Short-Term Visual Deprivation,Tactile Acuity,and Haptic Solid Shape Discrimination

Previous psychophysical studies have reported conflicting results concerning the effects of short-term visual deprivation upon tactile acuity. Some studies have found that 45 to 90 minutes of total light deprivation produce significant improvements in participants'' tactile acuity as measured with a grating orientation discrimination task. In contrast, a single 2011 study found no such improvement while attempting to replicate these earlier findings. A primary goal of the current experiment was to resolve this discrepancy in the literature by evaluating the effects of a 90-minute period of total light deprivation upon tactile grating orientation discrimination. We also evaluated the potential effect of short-term deprivation upon haptic 3-D shape discrimination using a set of naturally-shaped solid objects. According to previous research, short-term deprivation enhances performance in a tactile 2-D shape discrimination task – perhaps a similar improvement also occurs for haptic 3-D shape discrimination. The results of the current investigation demonstrate that not only does short-term visual deprivation not enhance tactile acuity, it additionally has no effect upon haptic 3-D shape discrimination. While visual deprivation had no effect in our study, there was a significant effect of experience and learning for the grating orientation task – the participants'' tactile acuity improved over time, independent of whether they had, or had not, experienced visual deprivation.     

A Tactile Automated Passive-Finger Stimulator (TAPS)

Although tactile spatial acuity tests are used in both neuroscience research and clinical assessment, few automated devices exist for delivering controlled spatially structured stimuli to the skin. Consequently, investigators often apply tactile stimuli manually. Manual stimulus application is time consuming, requires great care and concentration on the part of the investigator, and leaves many stimulus parameters uncontrolled. We describe here a computer-controlled tactile stimulus system, the Tactile Automated Passive-finger Stimulator (TAPS), that applies spatially structured stimuli to the skin, controlling for onset velocity, contact force, and contact duration. TAPS is a versatile, programmable system, capable of efficiently conducting a variety of psychophysical procedures. We describe the components of TAPS, and show how TAPS is used to administer a two-interval forced-choice tactile grating orientation test.Corresponding Author: Daniel Goldreich     

A linear model for symmetric receptive fields: implications for classification tests with flashed and moving images

The purpose of this study was to explore the effects of spatial and temporal properties on the expected responses of visual neurons that have linear receptive fields (RFs), particularly those having a mirror symmetric distribution of spatial subregions. Receptive fields that are symmetric in at least one spatial dimension occur in neurons of the retina, the lateral geniculate nucleus (LGN), and the visual cortex of mammals. Responses to flashing bars, moving bars, and moving edges were studied for different configurations of an analog RF model in which spatial and temporal aspects were varied independently. Responses of the model at intermediate stimulus speeds were found to agree with responses in the literature for X and Y units of the LGN and often for simple units of the visual cortex. In particular, having separated regions of response to light and dark edges, an identifying property of simple cells, was found to be a linear consequence of RF regions responding inversely to stimuli of opposite polarity. Model differences from responses of cortical complex units show that a linear model cannot mimic their responses, and imply that complex units employ major nonlinearities in coding image polarity (light vs dark), which signifies a nonlinearity in coding intensity. Because sudden flux changes inherent in flashing bars test mainly temporal RF properties, and slowly moving edges test mainly spatial properties, these two tests form a useful minimal set with which to describe and classify RFs. The usefulness of this set derives both from its sensitivity to spatial and temporal variables, and from the correlation between the linearity of a cell's processing of stimulus intensity and its RF classification.     

Short-term visual deprivation does not enhance passive tactile spatial acuity

An important unresolved question in sensory neuroscience is whether, and if so with what time course, tactile perception is enhanced by visual deprivation. In three experiments involving 158 normally sighted human participants, we assessed whether tactile spatial acuity improves with short-term visual deprivation over periods ranging from under 10 to over 110 minutes. We used an automated, precisely controlled two-interval forced-choice grating orientation task to assess each participant's ability to discern the orientation of square-wave gratings pressed against the stationary index finger pad of the dominant hand. A two-down one-up staircase (Experiment 1) or a Bayesian adaptive procedure (Experiments 2 and 3) was used to determine the groove width of the grating whose orientation each participant could reliably discriminate. The experiments consistently showed that tactile grating orientation discrimination does not improve with short-term visual deprivation. In fact, we found that tactile performance degraded slightly but significantly upon a brief period of visual deprivation (Experiment 1) and did not improve over periods of up to 110 minutes of deprivation (Experiments 2 and 3). The results additionally showed that grating orientation discrimination tends to improve upon repeated testing, and confirmed that women significantly outperform men on the grating orientation task. We conclude that, contrary to two recent reports but consistent with an earlier literature, passive tactile spatial acuity is not enhanced by short-term visual deprivation. Our findings have important theoretical and practical implications. On the theoretical side, the findings set limits on the time course over which neural mechanisms such as crossmodal plasticity may operate to drive sensory changes; on the practical side, the findings suggest that researchers who compare tactile acuity of blind and sighted participants should not blindfold the sighted participants.     

