Body-part specificity for learning of multiple prior distributions in human coincidence timing.

During timing tasks, the brain learns the statistical distribution of target intervals and integrates this prior knowledge with sensory inputs to optimise task performance. Daily events can have different temporal statistics (e.g., fastball/slowball in baseball batting), making it important to learn and retain multiple priors. However, the rules governing this process are not yet understood. Here, we demonstrate that the learning of multiple prior distributions in a coincidence timing task is characterised by body-part specificity. In our experiments, two prior distributions (short and long intervals) were imposed on participants. When using only one body part for timing responses, regardless of the priors, participants learned a single prior by generalising over the two distributions. However, when the two priors were assigned to different body parts, participants concurrently learned the two independent priors. Moreover, body-part specific prior acquisition was faster when the priors were assigned to anatomically distant body parts (e.g., hand/foot) than when they were assigned to close body parts (e.g., index/middle fingers). This suggests that the body-part specific learning of priors is organised according to somatotopy.

The spatial properties of adaptation-induced distance compression.

Exposure to a dynamic texture reduces the perceived separation between objects, altering the mapping between physical relations in the environment and their neural representations. Here we investigated the spatial tuning and spatial frame of reference of this aftereffect to understand the stage(s) of processing where adaptation-induced changes occur. In Experiment 1, we measured apparent separation at different positions relative to the adapted area, revealing a strong but tightly tuned compression effect. We next tested the spatial frame of reference of the effect, either by introducing a gaze shift between adaptation and test phase (Experiment 2) or by decoupling the spatial selectivity of adaptation in retinotopic and world-centered coordinates (Experiment 3). Results across the two experiments indicated that both retinotopic and world-centered adaptation effects can occur independently. Spatial attention to the location of the adaptor alone could not account for the world-centered transfer we observed, and retinotopic adaptation did not transfer to world-centered coordinates after a saccade (Experiment 4). Finally, we found that aftereffects in different reference frames have a similar, narrow spatial tuning profile (Experiment 5). Together, our results suggest that the neural representation of local separation resides early in the visual cortex, but it can also be modulated by activity in higher visual areas.

Multiple spatial reference frames underpin perceptual recalibration to audio-visual discrepancies.

In dynamic multisensory environments, the perceptual system corrects for discrepancies arising between modalities. For instance, in the ventriloquism aftereffect (VAE), spatial disparities introduced between visual and auditory stimuli lead to a perceptual recalibration of auditory space. Previous research has shown that the VAE is underpinned by multiple recalibration mechanisms tuned to different timescales, however it remains unclear whether these mechanisms use common or distinct spatial reference frames. Here we asked whether the VAE operates in eye- or head-centred reference frames across a range of adaptation timescales, from a few seconds to a few minutes. We developed a novel paradigm for selectively manipulating the contribution of eye- versus head-centred visual signals to the VAE by manipulating auditory locations relative to either the head orientation or the point of fixation. Consistent with previous research, we found both eye- and head-centred frames contributed to the VAE across all timescales. However, we found no evidence for an interaction between spatial reference frames and adaptation duration. Our results indicate that the VAE is underpinned by multiple spatial reference frames that are similarly leveraged by the underlying time-sensitive mechanisms.

Learning to silence saccadic suppression.

