Stimulation and single-unit studies of velocity storage in the vestibular and prepositus hypoglossi nuclei of the monkey
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With our stimulus conditions we were unable to record more than 2-3 beats of OKAN; therefore direct comparison to the data recorded from monkeys is not possible. We did, however, see cross-coupling in OKN. In monkeys, cross-coupling predominates in OKAN, indicating that velocity storage underlies this phenomenon. We consistently saw the axis of response shift towards the spatial vertical. This implies that although OKAN was weak, velocity storage contributed a representation of the spatial vertical to OKN that is dependent on the axis of the head or body with respect to gravity.
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1. A simplified three-dimensional state space model of visual vestibular interaction was formulated. Matrix and dynamical system operators representing coupling from the semicircular canals and the visual system to the velocity storage integrator were incorporated into the model. 2. It was postulated that the system matrix for a tilted position was a composition of two linear transformations of the system matrix for the upright position. One transformation modifies the eigenvalues of the system matrix while another rotates the pitch and roll eigenvectors with the head, while maintaining the yaw axis eigenvector approximately spatially invariant. Using this representation, the response characteristics of the pitch, roll, and yaw eye velocity were obtained in terms of the eigenvalues and associated eigenvectors. 3. Using OKAN data obtained from monkeys and comparing to the model predictions, the eigenvalues and eigenvectors of the system matrix were identified as a function of tilt to the side or of tilt to the prone positions, using a modification of the Marquardt algorithm. The yaw eigenvector for right-side-down tilt and for downward pitch cross-coupling was approximately 30 degrees from the spatial vertical. For the prone position, the eigenvector was computed to be approximately 20 degrees relative to the spatial vertical. For both side-down and prone positions, oblique OKN induced along eigenvector directions generated OKAN which decayed to zero along a straight line with approximately a single time constant. This was verified by a spectral analysis of the residual sequence about the straight line fit to the decaying data. The residual sequence was associated with a narrow autocorrelation function and a wide power spectrum. 4. Parameters found using the Marquardt algorithm were incorporated into the model. Diagonal matrices in a head coordinate frame were introduced to represent the direct pathway and the coupling of the visual system to the integrator. Model simulations predicted the behavior of yaw and pitch OKN and OKAN when the animal was upright, as well as the cross-coupling in the tilted position. The trajectories in velocity space were also accurately simulated. 5. There were similarities between the monkey eigenvectors and human perception of the spatial vertical. For side-down tilts and downward eye velocity cross-coupling, there was only an Aubert (A) effect. For upward eye velocity cross-coupling there were both Muller (E) and Aubert (A) effects. The mean of the eigenvectors for upward and downward eye velocities overlay human 1 x g perceptual data.(ABSTRACT TRUNCATED AT 400 WORDS).
Multi-person teams are sometimes responsible for critical tasks, such as flying an airliner. Here we present a method using gaze tracking data to assess shared visual attention, a term we use to describe the situation where team members are attending to a common set of elements in the environment. Gaze data are quantized with respect to a set of N areas of interest (AOIs); these are then used to construct a time series of N dimensional vectors, with each vector component representing one of the AOIs, all set to 0 except for the component corresponding to the currently fixated AOI, which is set to 1. The resulting sequence of vectors can be averaged in time, with the result that each vector component represents the proportion of time that the corresponding AOI was fixated within the given time interval. We present two methods for comparing sequences of this sort, one based on computing the time-varying correlation of the averaged vectors, and another based on a chi-square test testing the hypothesis that the observed gaze proportions are drawn from identical probability distributions.We have evaluated the method using synthetic data sets, in which the behavior was modeled as a series of activities, each of which was modeled as a first-order Markov process. By tabulating distributions for pairs of identical and disparate activities, we are able to perform a receiver operating characteristic (ROC) analysis, allowing us to choose appropriate criteria and estimate error rates.We have applied the methods to data from airline crews, collected in a high-fidelity flight simulator (Haslbeck, Gontar Schubert, 2014). We conclude by considering the problem of automatic (blind) discovery of activities, using methods developed for text analysis.
