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At least 19 records

Spatiotemporal processing of linear acceleration: primary afferent and central vestibular neuron responses

Spatiotemporal convergence and two-dimensional (2-D) neural tuning have been proposed as a major neural mechanism in the signal processing of linear acceleration. To examine this hypothesis, we studied the firing properties of primary otolith afferents and central otolith neurons that respond exclusively to horizontal linear accelerations of the head (0.16-10 Hz) in alert rhesus monkeys. Unlike primary afferents, the majority of central otolith neurons exhibited 2-D spatial tuning to linear acceleration. As a result, central otolith dynamics vary as a function of movement direction. During movement along the maximum sensitivity direction, the dynamics of all central otolith neurons differed significantly from those observed for the primary afferent population. Specifically at low frequencies (</=0.5 Hz), the firing rate of the majority of central otolith neurons peaked in phase with linear velocity, in contrast to primary afferents that peaked in phase with linear acceleration. At least three different groups of central response dynamics were described according to the properties observed for motion along the maximum sensitivity direction. "High-pass" neurons exhibited increasing gains and phase values as a function of frequency. "Flat" neurons were characterized by relatively flat gains and constant phase lags (approximately 20-55 degrees ). A few neurons ("low-pass") were characterized by decreasing gain and phase as a function of frequency. The response dynamics of central otolith neurons suggest that the approximately 90 degrees phase lags observed at low frequencies are not the result of a neural integration but rather the effect of nonminimum phase behavior, which could arise at least partly through spatiotemporal convergence. Neither afferent nor central otolith neurons discriminated between gravitational and inertial components of linear acceleration. Thus response sensitivity was indistinguishable during 0.5-Hz pitch oscillations and fore-aft movements. The fact that otolith-only central neurons with "high-pass" filter properties exhibit semicircular canal-like dynamics during head tilts might have important consequences for the conclusions of previous studies of sensory convergence and sensorimotor transformations in central vestibular neurons.

NASA Discipline Neuroscience

Posture, locomotion, spatial orientation, and motion sickness as a function of space flight

This article summarizes a variety of newly published findings obtained by the Neuroscience Laboratory, Johnson Space Center, and attempts to place this work within a historical framework of previous results on posture, locomotion, motion sickness, and perceptual responses that have been observed in conjunction with space flight. In this context, we have taken the view that correct transduction and integration of signals from all sensory systems is essential to maintaining stable vision, postural and locomotor control, and eye-hand coordination as components of spatial orientation. The plasticity of the human central nervous system allows individuals to adapt to altered stimulus conditions encountered in a microgravity environment. However, until some level of adaptation is achieved, astronauts and cosmonauts often experience space motion sickness, disturbances in motion control and eye-hand coordination, unstable vision, and illusory motion of the self, the visual scene, or both. Many of the same types of disturbances encountered in space flight reappear immediately after crew members return to earth. The magnitude of these neurosensory, sensory-motor and perceptual disturbances, and the time needed to recover from them, tend to vary as a function of mission duration and the space travelers prior experience with the stimulus rearrangement of space flight. To adequately chart the development of neurosensory changes associated with space flight, we recommend development of enhanced eye movement systems and body position measurement. We also advocate the use of a human small radius centrifuge as both a research tool and as a means of providing on-orbit countermeasures that will lessen the impact of living for long periods of time with out exposure to altering gravito-inertial forces. Copyright 1998 Elsevier Science B.V.