Shape Invariant Coding of Motion Direction in Somatosensory Cortex

Invariant representations of stimulus features are thought to play an important role in producing stable percepts of objects. In the present study, we assess the invariance of neural representations of tactile motion direction with respect to other stimulus properties. To this end, we record the responses evoked in individual neurons in somatosensory cortex of primates, including areas 3b, 1, and 2, by three types of motion stimuli, namely scanned bars and dot patterns, and random dot displays, presented to the fingertips of macaque monkeys. We identify a population of neurons in area 1 that is highly sensitive to the direction of stimulus motion and whose motion signals are invariant across stimulus types and conditions. The motion signals conveyed by individual neurons in area 1 can account for the ability of human observers to discriminate the direction of motion of these stimuli, as measured in paired psychophysical experiments. We conclude that area 1 contains a robust representation of motion and discuss similarities in the neural mechanisms of visual and tactile motion processing.     

Orientation-Cue Invariant Population Responses to Contrast-Modulated and Phase-Reversed Contour Stimuli in Macaque V1 and V2

Visual scenes can be readily decomposed into a variety of oriented components, the processing of which is vital for object segregation and recognition. In primate V1 and V2, most neurons have small spatio-temporal receptive fields responding selectively to oriented luminance contours (first order), while only a subgroup of neurons signal non-luminance defined contours (second order). So how is the orientation of second-order contours represented at the population level in macaque V1 and V2? Here we compared the population responses in macaque V1 and V2 to two types of second-order contour stimuli generated either by modulation of contrast or phase reversal with those to first-order contour stimuli. Using intrinsic signal optical imaging, we found that the orientation of second-order contour stimuli was represented invariantly in the orientation columns of both macaque V1 and V2. A physiologically constrained spatio-temporal energy model of V1 and V2 neuronal populations could reproduce all the recorded population responses. These findings suggest that, at the population level, the primate early visual system processes the orientation of second-order contours initially through a linear spatio-temporal filter mechanism. Our results of population responses to different second-order contour stimuli support the idea that the orientation maps in primate V1 and V2 can be described as a spatial-temporal energy map.     

Neural mechanisms of tactile motion integration in somatosensory cortex

How are local motion signals integrated to form a global motion percept? We investigate the neural mechanisms of tactile motion integration by presenting tactile gratings and plaids to the fingertips of monkeys, using the tactile analogue of a visual monitor and recording the responses evoked in somatosensory cortical neurons. The perceived directions of the gratings and plaids are measured in parallel psychophysical experiments. We identify a population of somatosensory neurons that exhibit integration properties comparable to those induced by analogous visual stimuli in area MT and find that these neural responses account for the perceived direction of the stimuli across all stimulus conditions tested. The preferred direction of the neurons and the perceived direction of the stimuli can be predicted from the weighted average of the directions of the individual stimulus features, highlighting that the somatosensory system implements a vector average mechanism to compute tactile motion direction that bears striking similarities to its visual counterpart.     

Tactile rivalry demonstrated with an ambiguous apparent-motion quartet

When observers view ambiguous visual stimuli, their perception will often alternate between the possible interpretations, a phenomenon termed perceptual rivalry [1]. To induce perceptual rivalry in the tactile domain, we developed a new tactile illusion, based on the visual apparent-motion quartet [2]. Pairs of 200 ms vibrotactile stimuli were applied to the finger pad at intervals separated by 300 ms. The location of each successive stimulus pair alternated between the opposing diagonal corners of the approximately 1 cm(2) stimulation array. This stimulation sequence led all participants to report switches between the perception of motion traveling either up and down or left and right across their fingertip. Adaptation to tactile stimulation biased toward one direction caused subsequent ambiguous stimulation to be experienced in the opposing direction. In contrast, when consecutive trials of ambiguous stimulation were presented, motion was generally perceived in the direction consistent with the motion reported in the previous trial. Voluntary eye movements induced shifts in the tactile perception toward a motion axis aligned along a world-centered coordinate frame. Because the tactile quartet results in switching perceptual states despite unvaried sensory input, it is ideally suited to future studies of the neural processes associated with conscious tactile perception.     