Multi-person teams are sometimes responsible for critical tasks, such as flying an airliner. Here we present a method using gaze tracking data to assess shared visual attention, a term we use to describe the situation where team members are attending to a common set of elements in the environment. Gaze data are quantized with respect to a set of N areas of interest (AOIs); these are then used to construct a time series of N dimensional vectors, with each vector component representing one of the AOIs, all set to 0 except for the component corresponding to the currently fixated AOI, which is set to 1. The resulting sequence of vectors can be averaged in time, with the result that each vector component represents the proportion of time that the corresponding AOI was fixated within the given time interval. We present two methods for comparing sequences of this sort, one based on computing the time-varying correlation of the averaged vectors, and another based on a chi-square test testing the hypothesis that the observed gaze proportions are drawn from identical probability distributions. We have evaluated the method using synthetic data sets, in which the behavior was modeled as a series of "activities," each of which was modeled as a first-order Markov process. By tabulating distributions for pairs of identical and disparate activities, we are able to perform a receiver operating characteristic (ROC) analysis, allowing us to choose appropriate criteria and estimate error rates. We have applied the methods to data from airline crews, collected in a high-fidelity flight simulator (Haslbeck, Gontar & Schubert, 2014). We conclude by considering the problem of automatic (blind) discovery of activities, using methods developed for text analysis.
Human behavior often consists of a series of distinct activities, each characterized by a unique signature of visual behavior. This is true even in a restricted domain, such as piloting an aircraft, where patterns of visual signatures might represent activities like communicating, navigating, and monitoring. We propose a novel analysis method for gaze-tracking data, to perform blind discovery of these activities based on their behavioral signatures. The method is in some respects similar to recurrence analysis, but here we compare not individual fixations, but groups of fixations aggregated over a fixed time interval. The duration of this interval is a parameter that we will refer to as τ. We assume that the environment has been divided into a set of N different areas-of-interest (AOIs). For a given interval of time of duration τ, we compute the proportion of time spent fixating each AOI, resulting in an N-dimensional vector. These proportions can be converted to counts by multiplying by τ divided by the average fixation duration (another parameter that we fix at 280 milliseconds). We compare different intervals by computing the chi-square statistic. The p-value associated with the statistic is the likelihood of observing the data under the hypothesis that the data in the two intervals were generated by a single process with a single set of probabilities governing the fixation of each AOI. We have investigated the method using a set of 10 synthetic "activities," that sample 4 AOIs. Four of these activities visit 3 of the 4 AOIs, with equal probability; as there are four different ways to leave-one- out, there are four such activities. Similarly, there are six different activities that leave-two-out. Sequences of simulated behavior were generated by running each activity for 40 seconds, in sequence, for a total of 6.7 minutes. The figure to the right shows the matrix of chi-square statistics, using a value of 2.8 seconds for τ, corresponding to 10 fixations. Low values (dark) indicate poor evidence for activity differences, while high values (bright) indicate strong evidence. The dark squares along the main diagonal each correspond to the forty second intervals in which the activity was held constant; the 4x4 block at the lower left corresponds to the four leave-one-out activities, while the 6x6 block in the upper right corresponds to the leave-two-out activities. (The anti-diagonal pattern of white squares indicates those activity pairs that share no AOIs.) The chi-square values can be binarized by choosing a particular significance level; we are interested in grouping bins that represent the same activity, effectively accepting the null hypothesis. Therefore, we may adopt a relatively lax criterion; for example, choosing a p-value of 0.2 means that two behaviors that have only a 1-in-5 chance of being produced by a single activity might nevertheless be clustered together. We have explored several methods to perform clustering on the data and solving for the activity probabilities. Greedy methods begin by selecting the time bin that is similar to the most (or least) other bins, and then forming a cluster from it and all other non-discriminable bins. These methods show mediocre performance, as they do not take into account temporal contiguity. Preliminary results indicate that methods that "grow" clusters in time from seed points perform better.