Flight Experiment

Perception of tilt (somatogravic illusion) in response to sustained linear acceleration during space flight

During the 1998 Neurolab mission (STS-90), four astronauts were exposed to interaural and head vertical (dorsoventral) linear accelerations of 0.5 g and 1 g during constant velocity rotation on a centrifuge, both on Earth and during orbital space flight. Subjects were oriented either left-ear-out or right-ear-out (Gy centrifugation), or lay supine along the centrifuge arm with their head off-axis (Gz centrifugation). Pre-flight centrifugation, producing linear accelerations of 0.5 g and 1 g along the Gy (interaural) axis, induced illusions of roll-tilt of 20 degrees and 34 degrees for gravito-inertial acceleration (GIA) vector tilts of 27 degrees and 45 degrees , respectively. Pre-flight 0.5 g and 1 g Gz (head dorsoventral) centrifugation generated perceptions of backward pitch of 5 degrees and 15 degrees , respectively. In the absence of gravity during space flight, the same centrifugation generated a GIA that was equivalent to the centripetal acceleration and aligned with the Gy or Gz axes. Perception of tilt was underestimated relative to this new GIA orientation during early in-flight Gy centrifugation, but was close to the GIA after 16 days in orbit, when subjects reported that they felt as if they were 'lying on side'. During the course of the mission, inflight roll-tilt perception during Gy centrifugation increased from 45 degrees to 83 degrees at 1 g and from 42 degrees to 48 degrees at 0.5 g. Subjects felt 'upside-down' during in-flight Gz centrifugation from the first in-flight test session, which reflected the new GIA orientation along the head dorsoventral axis. The different levels of in-flight tilt perception during 0.5 g and 1 g Gy centrifugation suggests that other non-vestibular inputs, including an internal estimate of the body vertical and somatic sensation, were utilized in generating tilt perception. Interpretation of data by a weighted sum of body vertical and somatic vectors, with an estimate of the GIA from the otoliths, suggests that perception weights the sense of the body vertical more heavily early in-flight, that this weighting falls during adaptation to microgravity, and that the decreased reliance on the body vertical persists early post-flight, generating an exaggerated sense of tilt. Since graviceptors respond to linear acceleration and not to head tilt in orbit, it has been proposed that adaptation to weightlessness entails reinterpretation of otolith activity, causing tilt to be perceived as translation. Since linear acceleration during in-flight centrifugation was always perceived as tilt, not translation, the findings do not support this hypothesis.

STS-90 Shuttle Project

Defining the cortical visual systems: "what", "where", and "how"

The visual system historically has been defined as consisting of at least two broad subsystems subserving object and spatial vision. These visual processing streams have been organized both structurally as two distinct pathways in the brain, and functionally for the types of tasks that they mediate. The classic definition by Ungerleider and Mishkin labeled a ventral "what" stream to process object information and a dorsal "where" stream to process spatial information. More recently, Goodale and Milner redefined the two visual systems with a focus on the different ways in which visual information is transformed for different goals. They relabeled the dorsal stream as a "how" system for transforming visual information using an egocentric frame of reference in preparation for direct action. This paper reviews recent research from psychophysics, neurophysiology, neuropsychology and neuroimaging to define the roles of the ventral and dorsal visual processing streams. We discuss a possible solution that allows for both "where" and "how" systems that are functionally and structurally organized within the posterior parietal lobe.

NASA Discipline Space Human Factors

Expansion of direction space around the cardinal axes revealed by smooth pursuit eye movements

It is well established that perceptual direction discrimination shows an oblique effect; thresholds are higher for motion along diagonal directions than for motion along cardinal directions. Here, we compare simultaneous direction judgments and pursuit responses for the same motion stimuli and find that both pursuit and perceptual thresholds show similar anisotropies. The pursuit oblique effect is robust under a wide range of experimental manipulations, being largely resistant to changes in trajectory (radial versus tangential motion), speed (10 versus 25 deg/s), directional uncertainty (blocked versus randomly interleaved), and cognitive state (tracking alone versus concurrent tracking and perceptual tasks). Our data show that the pursuit oblique effect is caused by an effective expansion of direction space surrounding the cardinal directions and the requisite compression of space for other directions. This expansion suggests that the directions around the cardinal directions are in some way overrepresented in the visual cortical pathways that drive both smooth pursuit and perception.