Orientation acuity estimated with simultaneous and successive procedures.

Orientation discrimination thresholds for medium contrast sinewave grating stimuli were estimated by a forced-choice procedure for the two principal, and the two main oblique axes. The results obtained for simultaneous presentation with a centre/surround stimulus arrangement are compared with the threshold estimates obtained when the two stimuli to be discriminated do not overlap in time. We find that orientation acuity depends on the temporal relationship of the discriminanda. Estimates of orientation threshold with simultaneous presentation are lower on all axes tested than those obtained with successive presentation; the greatest difference in orientation acuity was found at obliques. The data support the hypothesis that the meridional anisotropy in orientation discrimination that is found with conventional, two-interval forced-choice methods is not entirely attributable to anatomical and physiological changes at the level of the striate cortex.     

Comparison of the mechanisms of latency shift in pattern reversal visual evoked potential induced by blurring and contrast reduction

Reduction of visual acuity or of the contrast of the stimulus induces a prolongation of the pattern reversal visual evoked potential (PR-VEP) latencies, perhaps because these conditions cause deterioration of the visual capacity to recognize objects and may preferentially activate the slower central retina channel. The PR-VEP was obtained with a video stimulator and 3 kinds of stimuli: total video field, video with a central scotoma and a restricted central stimulus. The subjects were tested under conditions of normal (20/20) and reduced visual acuity (20/200) with 14′ and 56′ checks and 60% contrast, and under conditions of normal visual acuity (20/20) with 14′ checks and with stimulus contrast of 60% and 25%. Blurring increased latencies and decreased amplitudes only with the 14′ checks stimulus but no with 56′ checks, and the amplitudes obtained with the central stimulus became greater than those obtained with a central scotoma. Reducing contrast increased only latency, and there was not difference between amplitudes obtained with a central stimulus or a central scotoma. We conclude that blurring small checks induces a preferential stimulation of receptors in the central retina, but the same effect was not observed when stimulus contrast was reduced.     

Orientation Transfer in Vernier and Stereoacuity Training

Human performance on various visual tasks can be improved substantially via training. However, the enhancements are frequently specific to relatively low-level stimulus dimensions. While such specificity has often been thought to be indicative of a low-level neural locus of learning, recent research suggests that these same effects can be accounted for by changes in higher-level areas–in particular in the way higher-level areas read out information from lower-level areas in the service of highly practiced decisions. Here we contrast the degree of orientation transfer seen after training on two different tasks—vernier acuity and stereoacuity. Importantly, while the decision rule that could improve vernier acuity (i.e. a discriminant in the image plane) would not be transferable across orientations, the simplest rule that could be learned to solve the stereoacuity task (i.e. a discriminant in the depth plane) would be insensitive to changes in orientation. Thus, given a read-out hypothesis, more substantial transfer would be expected as a result of stereoacuity than vernier acuity training. To test this prediction, participants were trained (7500 total trials) on either a stereoacuity (N = 9) or vernier acuity (N = 7) task with the stimuli in either a vertical or horizontal configuration (balanced across participants). Following training, transfer to the untrained orientation was assessed. As predicted, evidence for relatively orientation specific learning was observed in vernier trained participants, while no evidence of specificity was observed in stereo trained participants. These results build upon the emerging view that perceptual learning (even very specific learning effects) may reflect changes in inferences made by high-level areas, rather than necessarily fully reflecting changes in the receptive field properties of low-level areas.     

Representation of moving stimuli by somatosensory neurons.

The findings obtained in neurophysiological and psychophysical investigations using tactile stimuli that move at constant velocity across the skin are reviewed. For certain neurons in the postcentral gyrus of the cerebral cortex (S-I) of macaque monkeys, direction of stimulus motion is a "trigger feature" i.e., moving tactile stimuli evoke vigorous discharge activity in these neurons only if the stimuli are moved in a particular direction across the receptive field. This directional selectivity is maximal when stimulus velocity is between 5 and 50 cm/sec, and falls off rapidly at lower or higher velocities. The capacity for human subjects to correctly identify the direction of stimulus motion on the skin exhibits a similar dependence on stimulus velocity. The similar effects of velocity on neural and psychophysical measures of directional sensitivity support the idea that direction of stimulus motion on the skin can only be recognized if the moving stimulus optimally activates the group of S-I neurons for which that directions of simulus motion is the trigger feature.     