Human behavior often consists of a series of distinct activities, each characterized by a unique pattern of interaction with the visual environment. This is true even in a restricted domain, such as a pilot flying an airplane; in this case, activities with distinct visual signatures might be things like communicating, navigating, monitoring, etc. We propose a novel analysis method for gaze-tracking data, to perform blind discovery of these hypothetical activities. We compare, not individual fixations, but groups of fixations aggregated over a fixed time interval (Tau). We assume that the environment has been divided into a finite set of discrete areas-of-interest (AOIs). For a given time interval, we compute the proportion of time spent fixating each AOI, resulting in an N-dimensional vector, where N is the number of AOIs. These proportions can be converted to integer counts by multiplying by Tau divided by the average fixation duration, a parameter that we fix at 283 milliseconds. We compare different intervals by computing the chi-squared statistic. The p-value associated with the statistic is the likelihood of observing the data under the hypothesis that the data in the two intervals were generated by a single process with a single set of probabilities governing the fixation of each AOI. We cluster the intervals, first by merging adjacent intervals that are sufficiently similar, optionally shifting the boundary between non-merged intervals to maximize the difference. Then we compare and cluster non-adjacent intervals. The method is evaluated using synthetic data generated by a hand-crafted set of activities. While the method generally finds more activities than put into the simulation, we have obtained agreement as high as 80 percent between the inferred activity labels and ground truth.
Oculomotor behavior is a powerful tool to probe brain function and disease. Previous studies have shown how fixational drift velocities depend on eccentricity (Bertolini et al., 2013; Romano et al., 2017) and how fixational drift can adapt after prolonged eccentric fixation (Otero-Millan et al., 2019). Here, eighteen subjects completed two experiments where we measured drift velocity while fixating a briefly flashed target, to remove retinal slip cues. In the first experiment, we measured their baseline relationship between fixational drift velocity and eye position. In the second experiment, we investigated the adaptation of that relationship after prolonged fixation at a very eccentric position. This adaptation usually manifests by the presence of rebound nystagmus (RN) upon returning to central gaze. Our results replicate the finding that drift increases at central fixation in the direction consistent with rebound nystagmus, that is, towards the previously held eccentric position, highlighting that drift velocity depends on the history of eye positions. When examining the entire relationship between drift velocity and eye position we find a mostly linear relationship but with increased slope relative to the baseline condition. However, the adaptation of the integrator cannot be modeled by a simple change in leakiness, as that would not result in increased drift at central gaze. Other mechanisms of adaptation such as shifts in the position or velocity set-points for gaze holding could be present but are indistinguishable with the current data.
Space motion sickness has become an operational concern in manned space flight. Considerable evidence exists that head movements in free fall, especially pitch movements, are provocative until adaptation occurs. The question arises whether space motion sickness is an unique nosological entity or is due to body movements in a nonterrestrial force environment, a force environment for which the body's dynamic sensory-motor adaptions to 1 G are no longer appropriate. To evaluate this issue, subjects were asked to make controlled head movements during exposure to high gravitoinertial force levels, 1.8-2.0 G, in parabolic flight maneuvers. Head movements in pitch with eyes open were most evocative of motion sickness, yaw movements with eyes covered were least provocative. This pattern is identical to that which occurs when the same types of head movements are made in the free fall phase of parabolic maneuvers. It appears that space motion sickness is the consequence of prolonged exposure to a nonterrestrial force background rather than of exposure to free fall per se.