Visual Pathways/physiology

The relation of motion sickness to the spatial-temporal properties of velocity storage

Tilting the head in roll to or from the upright while rotating at a constant velocity (roll while rotating, RWR) alters the position of the semicircular canals relative to the axis of rotation. This produces vertical and horizontal nystagmus, disorientation, vertigo, and nausea. With recurrent exposure, subjects habituate and can make more head movements before experiencing overpowering motion sickness. We questioned whether promethazine lessened the vertigo or delayed the habituation, whether habituation of the vertigo was related to the central vestibular time constant, i.e., to the time constant of velocity storage, and whether the severity of the motion sickness was related to deviation of the axis of eye velocity from gravity. Sixteen subjects received promethazine and placebo in a double-blind, crossover study in two consecutive 4-day test series 1 month apart, termed series I and II. Horizontal and vertical eye movements were recorded with video-oculography while subjects performed roll head movements of approx. 45 degrees over 2 s to and from the upright position while being rotated at 138 degrees /s around a vertical axis. Motion sickness was scaled from 1 (no sickness) to an endpoint of 20, at which time the subject was too sick to continue or was about to vomit. Habituation was determined by the number of head movements that subjects made before reaching the maximum motion sickness score of 20. Head movements increased steadily in each session with repeated testing, and there was no difference between the number of head movements made by the promethazine and placebo groups. Horizontal and vertical angular vestibulo-ocular reflex (aVOR) time constants declined in each test, with the declines being closely correlated to the increase in the number of head movements. The strength of vertiginous sensation was associated with the amount of deviation of the axis of eye velocity from gravity; the larger the deviation of the eye velocity axis from gravity, the more severe the motion sickness. Thus, promethazine neither reduced the nausea associated with RWR, nor retarded or hastened habituation. The inverse relationship between the aVOR time constants and number of head movements to motion sickness, and the association of the severity of motion sickness with the extent, strength, and time of deviation of eye velocity from gravity supports the postulate that the spatiotemporal properties of velocity storage, which are processed between the nodulus and uvula of the vestibulocerebellum and the vestibular nuclei, are likely to represent the source of the conflict responsible for producing motion sickness.

NASA Discipline Neuroscience

Vestibular adaptation to space in monkeys

Otolith-induced eye movements of rhesus monkeys were studied before and after the 1989 COSMOS 2044 and the 1992 to 1993 COSMOS 2229 flights. Two animals flew in each mission for approximately 2 weeks. After flight, spatial orientation of the angular vestibulo-ocular reflex was altered. In one animal the time constant of postrotatory nystagmus, which had been shortened by head tilts with regard to gravity before flight, was unaffected by the same head tilts after flight. In another animal, eye velocity, which tended to align with a gravitational axis before flight, moved toward a body axis after flight. This shift of orientation disappeared by 7 days after landing. After flight, the magnitude of compensatory ocular counter-rolling was reduced by about 70% in both dynamic and static tilts. Modulation in vergence in response to naso-occipital linear acceleration during off-vertical axis rotation was reduced by more than 50%. These changes persisted for 11 days after recovery. An up and down asymmetry of vertical nystagmus was diminished for 7 days. Gains of the semicircular canal-induced horizontal and vertical angular vestibulo-ocular reflexes were unaffected in both flights, but the gain of the roll angular vestibulo-ocular reflex was decreased. These data indicate that there are short- and long-term changes in otolith-induced eye movements after adaptation to microgravity. These experiments also demonstrate the unique value of the monkey as a model for studying effects of vestibular adaptation in space. Eye movements can be measured in three dimensions in response to controlled vestibular and visual stimulation, and the results are directly applicable to human beings. Studies in monkeys to determine how otolith afferent input and central processing is altered by adaptation to microgravity should be an essential component of future space-related research.

