Criterion-free measurement of motion transparency perception at different speeds.

Transparency perception often occurs when objects within the visual scene partially occlude each other or move at the same time, at different velocities across the same spatial region. Although transparent motion perception has been extensively studied, we still do not understand how the distribution of velocities within a visual scene contribute to transparent perception. Here we use a novel psychophysical procedure to characterize the distribution of velocities in a scene that give rise to transparent motion perception. To prevent participants from adopting a subjective decision criterion when discriminating transparent motion, we used an "odd-one-out," three-alternative forced-choice procedure. Two intervals contained the standard-a random-dot-kinematogram with dot speeds or directions sampled from a uniform distribution. The other interval contained the comparison-speeds or directions sampled from a distribution with the same range as the standard, but with a notch of different widths removed. Our results suggest that transparent motion perception is driven primarily by relatively slow speeds, and does not emerge when only very fast speeds are present within a visual scene. Transparent perception of moving surfaces is modulated by stimulus-based characteristics, such as the separation between the means of the overlapping distributions or the range of speeds presented within an image. Our work illustrates the utility of using objective, forced-choice methods to reveal the mechanisms underlying motion transparency perception.

Position matching between the visual fields in strabismus.

The misalignment of visual input in strabismus disrupts positional judgments. We measured positional accuracy in the extrafoveal visual field (1°-7° eccentricity) of a large group of strabismic subjects and a normal control group to identify positional distortions associated with the direction of strabismus. Subjects performed a free localization task in which targets were matched in opposite hemifields whilst fixating on a central cross. The constant horizontal error of each response was taken as a measure of accuracy, in addition to radial and angular error. In monocular conditions, all stimuli were viewed by one eye; thus, the error reflected spatial bias. In dichoptic conditions, the targets were seen by separate eyes; thus, the error reflected the perceived stimulus shift produced by ocular misalignment in addition to spatial bias. In both viewing conditions, both groups showed reliable over- and underestimations of visual field position, here termed a compression of response coordinates. The normal group showed compression in the left periphery, regardless of eye of stimulation. The strabismic group showed a visual field-specific compression that was clearly associated with direction of strabismus. The variation in perceived shift of strabismic subjects was largely accounted for by the biases present in monocular viewing, suggesting that binocular correspondence was uniform in the tested region. The asymmetric strabismic compression could not be reproduced in normal subjects through prism viewing, and its presence across viewing conditions suggests a hemifield-specific change in spatial coding induced by long-standing ocular misalignment.

The effect of normal aging and age-related macular degeneration on perceptual learning.

We investigated whether perceptual learning could be used to improve peripheral word identification speed. The relationship between the magnitude of learning and age was established in normal participants to determine whether perceptual learning effects are age invariant. We then investigated whether training could lead to improvements in patients with age-related macular degeneration (AMD). Twenty-eight participants with normal vision and five participants with AMD trained on a word identification task. They were required to identify three-letter words, presented 10° from fixation. To standardize crowding across each of the letters that made up the word, words were flanked laterally by randomly chosen letters. Word identification performance was measured psychophysically using a staircase procedure. Significant improvements in peripheral word identification speed were demonstrated following training (71% ± 18%). Initial task performance was correlated with age, with older participants having poorer performance. However, older adults learned more rapidly such that, following training, they reached the same level of performance as their younger counterparts. As a function of number of trials completed, patients with AMD learned at an equivalent rate as age-matched participants with normal vision. Improvements in word identification speed were maintained at least 6 months after training. We have demonstrated that temporal aspects of word recognition can be improved in peripheral vision with training across a range of ages and these learned improvements are relatively enduring. However, training targeted at other bottlenecks to peripheral reading ability, such as visual crowding, may need to be incorporated to optimize this approach.

Estimation of cortical magnification from positional error in normally sighted and amblyopic subjects.