Prey-capture in the African clawed toad (Xenopus laevis): comparison of turning to visual and lateral line stimuli

Separately delivered visual and lateral line stimuli elicit similar but not identical orientation and approach by intact, sighted Xenopus. Response frequencies for visual stimuli declined sharply for distant or caudal stimuli while those for lateral line stimuli changed little. Turn angles correlated highly with stimulus angles but were smaller on average, so regression slopes were less than one. Regression slopes were smaller for visual than for lateral line stimuli, but this apparent difference was due to different distributions of stimulus distance interacting with the toad’s rotation center. Errors in final headings, most often under-rotations, did not differ by modality. Frequencies of lunges and arm capture movements were higher for visual stimuli both overall and especially for rostral proximal stimuli. The results demonstrate accurate orientation by sighted Xenopus to visual and lateral line stimuli; they are consistent with expectations based on in-register tectal maps. Orientation to lateral line stimuli is similar to previous results with blinded animals, revealing no heightened acuity in the latter. Modality differences indicate that the lateral line system is better for omnidirectional orientation and approach to distant stimuli whereas the visual system is more attuned to nearby rostral stimuli and more apt to mediate strikes.Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .     

The effects of cross-modal manipulations of attention on the detection of vibrotactile stimuli in humans

Although it is well known that attention to a visual or auditory stimulus can enhance its perception, less is known concerning the effects of attention on the perception of natural tactile stimuli. The present study was conducted to examine the magnitude of the effect of cross-modal manipulations of attention in human subjects on the detection of weak, low-frequency vibrotactile stimuli delivered to the glabrous skin of the finger pad of the right index finger via an Optacon. Three suprathreshold vibrotactile arrays (40 Hz), varying in the number of activated pegs and hence the area of skin stimulated, were used. Subjects were trained to detect the occurrence of vibrotactile or visual stimuli and to respond by pressing a foot pedal as quickly as possible thereafter. Two instructional lights were used to cue the subjects as to which stimulus modality they should attend, in three experimental conditions. In the first cue condition, the forthcoming stimulus modality was indicated by the illumination of its associated light. In the second cue condition, both instructional lights were illuminated, and the subjects were asked to divide their attention equally between the two modalities. In the third cue condition, the stimulus modality was falsely indicated by the illumination of the cue not associated with the stimulus to be presented. Reaction times (RTs) were calculated for each trial. For each modality, tactile and visual, the RTs varied significantly with the cue condition, with the mean RT changing in a graded manner across the experimental conditions (being shortest for the correctly cued condition, intermediate for the neutrally cued condition, and longest for the incorrectly cued condition.(ABSTRACT TRUNCATED AT 250 WORDS)     

Motions Add,Orientations Don’t,in the Human Visual System

Humans can distinguish between contours of similar orientation, and between directions of visual motion. There is consensus that both of these capabilities depend on selective activation of tuned neural channels. The bandwidths of these tuned channels are estimated here by modelling previously published empirical data. Human subjects were presented with a rapid stream of randomly oriented gratings, or randomly directed motions, and asked to respond when they saw a target stimulus. For the orientation task, subjects were less likely to respond when two preceding orientations were close to the target orientation but differed from each other, presumably due to a failure of summation. For the motion data, by contrast, subjects were more likely to respond when the vector sum of two previous directions was in the target direction. Fitting a cortical signal-processing model to these data showed that the direction bandwidth of motion sensors is about three times the bandwidth of orientation sensors, and that it is the large bandwidth that allows the summation of motion stimuli. The differing bandwidths of orientation and motion sensors presumably equip them for differing tasks, such as orientation discrimination and estimation of heading, respectively.     

A comparison of vernier acuity for narrowband and broadband stimuli

This study investigates the influence of contrast and exposure duration on vernier acuity thresholds for abutting and separated narrowband stimuli, and asks whether these data can predict broadband vernier performance. Vernier thresholds were determined for sinusoidal grating stimuli at two spatial frequencies (1 and 8 c/deg) across a range of contrasts (0.05-0.8) and exposure durations (35-2100 ms). Performance was assessed for the abutting configuration, and when a gap equivalent to 0.5 to 1.5 times the spatial period of the grating was introduced between the upper and lower halves of the grating. Vernier thresholds were also determined for a square-wave stimulus as a function of contrast (0.06 to 0.78). Exposure duration was fixed at 2100 ms. In addition, thresholds were determined at the appropriate contrast levels for the fundamental frequency (1.8 c/deg) of the square-wave, and for a number of the harmonics (3F, 5F, 7F, 9F). Our results provide support for filter models of vernier acuity by showing that vernier performance for abutting and closely-separated broadband stimuli represents the envelope of vernier sensitivity of those spatial frequency mechanisms that are activated by the broadband stimulus. In the case of high frequency grating stimuli presented for long exposure durations, vernier performance can be invariant across much of the contrast range. Despite this, however, contrast independence is not exhibited for abutting broadband stimuli because, within the broadband stimuli, the contrast of the higher harmonic components never reaches a level to reveal this plateau.     