Perceptual stability is facilitated by a decrease in visual sensitivity during rapid eye movements, called saccadic suppression. While a large body of evidence demonstrates that saccadic programming is plastic, little is known about whether the perceptual consequences of saccades can be modified. Here, we demonstrate that saccadic suppression is attenuated during learning on a standard visual detection-in-noise task, to the point that it is effectively silenced. Across a period of 7 days, 44 participants were trained to detect brief, low-contrast stimuli embedded within dynamic noise, while eye position was tracked. Although instructed to fixate, participants regularly made small fixational saccades. Data were accumulated over a large number of trials, allowing us to assess changes in performance as a function of the temporal proximity of stimuli and saccades. This analysis revealed that improvements in sensitivity over the training period were accompanied by a systematic change in the impact of saccades on performance-robust saccadic suppression on day 1 declined gradually over subsequent days until its magnitude became indistinguishable from zero. This silencing of suppression was not explained by learning-related changes in saccade characteristics and generalized to an untrained retinal location and stimulus orientation. Suppression was restored when learned stimulus timing was perturbed, consistent with the operation of a mechanism that temporarily reduces or eliminates saccadic suppression, but only when it is behaviorally advantageous to do so. Our results indicate that learning can circumvent saccadic suppression to improve performance, without compromising its functional benefits in other viewing contexts.

Impact of microsaccades on visual shape processing.

Sensitivity to subtle changes in the shape of visual objects has been attributed to the existence of global pooling mechanisms that integrate local form information across space. Although global pooling is typically demonstrated under steady fixation, other work suggests prolonged fixation can lead to a collapse of global structure. Here, we ask whether small ballistic eye movements that naturally occur during periods of fixation affect the global processing of radial frequency (RF) patterns-closed contours created by sinusoidally modulating the radius of a circle. Observers were asked to discriminate the shapes of circular patterns and RF-modulated patterns while fixational eye movements were recorded binocularly at 500 Hz. Microsaccades were detected using a velocity-based algorithm, allowing trials to be sorted according to the relative timing of stimulus and microsaccade onset. Results revealed clear perisaccadic changes in shape discrimination thresholds. Performance was impaired when microsaccades occurred close to stimulus onset, but facilitated when they occurred shortly afterward. In contrast, global integration of shape was unaffected by the timing of microsaccades. These findings suggest that microsaccades alter the discrimination sensitivity to briefly presented shapes but do not disrupt the spatial pooling of local form signals.NEW & NOTEWORTHY Microsaccades cause rapid displacement of visual images during fixation and dramatically alter the perception of basic image features. However, their effect on more complex aspects of visual processing is not well understood. Here, we demonstrate a dissociation in the impact of microsaccades on shape perception. Although overall shape discrimination performance is modulated around the time of microsaccades, the pooling efficiency of global mechanisms that combine local form information across space remains unaffected.

Publisher Correction: Temporal rate is not a distinct perceptual metric.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Temporal rate is not a distinct perceptual metric.

Sensory adaptation experiments have revealed the existence of 'rate after-effects' - adapting to a relatively fast rate makes an intermediate test rate feel slow, and adapting to a slow rate makes the same moderate test rate feel fast. The present work aims to deconstruct the concept of rate and clarify how exactly the brain processes a regular sequence of sensory signals. We ask whether rate forms a distinct perceptual metric, or whether it is simply the perceptual aggregate of the intervals between its component signals. Subjects were exposed to auditory or visual temporal rates (a 'slow' rate of 1.5 Hz and a 'fast' rate of 6 Hz), before being tested with single unfilled intervals of varying durations. Results show adapting to a given rate strongly influences the perceived duration of a single empty interval. This effect is robust across both interval reproduction and duration discrimination judgments. These findings challenge our understanding of rate perception. Specifically, they suggest that contrary to some previous assertions, the perception of sequence rate is strongly influenced by the perception of the sequence's component duration intervals.

Temporal modulation improves dynamic peripheral acuity.

