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Psychological and physiological study of auditory masking
Two part-task preflight adaptation trainers (PATs) are being developed at the NASA Johnson Space Center to preadapt astronauts to novel sensory stimulus conditions similar to those present in microgravity to facilitate adaptation to microgravity and readaptation to Earth. This activity is a major component of a general effort to develop countermeasures aimed at minimizing sensory and sensorimotor disturbances and Space Motion Sickness (SMS) associated with adaptation to microgravity and readaptation to Earth. Design principles for the development of the two trainers are discussed, along with a detailed description of both devices. In addition, a summary of four ground-based investigations using one of the trainers to determine the extent to which various novel sensory stimulus conditions produce changes in compensatory eye movement responses, postural equilibrium, motion sickness symptoms, and electrogastric responses are presented. Finally, a brief description of the general concept of dual-adopted states that underly the development of the PATs, and ongoing and future operational and basic research activities are presented.
A method which uses an instrument that detects the response of a long bone to a vibratory stimulus to quantify mechanical properties non-invasively was revised and validated for use in the tibia. Stored data from healthy men was reanalyzed and compared with values from non-human primates. The analysis uses the relationship K(sub b) = 48 EI/L(sup 3) where K(sub b) is the lateral stiffness of a beam with force applied midspan, E is the elastic modulus, I the geometric moment of inertia and L, the limb length. Values for stiffness (EI, Nm(sup2)), the Euler buckling load (P(sub cr) = EI (pi/L)(sup 2)), and bone sufficiency (S) which represents the axial load the bone can support, adjusted to BW (S=P(sub cr)/BW) were obtained. The interest precision of the method in relaxed men, 5.8%, and in sedated male monkeys, 4.3%, was based on repeated measures in the same subjects at 1 month intervals. The R tibias of 40 men, aged 38.6 +/- 7.3 yrs with BW 78.9 +/- 7.9 kg, showed average (+/- SD) L to be 35 +/- 2 cm, EI 222 +/- 71 Nm(sup 2), P(sub cr) 18.1 +/- 4.9 kN, and S 23.4 +/- 5.7 N. The R tibias of 24 Rhesus monkeys ranging in age from 2-12 years, BW 4.9 +/- 3 kg, showed L to be 14.7 +/- 1.9 cm, EI 6.0 +/- 4.8 Nm(sup 2), P(sub cr) 2.51 +/- 1.2 kN and S 57.3 N. These measurements indicate that the tibia of a terrestrial non-human primate, M. mulatta, has higher load carrying capacity for the level of body weights in the species than the human bone.
To study task switching when subjects need not inhibit inappropriate responses, we examine tasks with non-overlapping stimulus sets (e.g. color patches and uncolored letters). A new three-task paradigm permits the dissociation of several otherwise confounded variables. We find that performance declines monotonically with increasing time since last performance of a task. Adjusting for the effects of this factor permits a fresh assessment of the relationship between task expectancy and recency (Ruthruff, Remington & Johnston, 1996).
Our learning model has memory templates representing the target-plus-noise and noise-alone stimulus sets. The best correlating template determines the response. The correlations and the feedback participate in the additive template updating rule. The model can predict the relative thresholds for detection in random, fixed and twin noise.
The development of the leaf-sheath pulvinus of oat (Avena sativa L. cv. Victory) was studied in terms of its competency to respond to gravistimulation. Stages of onset of competency, maximum competency and loss of competency were identified, using the length of the supertending internode as a developmental marker. During the early phases in the onset of competency, the latency period between stimulus and graviresponse decreased and the steady state response rate increased significantly. When fully competent, the latency period remained constant as the plant continued to develop, suggesting that the latency period is relatively insensitive to quantitative changes (e.g., in carbohydrate or nutrient availability) at the cell level within the plant. In contrast, the response rate was found to increase with plant development, indicating that graviresponse rate is more strongly influenced by quantitative cellular changes. The total possible graviresponse of a single oat pulvinus was confirmed to be significantly less than the original presentation angle. This was shown to not result from a loss of competency, since the graviresponse could be reinitiated by increasing the presentation angle. As a result of the low overall graviresponse of individual pulvini, two or more pulvini are required to bring the plant apex to the vertical. This was determined to occur though the sequential, rather than simultaneous, action of successive pulvini, since a given pulvinus lost competency to gravirespond shortly after the next pulvinus became fully competent.
Mechanisms for the induction of motor learning in the vestibulo-ocular reflex (VOR) were evaluated by recording the patterns of neural activity elicited in the cerebellum by a range of stimuli that induce learning. Patterns of climbing-fiber, vestibular, and Purkinje cell simple-spike signals were examined during sinusoidal head movement paired with visual image movement at stimulus frequencies from 0.5 to 10 Hz. A comparison of simple-spike and vestibular signals contained the information required to guide learning only at low stimulus frequencies, and a comparison of climbing-fiber and simple-spike signals contained the information required to guide learning only at high stimulus frequencies. Learning could be guided by comparison of climbing-fiber and vestibular signals at all stimulus frequencies tested, but only if climbing fiber responses were compared with the vestibular signals present 100 msec earlier. Computational analysis demonstrated that this conclusion is valid even if there is a broad range of vestibular signals at the site of plasticity. Simulations also indicated that the comparison of vestibular and climbing-fiber signals across the 100 msec delay must be implemented by a subcellular "eligibility" trace rather than by neural circuits that delay the vestibular inputs to the site of plasticity. The results suggest two alternative accounts of learning in the VOR. Either there are multiple mechanisms of learning that use different combinations of neural signals to drive plasticity, or there is a single mechanism tuned to climbing-fiber activity that follows activity in vestibular pathways by approximately 100 msec.