It is pointed out that head movements play an important role in gaze. The interaction between eye and head movements involves both their shared role in directing gaze and the compensatory vestibular ocular reflex. The dynamics of head trajectories are discussed, taking into account the use of parameterization to obtain the peak velocity, peak accelerations, the times of these extrema, and the duration of the movement. Attention is given to the main sequence, neck muscle EMG and details of the head-movement trajectory, types of head model accelerations, the latency of eye and head movement in coordinated gaze, gaze latency as a function of various factors, and coordinated gaze types. Clinical examples of gaze-plane analysis are considered along with the instantaneous change of compensatory eye movement (CEM) gain, and aspects of variability.
The goal of the present study was to determine if adaptive modulation of vestibulo-ocular reflex (VOR) function is associated with commensurate alterations in manual target localization. To measure the effects of adapted VOR on manual responses we developed the Vestibular-Contingent Pointing Test (VCP). In the VCP test, subjects pointed to a remembered target following passive whole body rotation in the dark. In the first experiment, subjects performed VCP before and after wearing 0.5X minifying lenses that adaptively attenuate horizontal VOR gain. Results showed that adaptive reduction in horizontal VOR gain was accompanied by a commensurate change in VCP performance. In the second experiment, bilaterally labyrinthine deficient (LD) subjects were tested to confirm that vestibular cues were central to the spatial coding of both eye and hand movements during VCP. LD subjects performed significantly worse than normal subjects. These results demonstrate that adaptive change in VOR can lead to alterations in manual target localization.
Psychological and physiological models for describing biological processes of controlled eye and forearm movement during target tracking operations
Stabilization of the eyes and head during body movements is important for maintaining balance and keeping the images of objects stationary on our retinas. Impairment of this ability can lead to disorientation and reduced performance in sensorimotor tasks such as piloting of spacecraft. In the absence of a normal earth gravity field, the dynamics of head stabilization, and the interpretation of vestibular signals that sense gravity and linear acceleration, are subject to change. Transitions between different gravitoinertial force environments - as during different phases of space flight - provide an extreme test of the adaptive mechanisms that maintain these reflexive abilities. It is vitally important to determine human adaptive capabilities in such a circumstance, so that we can know to what extent the sensorimotor skills acquired in one gravity environment will transfer to others. Our work lays the foundation for understanding these capabilities, and for determining how we can aid the processes of adaptation and readaptation. An integrated set of experiments addresses this issue. We use the general approach of adapting some type of reflexive eye movement (saccades, the angular vestibulo-ocular reflex (AVOR), the linear vestibulo-ocular reflex (LVOR)), or the vestibulo-collic reflex (VCR), to a particular change in gain or phase in one condition of gravitoiner-tial force, and adapting to a different gain or phase (or asking for no change) in a second gravitoinertial force condition, and then seeing if the gravitoinertial force itself - the context cue - can recall the previously learned adapted responses. The majority of the experiments in the laboratory use the direction of vertical gaze or the direction of gravity (head tilt) as the context cue. This allows us to study context-specificity in a ground-based setting. One set of experiments, to be performed in parabolic flight, specifically uses the magnitude of gravitoinertial force as a context cue. This is a much better analog of the situation encountered in space flight. Various experiments investigate the behavioral properties, neurophysiological basis, and anatomical substrate of context-specific learning mechanisms. We use otolith (gravity) signals as the contextual cue for switching between adapted states of the saccadic system, the angular and linear vestibulo-ocular reflexes, and the VCR. (By LVOR we mean the oculomotor response - horizontal, vertical, and torsional - to linear translation of the head and body.) We are studying the effect of context on adaptation of saccade gain, phase and gain of the AVOR and LVOR, on ocular counterrolling (OCR) in response to static head tilt, and on head/neck reflexes (VCR) in response to rotation in different orientations. Such research is particularly germane to potential problems of postural and oculomotor control upon exposure to different gravitational environments.
In sequentially-scanned liquid-crystal displays the chromatic components of color pixels are distributed in time. For such displays eye, head, display, and image-object movements can cause the individual color elements to be visible. We analyze conditions (scan designs, types of eye movement) likely to produce color breakup.