Macular degeneration and related visual disorders greatly limit foveal function, resulting in reliance on the peripheral retina for tasks requiring fine spatial vision. Here we investigate stimulus manipulations intended to maximize peripheral acuity for dynamic targets. Acuity was measured using a single interval orientation discrimination task at 10° eccentricity. Two types of image motion were investigated along with two different forms of temporal manipulation. Smooth object motion was generated by translating targets along an isoeccentric path at a constant speed (0-20°/s). Ocular motion was simulated by jittering target location using previously recorded fixational eye movement data, amplified by a variable gain factor (0-8). In one stimulus manipulation, the sequence was temporally subsampled by displaying the target on an evenly spaced subset of video frames. In the other, the contrast polarity of the stimulus was reversed at a variable rate. We found that threshold under object motion was improved at all speeds by reversing contrast polarity, while temporal subsampling improved resolution at high speeds but impaired performance at low speeds. With simulated ocular motion, thresholds were consistently improved by contrast polarity reversal, but impaired by temporal subsampling. We find that contrast polarity reversal and temporal subsampling produce differential effects on peripheral acuity. Applying contrast polarity reversal may offer a relatively simple image manipulation that could enhance visual performance in individuals with central vision loss.

Distinct mechanisms govern recalibration to audio-visual discrepancies in remote and recent history.

To maintain perceptual coherence, the brain corrects for discrepancies between the senses. If, for example, lights are consistently offset from sounds, representations of auditory space are remapped to reduce this error (spatial recalibration). While recalibration effects have been observed following both brief and prolonged periods of adaptation, the relative contribution of discrepancies occurring over these timescales is unknown. Here we show that distinct multisensory recalibration mechanisms operate in remote and recent history. To characterise the dynamics of this spatial recalibration, we adapted human participants to audio-visual discrepancies for different durations, from 32 to 256 seconds, and measured the aftereffects on perceived auditory location. Recalibration effects saturated rapidly but decayed slowly, suggesting a combination of transient and sustained adaptation mechanisms. When long-term adaptation to an audio-visual discrepancy was immediately followed by a brief period of de-adaptation to an opposing discrepancy, recalibration was initially cancelled but subsequently reappeared with further testing. These dynamics were best fit by a multiple-exponential model that monitored audio-visual discrepancies over distinct timescales. Recent and remote recalibration mechanisms enable the brain to balance rapid adaptive changes to transient discrepancies that should be quickly forgotten against slower adaptive changes to persistent discrepancies likely to be more permanent.

Adaptation reveals multi-stage coding of visual duration.

In conflict with historically dominant models of time perception, recent evidence suggests that the encoding of our environment's temporal properties may not require a separate class of neurons whose raison d'être is the dedicated processing of temporal information. If true, it follows that temporal processing should be imbued with the known selectivity found within non-temporal neurons. In the current study, we tested this hypothesis for the processing of a poorly understood stimulus parameter: visual event duration. We used sensory adaptation techniques to generate duration aftereffects: bidirectional distortions of perceived duration. Presenting adapting and test durations to the same vs different eyes utilises the visual system's anatomical progression from monocular, pre-cortical neurons to their binocular, cortical counterparts. Duration aftereffects exhibited robust inter-ocular transfer alongside a small but significant contribution from monocular mechanisms. We then used novel stimuli which provided duration information that was invisible to monocular neurons. These stimuli generated robust duration aftereffects which showed partial selectivity for adapt-test changes in retinal disparity. Our findings reveal distinct duration encoding mechanisms at monocular, depth-selective and depth-invariant stages of the visual hierarchy.

Estimation of Contrast Sensitivity From Fixational Eye Movements.

Purpose: Even during steady fixation, people make small eye movements such as microsaccades, whose rate is altered by presentation of salient stimuli. Our goal was to develop a practical method for objectively and robustly estimating contrast sensitivity from microsaccade rates in a diverse population. Methods: Participants, recruited to cover a range of contrast sensitivities, were visually normal (n = 19), amblyopic (n = 10), or had cataract (n = 9). Monocular contrast sensitivity was estimated behaviorally while binocular eye movements were recorded during interleaved passive trials. A probabilistic inference approach was used to establish the likelihood of observed microsaccade rates given the presence or absence of a salient stimulus. Contrast sensitivity was estimated from a function fitted to the scaled log-likelihood ratio of the observed microsaccades in the presence or absence of a salient stimulus across a range of contrasts. Results: Microsaccade rate signature shapes were heterogeneous; nevertheless, estimates of contrast sensitivity could be obtained in all participants. Microsaccade-estimated contrast sensitivity was unbiased compared to behavioral estimates (1.2% mean), with which they were strongly correlated (Spearman's ρ 0.74, P < 0.001, median absolute difference 7.6%). Measurement precision of microsaccade-based contrast sensitivity estimates was worse than that of behavioral estimates, requiring more than 20 times as many presentations to equate precision. Conclusions: Microsaccade rate signatures are heterogeneous in shape when measured across populations with a broad range of contrast sensitivities. Contrast sensitivity can be robustly estimated from rate signatures by probabilistic inference, but more stimulus presentations are currently required to achieve similarly precise estimates to behavioral techniques.

