in velocity storage time constants with head position re gravity. Time constants were obtained from optokinetic after-nystagmus, a direct representation of velocity storage. When the orienting lVOR was combined with models of the compensatory lVOR and velocity estimator from sequential otolith activation to generate the bias component, the model accurately predicted eye position and velocity in three dimensions. These data support the postulates that OVAR generates compensatory eye velocity through activation of velocity storage and that oscillatory components arise predominantly through lVOR orientation mechanisms.

Animals↗
PubMed2002

Spatial orientation of caloric nystagmus in semicircular canal-plugged monkeys.

We studied caloric nystagmus before and after plugging all six semicircular canals to determine whether velocity storage contributed to the spatial orientation of caloric nystagmus. Monkeys were stimulated unilaterally with cold ( approximately 20 degrees C) water while upright, supine, prone, right-side down, and left-side down. The decline in the slow phase velocity vector was determined over the last 37% of the nystagmus, at a time when the response was largely due to activation of velocity storage. Before plugging, yaw components varied with the convective flow of endolymph in the lateral canals in all head orientations. Plugging blocked endolymph flow, eliminating convection currents. Despite this, caloric nystagmus was readily elicited, but the horizontal component was always toward the stimulated (ipsilateral) side, regardless of head position relative to gravity. When upright, the slow phase velocity vector was close to the yaw and spatial vertical axes. Roll components became stronger in supine and prone positions, and vertical components were enhanced in side down positions. In each case, this brought the velocity vectors toward alignment with the spatial vertical. Consistent with principles governing the orientation of velocity storage, when the yaw component of the velocity vector was positive, the cross-coupled pitch or roll components brought the vector upward in space. Conversely, when yaw eye velocity vector was downward in the head coordinate frame, i.e., negative, pitch and roll were downward in space. The data could not be modeled simply by a reduction in activity in the ipsilateral vestibular nerve, which would direct the velocity vector along the roll direction. Since there is no cross coupling from roll to yaw, velocity storage alone could not rotate the vector to fit the data. We postulated, therefore, that cooling had caused contraction of the endolymph in the plugged canals. This contraction would deflect the cupula toward the plug, simulating ampullofugal flow of endolymph. Inhibition and excitation induced by such cupula deflection fit the data well in the upright position but not in lateral or prone/supine conditions. Data fits in these positions required the addition of a spatially orientated, velocity storage component. We conclude, therefore, that three factors produce cold caloric nystagmus after canal plugging: inhibition of activity in ampullary nerves, contraction of endolymph in the stimulated canals, and orientation of eye velocity to gravity through velocity storage. Although the response to convection currents dominates the normal response to caloric stimulation, velocity storage probably also contributes to the orientation of eye velocity.

Animals↗