Cosmos 2229 Project

Primate translational vestibuloocular reflexes. I. High-frequency dynamics and three-dimensional properties during lateral motion

The dynamics and three-dimensional (3-D) properties of the primate translational vestibuloocular reflex (trVOR) for high-frequency (4-12 Hz, +/-0.3-0.4 g) lateral motion were investigated during near-target viewing at center and eccentric targets. Horizontal response gains increased with frequency and depended on target eccentricity. The larger the horizontal and vertical target eccentricity, the steeper the dependence of horizontal response gain on frequency. In addition to horizontal eye movements, robust torsional response components also were present at all frequencies. During center-target fixation, torsional response phase was opposite (anticompensatory) to that expected for an "apparent" tilt response. Instead torsional response components depended systematically on vertical-target eccentricity, increasing in amplitude when looking down and reversing phase when looking up. As a result the trVOR eye velocity vector systematically tilted away from a purely horizontal direction, through an angle that increased with vertical eccentricity with a slope of approximately 0.7. This systematic dependence of torsional eye velocity tilt on vertical eye position suggests that the trVOR might follow the 3-D kinematic requirements that have been shown to govern visually guided eye movements and near-target fixation.

NASA Discipline Neuroscience

Neural readaptation to Earth's gravity following return from space

The consequence of exposure to microgravity on the otolith organs was studied by recording the responses of vestibular nerve afferents supplying the utricular otolith organ to inertial accelerations in four toadfish, Opsanus tau, sequentially for 5 days following two National Aeronautics and Space Administration shuttle orbital flights. Within the first day postflight, the magnitude of response to an applied translation was on average three times greater than for controls. The reduced gravitational acceleration in orbit apparently resulted in an upregulation of the sensitivity of utricular afferents. By 30 h postflight, responses were statistically similar to control. The time course of return to normal afferent sensitivity parallels the reported decrease in vestibular disorientation in astronauts following return from space.

manned

Rapid motor learning in the translational vestibulo-ocular reflex

Motor learning was induced in the translational vestibulo-ocular reflex (TVOR) when monkeys were repeatedly subjected to a brief (0.5 sec) head translation while they tried to maintain binocular fixation on a visual target for juice rewards. If the target was world-fixed, the initial eye speed of the TVOR gradually increased; if the target was head-fixed, the initial eye speed of the TVOR gradually decreased. The rate of learning acquisition was very rapid, with a time constant of approximately 100 trials, which was equivalent to <1 min of accumulated stimulation. These learned changes were consolidated over >or=1 d without any reinforcement, indicating induction of long-term synaptic plasticity. Although the learning generalized to targets with different viewing distances and to head translations with different accelerations, it was highly specific for the particular combination of head motion and evoked eye movement associated with the training. For example, it was specific to the modality of the stimulus (translation vs rotation) and the direction of the evoked eye movement in the training. Furthermore, when one eye was aligned with the heading direction so that it remained motionless during training, learning was not expressed in this eye, but only in the other nonaligned eye. These specificities show that the learning sites are neither in the sensory nor the motor limb of the reflex but in the sensory-motor transformation stage of the reflex. The dependence of the learning on both head motion and evoked eye movement suggests that Hebbian learning may be one of the underlying cellular mechanisms.

Non-NASA Center

Plant biology in space: proceedings of the International Workshop, Bad Honnef, Germany, June 24-27, 1996

Papers presented at the International Workshop on Plant Biology in Space include reviews, reports, and perspectives related to plant gravitational biology. Presentations focused on nine subject areas: gravitropism in unicellular plants, gravitropism in fungi, cell development, gravity perception in multicellular plants, gravity responses in multicellular plants, plant reproduction, evaluation of a clinostat for weightlessness simulation, biological life support systems, and future research.