Visual crowding is unaffected by adaptation-induced spatial compression.

It has recently been shown that adapting to a densely textured stimulus alters the perception of visual space, such that the distance between two points subsequently presented in the adapted region appears reduced (Hisakata, Nishida, & Johnston, 2016). We asked whether this form of adaptation-induced spatial compression alters visual crowding. To address this question, we first adapted observers to a dynamic dot texture presented within an annular region surrounding the test location. Following adaptation, observers perceived a test array comprised of multiple oriented dot dipoles as spatially compressed, resulting in an overall reduction in perceived size. We then tested to what extent this spatial compression influences crowding by measuring orientation discrimination of a single dipole flanked by randomly oriented dipoles across a range of separations. Following adaptation, we found that the magnitude of crowding was predicted by the physical rather than perceptual separation between center and flanking dipoles. These findings contrast with previous studies in which crowding has been shown to increase when motion-induced position shifts act to reduce apparent separation (Dakin, Greenwood, Carlson, & Bex, 2011; Maus, Fischer, & Whitney, 2011).

Selective modulation of visual sensitivity during fixation.

During periods of steady fixation, we make small-amplitude ocular movements, termed microsaccades, at a rate of 1-2 every second. Early studies provided evidence that visual sensitivity is reduced during microsaccades-akin to the well-established suppression associated with larger saccades. However, the results of more recent work suggest that microsaccades may alter retinal input in a manner that enhances visual sensitivity to some stimuli. Here we parametrically varied the spatial frequency of a stimulus during a detection task and tracked contrast sensitivity as a function of time relative to microsaccades. Our data reveal two distinct modulations of sensitivity: suppression during the eye movement itself and facilitation after the eye has stopped moving. The magnitude of suppression and facilitation of visual sensitivity is related to the spatial content of the stimulus: suppression is greatest for low spatial frequencies, while sensitivity is enhanced most for stimuli of 1-2 cycles/°, spatial frequencies at which we are already most sensitive in the absence of eye movements. We present a model in which the tuning of suppression and facilitation is explained by delayed lateral inhibition between spatial frequency channels. Our data show that eye movements actively modulate visual sensitivity even during fixation: the detectability of images at different spatial scales can be increased or decreased depending on when the image occurs relative to a microsaccade. NEW & NOTEWORTHY Given the frequency with which we make microsaccades during periods of fixation, it is vital that we understand how they affect visual processing. We demonstrate two selective modulations of contrast sensitivity that are time-locked to the occurrence of a microsaccade: suppression of low spatial frequencies during each eye movement and enhancement of higher spatial frequencies after the eye has stopped moving. These complementary changes may arise naturally because of sluggish gain control between spatial channels.

Rate after-effects fail to transfer cross-modally: Evidence for distributed sensory timing mechanisms.