Introduction. This joint European Space Agency/NASA pre- and post-flight study investigates the influence of exposure to microgravity on the subjective straight ahead (SSA) in crewmembers returning from long-duration expeditions to the International Space Station (ISS). The SSA is a measure of the internal representation of body orientation and to be influenced by stimulation of sensory systems involved in postural control. The use of a vibrotactile sensory aid to correct the representation of body tilted relative to gravity is also tested as a countermeasure. This study addresses the sensorimotor research gap to "determine the changes in sensorimotor function over the course of a mission and during recovery after landing." Research Plans. The ISS study will involve eight crewmembers who will participate in three pre-flight sessions (between 120 and 60 days before launch) and then three post-flight sessions on R plus 0/1 day, R plus 4 days, and R plus 8 days. Sixteen control subjects were also tested during three sessions to evaluate the effects of repeated testing and to establish normative values. The experimental protocol includes measurements of gaze and arm movements during the following tasks: (1) Near & Far Fixation: The subject is asked to look at actual targets in the true straight-ahead direction or to imagine these targets in the dark. Targets are located at near distance (arm's length) and far distance (beyond 2 meters). This task is successively performed with the subject's body aligned with the gravitational vertical, and with the subject's body tilted in pitch relative to the gravitational vertical using a tilt chair. Measures are then compared with and without a vibrotactile sensory aid that indicates how far one has tilted relative to the vertical; (2) Eye and Arm Movements: The subject is asked to look and point in the SSA direction in darkness and then make horizontal and vertical eye or arm movements, relative to Earth coordinates (allocentric) and to the subject's head/body reference (egocentric). This task is successively performed with the subject's body aligned with the gravitational vertical, and with subject's body tilted in roll using a tilt chair; (3) Linear Vestibulo-Ocular Reflex: The subject is asked to fixate actual visual targets at near and far distances in the true straight-ahead direction, and to evaluate the distance of these targets. The subject is asked to continue fixating the same imagined targets in darkness while he/she is passively accelerated up and down on a spring-loaded vertical linear accelerator. Results. In the control subject population, the perceived tilt angles, translations, and distances were remarkably close to the actual values. The pointing tasks indicated that the orientation of arm saccades was influenced by both the gravitational vertical and the body idiotropic vector. Repeating the testing did not reveal any significant changes. Preliminary results obtained in three crewmembers before and after flight will also be presented. Applications. A change in an individual's egocentric reference might have negative consequences on evaluating the direction of an approaching object or on the accuracy of reaching movements or locomotion. Consequently, investigating how microgravity affects the target location will have theoretical, operational, and even clinical implications for future space exploration missions. The use of vibrotactile feedback as a sensorimotor countermeasure is applicable to balance therapy applications for patients with vestibular loss and the elderly to mitigate risks due to loss of spatial orientation.
The reafference model has frequently been used to explain spatial constancy during eye and head movements. We have found that its basic concepts also form part of the information processing necessary for the control and recalibration of reaching movements. Reaching was studied in a novel force environment--a rotating room that creates centripetal forces of the type that could someday substitute for gravity in space flight, and Coriolis forces which are side effects of rotation. We found that inertial, noncontacting Coriolis forces deviate the path and endpoint of reaching movements, a finding that shows the inadequacy of equilibrium position models of movement control. Repeated movements in the rotating room quickly lead to normal movement patterns and to a failure to perceive the perturbing forces. The first movements made after rotation stops, without Coriolis forces present, show mirror-image deviations and evoke perception of a perturbing force even though none is present. These patterns of sensorimotor control and adaptation can largely be explained on the basis of comparisons of efference copy, reafferent muscle spindle, and cutaneous mechanoreceptor signals. We also describe experiments on human locomotion using an apparatus similar to that which Mittelstaedt used to study the optomotor response of the Eristalis fly. These results show that the reafference principle relates as well to the perception of the forces acting on and exerted by the body during voluntary locomotion.