NASA Discipline Plant Biology

Hippocampal place cell instability after lesions of the head direction cell network

The occurrence of cells that encode spatial location (place cells) or head direction (HD cells) in the rat limbic system suggests that these cell types are important for spatial navigation. We sought to determine whether place fields of hippocampal CA1 place cells would be altered in animals receiving lesions of brain areas containing HD cells. Rats received bilateral lesions of anterodorsal thalamic nuclei (ADN), postsubiculum (PoS), or sham lesions, before place cell recording. Although place cells from lesioned animals did not differ from controls on many place-field characteristics, such as place-field size and infield firing rate, the signal was significantly degraded with respect to measures of outfield firing rate, spatial coherence, and information content. Surprisingly, place cells from lesioned animals were more likely modulated by the directional heading of the animal. Rotation of the landmark cue showed that place fields from PoS-lesioned animals were not controlled by the cue and shifted unpredictably between sessions. Although fields from ADN-lesioned animals tended to have less landmark control than fields from control animals, this impairment was mild compared with cells recorded from PoS-lesioned animals. Removal of the prominent visual cue also led to instability of place-field representations in PoS-lesioned, but not ADN-lesioned, animals. Together, these findings suggest that an intact HD system is not necessary for the maintenance of place fields, but lesions of brain areas that convey the HD signal can degrade this signal, and lesions of the PoS might lead to perceptual or mnemonic deficits, leading to place-field instability between sessions.

NASA Program Biomedical Research and Countermeasur

Characteristics of covert and overt visual orienting: Evidence from attentional and oculomotor capture

Five visual search experiments found oculomotor and attentional capture consistent with predictions of contingent orienting, contrary to claims that oculomotor capture is purely stimulus driven. Separate saccade and attend-only conditions contained a color target appearing either singly, with an onset or color distractor, or both. In singleton mode, onsets produced oculomotor and attentional capture. In feature mode, capture was absent or greatly reduced, providing evidence for top-down modulation of both types of capture. Although attentional capture by color abstractors was present throughout, oculomotor capture by color occurred only when accompanied by transient change, providing evidence for a dissociation between oculomotor and attentional capture. Oculomotor and attentional capture appear to be mediated by top-down attentional control settings, but transient change may be necessary for oculomotor capture. ((c) 2003 APA, all rights reserved).

Clinical Trial

Recovery trajectories of vestibulopathic subjects after perturbations during locomotion

We compared the mediolateral (M/L) responses to perturbations during locomotion of vestibulopathic (VP) subjects to those of controls. Eight subjects with unilateral vestibular loss (100% Reduced Vestibular Response from the caloric test) resulting from surgery for vestibular schwannoma and 11 controls were selected for this study. Despite their known vestibulopathy, all VP subjects scored within the normal range on computerized dynamic posturography Sensory Organization Tests. During gait, subjects were given surface perturbations of the right support-phase foot in two possible directions (forward-right and backward-left) at two possible magnitudes (5 and 10 cm) that were randomly mixed with trials having no perturbations. M/L stability was quantified by estimating the length of the M/L moment arm between the support foot and the trunk, and the M/L accelerations of the sternum and the head. The VP group had greater changes (p < 0.05) in their moment arm responses compared to controls. The number of steps that it took for the moment arm oscillations to return to normal and the variability in the moment arms were greater for the VP group. Differences in the sternum and head accelerations between VP and control groups were not as consistent, but there was a trend toward greater response deviations in the VP group for all 4 perturbation types. Increased response magnitude and variability of the VP group is consistent with an increase in their sensory noise of vestibular inputs due to the surgical lesion. Another possibility is a reduced sensitivity to motion inputs. This perturbation approach may prove useful for characterizing subtle vestibulopathies and similar changes in the human orientation mechanism after exposure to microgravity.