Accurate time perception is critical for a number of human behaviours, such as understanding speech and the appreciation of music. However, it remains unresolved whether sensory time perception is mediated by a central timing component regulating all senses, or by a set of distributed mechanisms, each dedicated to a single sensory modality and operating in a largely independent manner. To address this issue, we conducted a range of unimodal and cross-modal rate adaptation experiments, in order to establish the degree of specificity of classical after-effects of sensory adaptation. Adapting to a fast rate of sensory stimulation typically makes a moderate rate appear slower (repulsive after-effect), and vice versa. A central timing hypothesis predicts general transfer of adaptation effects across modalities, whilst distributed mechanisms predict a high degree of sensory selectivity. Rate perception was quantified by a method of temporal reproduction across all combinations of visual, auditory and tactile senses. Robust repulsive after-effects were observed in all unimodal rate conditions, but were not observed for any cross-modal pairings. Our results show that sensory timing abilities are adaptable but, crucially, that this change is modality-specific - an outcome that is consistent with a distributed sensory timing hypothesis.

New insights into the role of motion and form vision in neurodevelopmental disorders.

A selective deficit in processing the global (overall) motion, but not form, of spatially extensive objects in the visual scene is frequently associated with several neurodevelopmental disorders, including preterm birth. Existing theories that proposed to explain the origin of this visual impairment are, however, challenged by recent research. In this review, we explore alternative hypotheses for why deficits in the processing of global motion, relative to global form, might arise. We describe recent evidence that has utilised novel tasks of global motion and global form to elucidate the underlying nature of the visual deficit reported in different neurodevelopmental disorders. We also examine the role of IQ and how the sex of an individual can influence performance on these tasks, as these are factors that are associated with performance on global motion tasks, but have not been systematically controlled for in previous studies exploring visual processing in clinical populations. Finally, we suggest that a new theoretical framework is needed for visual processing in neurodevelopmental disorders and present recommendations for future research.

Motion-based super-resolution in the peripheral visual field.

Improvements in foveal acuity for moving targets have been interpreted as evidence for the ability of the visual system to combine information over space and time, in order to reconstruct the image at a higher resolution (super-resolution). Here, we directly test whether this occurs in the peripheral visual field and discuss its potential for improving functional capacity in ocular disease. The effect of motion on visual acuity was first compared under conditions in which performance was limited either by natural undersampling in the retinal periphery or by the presence of overlaid masks with opaque elements to simulate retinal loss. To equate the information content of moving and static sequences, we next manipulated the dynamic properties of the masks. Finally, we determined the dependence of motion-related improvements on the object of motion (target or mask) and its trajectory (smooth or jittered). Motion improved visual acuity for masked but not unmasked peripheral targets. Equating the information content of moving and static conditions removed some but not all of this benefit. Residual motion-related improvements were largest in conditions in which the target moved along a consistent and predictable path. Our results show that motion can improve peripheral acuity in situations in which performance is limited by abnormal undersampling. These findings are consistent with the operation of a super-resolution system and could have important implications for any pathology that alters the regular sampling properties of the retinal mosaic.

Visual perception in dyslexia is limited by sub-optimal scale selection.

Readers with dyslexia are purported to have a selective visual impairment but the underlying nature of the deficit remains elusive. Here, we used a combination of behavioural psychophysics and biologically-motivated computational modeling to investigate if this deficit extends to object segmentation, a process implicated in visual word form recognition. Thirty-eight adults with a wide range of reading abilities were shown random-dot displays spatially divided into horizontal segments. Adjacent segments contained either local motion signals in opposing directions or analogous static form cues depicting orthogonal orientations. Participants had to discriminate these segmented patterns from stimuli containing identical motion or form cues that were spatially intermingled. Results showed participants were unable to perform the motion or form task reliably when segment size was smaller than a spatial resolution (acuity) limit that was independent of reading skill. Coherence thresholds decreased as segment size increased, but for the motion task the rate of improvement was shallower for readers with dyslexia and the segment size where performance became asymptotic was larger. This suggests that segmentation is impaired in readers with dyslexia but only on tasks containing motion information. We interpret these findings within a novel framework in which the mechanisms underlying scale selection are impaired in developmental dyslexia.