In the present study, the sensitivity to sinusoidal linear accelerations in the plane of the utricular macula was tested in afferents. The head orientation relative to the translation axis was varied in order to determine the head position that elicited the maximal and minimal responses for each afferent. The response gain and phase values obtained to 0.5-Hz and 2-Hz linear acceleration stimuli were then plotted as a function of head orientation and a modified cosine function was fit to the data. From the best-fit cosine function, the predicted head orientations that would produce the maximal and minimal response gains were estimated. The estimated maximum response gains to linear acceleration in the utricular plane for the afferents varied between 75 and 1420 spikes s-1 g-1. The mean maximal gains for all afferents to 0.5-Hz and 2-Hz sinusoidal linear acceleration stimuli were 282 and 367 spikes s-1 g-1, respectively. The minimal response gains were essentially zero for most units. The response phases always led linear acceleration and remained constant for each afferent, regardless of head orientation. These response characteristics indicate that otolith afferents are cosine tuned and behave as one-dimensional linear accelerometers. The directions of maximal sensitivity to linear acceleration for the afferents varied throughout the plane of the utricle; however, most vectors were directed out of the opposite ear near the interaural axis. The response dynamics of the afferents were tested using stimulus frequencies ranging between 0.25 Hz and 10 Hz (0.1 g peak acceleration). Across stimulus frequencies, most afferents had increasing gains and constant phase values. These dynamic properties for individual afferents were fit with a simple transfer function that included three parameters: a mechanical time constant, a gain constant, and a fractional order distributed adaptation operator.
The present study demonstrates that the general capability of skeletal muscle to maintain its proteins decreases rapidly in response to space flight. The present findings suggest further that the magnitude of enzymatic and cell volumes changes in response to space flight depend on several factors including the muscle and its fiber type composition. It appears that in order to associate physiological relevance to the observed enzymatic changes, cell volume should be considered also. Although it remains unclear as to the stimulus, or lack of stimulus, that triggers the rapid changes in muscle proteins in response to space flight, ground-based models of muscle atrophy suggest that the reduction in mechanical loading of muscle may be more important than the total amount of activation over a 24-hr period.
Meaningful comparisons between single stimulus EEG records and average evoked potential
Cortical gamma band oscillations have been recorded in sensory cortices of cats and monkeys, and are thought to aid in perceptual binding. Gamma activity has also been recorded in the rat hippocampus and entorhinal cortex, where it has been shown, that field gamma power is modulated at theta frequency. Since the power of gamma activity in the sensory cortices is not constant (gamma-bursts). we decided to examine the relationship between gamma power and the phase of low frequency oscillation in the auditory cortex of the awake macaque. Macaque monkeys were surgically prepared for chronic awake electrophysiological recording. During the time of the experiments. linear array multielectrodes were inserted in area AI to obtain laminar current source density (CSD) and multiunit activity profiles. Instantaneous theta and gamma power and phase was extracted by applying the Morlet wavelet transformation to the CSD. Gamma power was averaged for every 1 degree of low frequency oscillations to calculate power-phase relation. Both gamma and theta-delta power are largest in the supragranular layers. Power modulation of gamma activity is phase locked to spontaneous, as well as stimulus-related local theta and delta field oscillations. Our analysis also revealed that the power of theta oscillations is always largest at a certain phase of delta oscillation. Auditory stimuli produce evoked responses in the theta band (Le., there is pre- to post-stimulus addition of theta power), but there is also indication that stimuli may cause partial phase re-setting of spontaneous delta (and thus also theta and gamma) oscillations. We also show that spontaneous oscillations might play a role in the processing of incoming sensory signals by 'preparing' the cortex.
Head movements in the sagittal pitch plane typically involve off-axis rotation requiring both vertical and horizontal vergence ocular reflexes to compensate for angular and translational motion relative to visual targets of interest. The purpose of this study was to compare passive pitch VOR responses during rotation about an Earth-vertical axis (canal only cues) with off-axis rotation (canal and otolith cues). Methods. Eleven human subjects were oscillated sinusoidally at 0.13, 0.3 and 0.56 Hz while lying left-side down with the interaural axis either aligned with the axis of rotation or offset by 50 cm. In a second set of measurements, twelve subjects were also tested during sinusoidally varying centrifugation over the same frequency range. The modulation of vertical and horizontal vergence ocular responses was measured with a binocular videography system. Results. Off-axis pitch rotation enhanced the vertical VOR at lower frequencies and enhanced the vergence VOR at higher frequencies. During sinusoidally varying centrifugation, the opposite trend was observed for vergence, with both vertical and vergence vestibulo-ocular reflexes being suppressed at the highest frequency. Discussion. These differential effects of off-axis rotation over the 0.13 to 0.56 Hz range are consistent with the hypothesis that otolith-ocular reflexes are segregated in part on the basis of stimulus frequency. At the lower frequencies, tilt otolith-ocular responses compensate for declining canal input. At higher frequencies, translational otolith-ocular reflexes compensate for declining visual contributions to the kinematic demands required for fixating near targets.