We describe a method for deriving the linear cortical magnification factor from positional error across the visual field. We compared magnification obtained from this method between normally sighted individuals and amblyopic individuals, who receive atypical visual input during development. The cortical magnification factor was derived for each subject from positional error at 32 locations in the visual field, using an established model of conformal mapping between retinal and cortical coordinates. Magnification of the normally sighted group matched estimates from previous physiological and neuroimaging studies in humans, confirming the validity of the approach. The estimate of magnification for the amblyopic group was significantly lower than the normal group: by 4.4 mm deg(-1) at 1° eccentricity, assuming a constant scaling factor for both groups. These estimates, if correct, suggest a role for early visual experience in establishing retinotopic mapping in cortex. We discuss the implications of altered cortical magnification for cortical size, and consider other neural changes that may account for the amblyopic results.

Characterizing the effects of multidirectional motion adaptation.

Recent sensory experience can alter our perception and change the response characteristics of sensory neurons. These effects of sensory adaptation are a ubiquitous property of perceptual systems and are believed to be of fundamental importance to sensory coding. Yet we know little about how adaptation to stimulus ensembles affects our perception of the environment as most psychophysical experiments employ adaptation protocols that focus on prolonged exposure to a single visual attribute. Here, we investigate how concurrent adaptation to multiple directions of motion affects perception of subsequently presented motion using the direction aftereffect. In different conditions, observers adapted to a stimulus ensemble comprised of dot directions sampled from different distributions or to bidirectional motion. Increasing the variance of normally distributed directions reduced the magnitude of the peak direction aftereffect and broadened its tuning profile. Sampling of asymmetric Gaussian and uniform distributions resulted in shifts of direction aftereffect tuning profiles consistent with changes in the perceived global direction of the adapting stimulus. Adding dots in a direction opposite or orthogonal to a unidirectional adapting stimulus led to a pronounced reduction in the direction aftereffect. A simple population-coding model, in which adaptation selectively alters the responsivity of direction-selective neurons, can accommodate the effects of multidirectional adaptation on the perceived direction of motion.

Perceptual learning reconfigures the effects of visual adaptation.

Our sensory experiences over a range of different timescales shape our perception of the environment. Two particularly striking short-term forms of plasticity with manifestly different time courses and perceptual consequences are those caused by visual adaptation and perceptual learning. Although conventionally treated as distinct forms of experience-dependent plasticity, their neural mechanisms and perceptual consequences have become increasingly blurred, raising the possibility that they might interact. To optimize our chances of finding a functionally meaningful interaction between learning and adaptation, we examined in humans the perceptual consequences of learning a fine discrimination task while adapting the neurons that carry most information for performing this task. Learning improved discriminative accuracy to a level that ultimately surpassed that in an unadapted state. This remarkable improvement came at a price: adapting directions that before learning had little effect elevated discrimination thresholds afterward. The improvements in discriminative accuracy grew quickly and surpassed unadapted levels within the first few training sessions, whereas the deterioration in discriminative accuracy had a different time course. This learned reconfiguration of adapted discriminative accuracy occurred without a concomitant change to the characteristic perceptual biases induced by adaptation, suggesting that the system was still in an adapted state. Our results point to a functionally meaningful push-pull interaction between learning and adaptation in which a gain in sensitivity in one adapted state is balanced by a loss of sensitivity in other adapted states.

Perceptual learning reduces crowding in amblyopia and in the normal periphery.

Amblyopia is a developmental visual disorder of cortical origin, characterized by crowding and poor acuity in central vision of the affected eye. Crowding refers to the adverse effects of surrounding items on object identification, common only in normal peripheral but not central vision. We trained a group of adult human amblyopes on a crowded letter identification task to assess whether the crowding problem can be ameliorated. Letter size was fixed well above the acuity limit, and letter spacing was varied to obtain spacing thresholds for central target identification. Normally sighted observers practiced the same task in their lower peripheral visual field. Independent measures of acuity were taken in flanked and unflanked conditions before and after training to measure crowding ratios at three fixed letter separations. Practice improved the letter spacing thresholds of both groups on the training task, and crowding ratios were reduced after posttest. The reductions in crowding in amblyopes were associated with improvements in standard measures of visual acuity. Thus, perceptual learning reduced the deleterious effects of crowding in amblyopia and in the normal periphery. The results support the effectiveness of plasticity-based approaches for improving vision in adult amblyopes and suggest experience-dependent effects on the cortical substrates of crowding.