Encoding of rapid time-varying information is impaired in poor readers.

A characteristic set of eye movements and fixations are made during reading, so the position of words on the retinae is constantly being updated. Effective decoding of print requires this temporal stream of visual information to be segmented or parsed into its constituent units (e.g., letters or words). Poor readers' difficulties with word recognition could arise at the point of segmenting time-varying visual information, but the mechanisms underlying this process are little understood. Here, we used random-dot displays to explore the effects of reading ability on temporal segmentation. Thirty-eight adult readers viewed test stimuli that were temporally segmented by constraining either local motions or analogous form cues to oscillate back and fourth at each of a range of rates. Participants had to discriminate these segmented patterns from comparison stimuli containing the same motion and form cues but these were temporally intermingled. Results showed that the motion and form tasks could not be performed reliably when segment duration was shorter than a temporal resolution (acuity) limit. The acuity limits for both tasks were significantly and negatively correlated with reading scores. Importantly, the minimum segment duration needed to detect the temporally segmented stimuli was longer in relatively poor readers than relatively good readers. This demonstrates that adult poor readers have difficulty segmenting temporally changing visual input particularly at short segment durations. These results are consistent with evidence suggesting that precise encoding of rapid time-varying information is impaired in developmental dyslexia.

Generalization of prior information for rapid Bayesian time estimation.

To enable effective interaction with the environment, the brain combines noisy sensory information with expectations based on prior experience. There is ample evidence showing that humans can learn statistical regularities in sensory input and exploit this knowledge to improve perceptual decisions and actions. However, fundamental questions remain regarding how priors are learned and how they generalize to different sensory and behavioral contexts. In principle, maintaining a large set of highly specific priors may be inefficient and restrict the speed at which expectations can be formed and updated in response to changes in the environment. However, priors formed by generalizing across varying contexts may not be accurate. Here, we exploit rapidly induced contextual biases in duration reproduction to reveal how these competing demands are resolved during the early stages of prior acquisition. We show that observers initially form a single prior by generalizing across duration distributions coupled with distinct sensory signals. In contrast, they form multiple priors if distributions are coupled with distinct motor outputs. Together, our findings suggest that rapid prior acquisition is facilitated by generalization across experiences of different sensory inputs but organized according to how that sensory information is acted on.

Why is the processing of global motion impaired in adults with developmental dyslexia?

Individuals with dyslexia are purported to have a selective dorsal stream impairment that manifests as a deficit in perceiving visual global motion relative to global form. However, the underlying nature of the visual deficit in readers with dyslexia remains unclear. It may be indicative of a difficulty with motion detection, temporal processing, or any task that necessitates integration of local visual information across multiple dimensions (i.e. both across space and over time). To disentangle these possibilities we administered four diagnostic global motion and global form tasks to a large sample of adult readers (N=106) to characterise their perceptual abilities. Two sets of analyses were conducted. First, to investigate if general reading ability is associated with performance on the visual tasks across the entire sample, a composite reading score was calculated and entered into a series of continuous regression analyses. Next, to investigate if the performance of readers with dyslexia differs from that of good readers on the visual tasks we identified a group of forty-three individuals for whom phonological decoding was specifically impaired, consistent with the dyslexic profile, and compared their performance with that of good readers who did not exhibit a phonemic deficit. Both analyses yielded a similar pattern of results. Consistent with previous research, coherence thresholds of poor readers were elevated on a random-dot global motion task and a spatially one-dimensional (1-D) global motion task, but no difference was found on a static global form task. However, our results extend those of previous studies by demonstrating that poor readers exhibited impaired performance on a temporally-defined global form task, a finding that is difficult to reconcile with the dorsal stream vulnerability hypothesis. This suggests that the visual deficit in developmental dyslexia does not reflect an impairment detecting motion per se. It is better characterised as a difficulty processing temporal information, which is exacerbated when local visual cues have to be integrated across multiple (>2) dimensions.