Non-random oscillation in electroencephalographic alpha blocking response
For monocular viewing, the fluctuations in accommodative lens power in the frequency range from 0.5 to 3 Hz were found to be considerably greater than those in accommodative vergence movements of the covered eye. Considering the close synkinesis between these motor responses for step changes or slow variations in accommodative stimulus, this finding is unexpected. This apparent lack of synkinesis is found to result mainly from the fact that the decrease in small-signal linear gain with increasing frequency is more rapid in the case of the accommodative vergence system than in the case of the accommodation system, rather than from some nonlinear phenomenon.
When one tone burst (the conditioner) preceeds another (the target) by 100 ms, target loudness is enhanced if the conditioner is more intense and decreased if it is less intense. We show here that similar loudness enhancements and decrements occur when the conditioner follows the target. In all instances, monaural loudness enhancements (in which the conditioner and target are delivered to the same ear) are greater than the dichotic enhancements (in which the conditioner is presented contralaterally), but the decrements, which are smaller than the enhancements, are similar in magnitude. Loudness enhancements and decrements are similar to sequential loudness effects and central tendency effects; the major difference is the relatively very large increases in loudness obtainable in loudness enhancement experiments. We outline a mechanism to account for these loudness phenomena and suggest that this mechanism is responsible for similar perceptual effects that occur in other stimulus dimensions and modalities.
A large number of roll rate stimuli, covering rates from zero to plus or minus 25 deg/sec, were presented to subjects in random order at 2 sec intervals. Subjects were to make estimates of magnitude of perceived roll rate stimuli presented on either a central display, on displays in the peripheral ield of vision, or on all displays simultaneously. Response was by way of a digital keyboard device, stimulus exposition times were varied. The present experiment differs from earlier perception tasks by the same authors in that mean rate perception error (and standard deviation) was obtained as a function of rate stimulus magnitude, whereas the earlier experiments only yielded mean absolute error magnitude. Moreover, in the present experiment, all stimulus rates had an equal probability of occurrence, whereas the earlier tests featured a Gaussian stimulus probability density function. Results yield a ood illustration of the nonlinear functions relating rate presented to rate perceived by human observers or operators.
We have previously proposed a computational neural-network model by which the complex patterns of retinal image motion generated during locomotion (optic flow) can be processed by specialized detectors acting as templates for specific instances of self-motion. The detectors in this template model respond to global optic flow by sampling image motion over a large portion of the visual field through networks of local motion sensors with properties similar to neurons found in the middle temporal (MT) area of primate extrastriate visual cortex. The model detectors were designed to extract self-translation (heading), self-rotation, as well as the scene layout (relative distances) ahead of a moving observer, and are arranged in cortical-like heading maps to perform this function. Heading estimation from optic flow has been postulated by some to be implemented within the medial superior temporal (MST) area. Others have questioned whether MST neurons can fulfill this role because some of their receptive-field properties appear inconsistent with a role in heading estimation. To resolve this issue, we systematically compared MST single-unit responses with the outputs of model detectors under matched stimulus conditions. We found that the basic physiological properties of MST neurons can be explained by the template model. We conclude that MST neurons are well suited to support heading estimation and that the template model provides an explicit set of testable hypotheses which can guide future exploration of MST and adjacent areas within the primate superior temporal sulcus.
Tests of otolith function were performed pre-flight and post-flight on the science crew of the first Spacelab Mission with a rail-mounted linear acceleration sled. Four tests were performed using horizontal lateral (y-axis) acceleration: perception of linear motion, a closed loop nulling task, dynamic ocular torsion, and lateral eye deviations. The motion perception test measured the time to detect the onset and direction of near threshold accelerations. Post-flight measures of threshold and velocity constant obtained during the days immediately following the mission showed no consistent pattern of change among the four crewmen compared to their pre-flight baseline other than an increased variability of response. In the closed loop nulling task, crewmen controlled the motion of the sled and attempted to null a computer-generated random disturbance motion. When performed in the light, no difference in ability was noted between pre-flight and post-flight. In the dark, however, two of the four crewmen exhibited somewhat enhanced performance post-flight. Dynamic ocular torsion was measured in response to sinusoidal lateral acceleration which produces a gravitionertial stimulus equivalent to lateral head tilt without rotational movement of the head. Results available for two crewmen suggest a decreased amplitude of sinusoidal ocular torsion when measured on the day of landing (R+0) and an increasing amplitude when measured during the week following the mission.