Filling-in and suppression of visual perception from context: a Bayesian account of perceptual biases by contextual influences

Visual object recognition and sensitivity to image features are largely influenced by contextual inputs. We study influences by contextual bars on the bias to perceive or infer the presence of a target bar, rather than on the sensitivity to image features. Human observers judged from a briefly presented stimulus whether a target bar of a known orientation and shape is present at the center of a display, given a weak or missing input contrast at the target location with or without a context of other bars. Observers are more likely to perceive a target when the context has a weaker rather than stronger contrast. When the context can perceptually group well with the would-be target, weak contrast contextual bars bias the observers to perceive a target relative to the condition without contexts, as if to fill in the target. Meanwhile, high-contrast contextual bars, regardless of whether they group well with the target, bias the observers to perceive no target. A Bayesian model of visual inference is shown to account for the data well, illustrating that the context influences the perception in two ways: (1) biasing observers' prior belief that a target should be present according to visual grouping principles, and (2) biasing observers' internal model of the likely input contrasts caused by a target bar. According to this model, our data suggest that the context does not influence the perceived target contrast despite its influence on the bias to perceive the target's presence, thereby suggesting that cortical areas beyond the primary visual cortex are responsible for the visual inferences.     

Orientation processing by synaptic integration across first-order tactile neurons

Our ability to manipulate objects relies on tactile inputs from first-order tactile neurons that innervate the glabrous skin of the hand. The distal axon of these neurons branches in the skin and innervates many mechanoreceptors, yielding spatially-complex receptive fields. Here we show that synaptic integration across the complex signals from the first-order neuronal population could underlie human ability to accurately (< 3°) and rapidly process the orientation of edges moving across the fingertip. We first derive spiking models of human first-order tactile neurons that fit and predict responses to moving edges with high accuracy. We then use the model neurons in simulating the peripheral neuronal population that innervates a fingertip. We train classifiers performing synaptic integration across the neuronal population activity, and show that synaptic integration across first-order neurons can process edge orientations with high acuity and speed. In particular, our models suggest that integration of fast-decaying (AMPA-like) synaptic inputs within short timescales is critical for discriminating fine orientations, whereas integration of slow-decaying (NMDA-like) synaptic inputs supports discrimination of coarser orientations and maintains robustness over longer timescales. Taken together, our results provide new insight into the computations occurring in the earliest stages of the human tactile processing pathway and how they may be critical for supporting hand function.     

Perceptual Grouping Enhances Visual Plasticity

Visual perceptual learning, a manifestation of neural plasticity, refers to improvements in performance on a visual task achieved by training. Attention is known to play an important role in perceptual learning, given that the observer''s discriminative ability improves only for those stimulus feature that are attended. However, the distribution of attention can be severely constrained by perceptual grouping, a process whereby the visual system organizes the initial retinal input into candidate objects. Taken together, these two pieces of evidence suggest the interesting possibility that perceptual grouping might also affect perceptual learning, either directly or via attentional mechanisms. To address this issue, we conducted two experiments. During the training phase, participants attended to the contrast of the task-relevant stimulus (oriented grating), while two similar task-irrelevant stimuli were presented in the adjacent positions. One of the two flanking stimuli was perceptually grouped with the attended stimulus as a consequence of its similar orientation (Experiment 1) or because it was part of the same perceptual object (Experiment 2). A test phase followed the training phase at each location. Compared to the task-irrelevant no-grouping stimulus, orientation discrimination improved at the attended location. Critically, a perceptual learning effect equivalent to the one observed for the attended location also emerged for the task-irrelevant grouping stimulus, indicating that perceptual grouping induced a transfer of learning to the stimulus (or feature) being perceptually grouped with the task-relevant one. Our findings indicate that no voluntary effort to direct attention to the grouping stimulus or feature is necessary to enhance visual plasticity.