NASA Discipline Neuroscience

Training, transfer, and retention of three-dimensional spatial memory in virtual environments

Human orientation requires one to remember and visualize spatial arrangements of landmarks from different perspectives. Astronauts have reported difficulties remembering relationships between environmental landmarks when imagined in arbitrary 3D orientations. The present study investigated the effects of strategy training on humans' 1) ability to infer their orientation from landmarks presented ahead and below, 2) performance when subsequently learning a different array, and 3) retention of configurational knowledge over time. On the first experiment day, 24 subjects were tested in a virtual cubic chamber in which a picture of an animal was drawn on each wall. Through trial-by-trial exposures, they had to memorize the spatial relationships among the six pictures around them and learn to predict the direction to a specific picture when facing any view direction, and in any roll orientation. Half of the subjects ("strategy group") were taught methods for remembering picture groupings, while the remainder received no such training ("control group"). After learning one picture array, the procedure was repeated in a second. Accuracy (% correct) and response time learning curves were measured. Performance for the second array and configurational memory of both arrays were also retested 1, 7, and 30 days later. Results showed that subjects "learned how to learn" this generic 3D spatial memory task regardless of their relative orientation to the environment, that ability and configurational knowledge was retained for at least a month, that figure rotation ability and field independence correlate with performance, and that teaching subjects specific strategies in advance significantly improves performance. Training astronauts to perform a similar generic 3D spatial memory task, and suggesting strategies in advance, may help them orient in three dimensions.

Non-NASA Center

Spatial updating in human parietal cortex

Single neurons in monkey parietal cortex update visual information in conjunction with eye movements. This remapping of stimulus representations is thought to contribute to spatial constancy. We hypothesized that a similar process occurs in human parietal cortex and that we could visualize it with functional MRI. We scanned subjects during a task that involved remapping of visual signals across hemifields. We observed an initial response in the hemisphere contralateral to the visual stimulus, followed by a remapped response in the hemisphere ipsilateral to the stimulus. We ruled out the possibility that this remapped response resulted from either eye movements or visual stimuli alone. Our results demonstrate that updating of visual information occurs in human parietal cortex.

NASA Discipline Neuroscience

Stimulus information contaminates summation tests of independent neural representations of features

Many models of visual processing assume that visual information is analyzed into separable and independent neural codes, or features. A common psychophysical test of independent features is known as a summation study, which measures performance in a detection, discrimination, or visual search task as the number of proposed features increases. Improvement in human performance with increasing number of available features is typically attributed to the summation, or combination, of information across independent neural coding of the features. In many instances, however, increasing the number of available features also increases the stimulus information in the task, as assessed by an optimal observer that does not include the independent neural codes. In a visual search task with spatial frequency and orientation as the component features, a particular set of stimuli were chosen so that all searches had equivalent stimulus information, regardless of the number of features. In this case, human performance did not improve with increasing number of features, implying that the improvement observed with additional features may be due to stimulus information and not the combination across independent features.

NASA Discipline Space Human Factors

Spatial learning and memory is preserved in rats after early development in a microgravity environment

This study evaluated the cognitive mapping abilities of rats that spent part of their early development in a microgravity environment. Litters of male and female Sprague-Dawley rat pups were launched into space aboard the National Aeronautics and Space Administration space shuttle Columbia on postnatal day 8 or 14 and remained in space for 16 days. These animals were designated as FLT groups. Two age-matched control groups remained on Earth: those in standard vivarium housing (VIV) and those in housing identical to that aboard the shuttle (AGC). On return to Earth, animals were tested in three different tasks that measure spatial learning ability, the Morris water maze (MWM), and a modified version of the radial arm maze (RAM). Animals were also tested in an open field apparatus to measure general activity and exploratory activity. Performance and search strategies were evaluated in each of these tasks using an automated tracking system. Despite the dramatic differences in early experience, there were remarkably few differences between the FLT groups and their Earth-bound controls in these tasks. FLT animals learned the MWM and RAM as quickly as did controls. Evaluation of search patterns suggested subtle differences in patterns of exploration and in the strategies used to solve the tasks during the first few days of testing, but these differences normalized rapidly. Together, these data suggest that development in an environment without gravity has minimal long-term impact on spatial learning and memory abilities. Any differences due to development in microgravity are quickly reversed after return to earth normal gravity.

NASA Discipline Neuroscience