Neural computations governing spatiotemporal pooling of visual motion signals in humans.

The brain estimates visual motion by decoding the responses of populations of neurons. Extracting unbiased motion estimates from early visual cortical neurons is challenging because each neuron contributes an ambiguous (local) representation of the visual environment and inherently variable neural response. To mitigate these sources of noise, the brain can pool across large populations of neurons, pool the response of each neuron over time, or a combination of the two. Recent psychophysical and physiological work points to a flexible motion pooling system that arrives at different computational solutions over time and for different stimuli. Here we ask whether a single, likelihood-based computation can accommodate the flexible nature of spatiotemporal motion pooling in humans. We examined the contribution of different computations to human observers' performance on two global visual motion discriminations tasks, one requiring the combination of motion directions over time and another requiring their combination in different relative proportions over space and time. Observers' perceived direction of global motion was accurately predicted by a vector average readout of direction signals accumulated over time and a maximum likelihood readout of direction signals combined over space, consistent with the notion of a flexible motion pooling system that uses different computations over space and time. Additional simulations of observers' performance with a population decoding model revealed a more parsimonious solution: flexible spatiotemporal pooling could be accommodated by a single computation that optimally pools motion signals across a population of neurons that accumulate local motion signals on their receptive fields at a fixed rate over time.

Transfer of perceptual learning between different visual tasks.

Practice in most sensory tasks substantially improves perceptual performance. A hallmark of this 'perceptual learning' is its specificity for the basic attributes of the trained stimulus and task. Recent studies have challenged the specificity of learned improvements, although transfer between substantially different tasks has yet to be demonstrated. Here, we measure the degree of transfer between three distinct perceptual tasks. Participants trained on an orientation discrimination, a curvature discrimination, or a 'global form' task, all using stimuli comprised of multiple oriented elements. Before and after training they were tested on all three and a contrast discrimination control task. A clear transfer of learning was observed, in a pattern predicted by the relative complexity of the stimuli in the training and test tasks. Our results suggest that sensory improvements derived from perceptual learning can transfer between very different visual tasks.

Can perceptual learning be used to treat amblyopia beyond the critical period of visual development?

BACKGROUND: Amblyopia presents early in childhood and affects approximately 3% of western populations. The monocular visual acuity loss is conventionally treated during the 'critical periods' of visual development by occluding or penalising the fellow eye to encourage use of the amblyopic eye. Despite the measurable success of this approach in many children, substantial numbers of people still suffer with amblyopia later in life because either they were never diagnosed in childhood, did not respond to the original treatment, the amblyopia was only partially remediated, or their acuity loss returned after cessation of treatment. PURPOSE: In this review, we consider whether the visual deficits of this largely overlooked amblyopic group are amenable to conventional and innovative therapeutic interventions later in life, well beyond the age at which treatment is thought to be effective. RECENT FINDINGS: There is a considerable body of evidence that residual plasticity is present in the adult visual brain and this can be harnessed to improve function in adults with amblyopia. Perceptual training protocols have been developed to optimise visual gains in this clinical population. Results thus far are extremely encouraging; marked visual improvements have been demonstrated, the perceptual benefits transfer to new visual tasks and appear to be relatively enduring. The essential ingredients of perceptual training protocols are being incorporated into video game formats, facilitating home-based interventions. SUMMARY: Many studies support perceptual training as a tool for improving vision in amblyopes beyond the critical period. Should this novel form of treatment stand up to the scrutiny of a randomised controlled trial, clinicians may need to re-evaluate their therapeutic approach to adults with amblyopia.

Linking Multi-Modal MRI to Clinical Measures of Visual Field Loss After Stroke.

Loss of vision across large parts of the visual field is a common and devastating complication of cerebral strokes. In the clinic, this loss is quantified by measuring the sensitivity threshold across the field of vision using static perimetry. These methods rely on the ability of the patient to report the presence of lights in particular locations. While perimetry provides important information about the intactness of the visual field, the approach has some shortcomings. For example, it cannot distinguish where in the visual pathway the key processing deficit is located. In contrast, brain imaging can provide important information about anatomy, connectivity, and function of the visual pathway following stroke. In particular, functional magnetic resonance imaging (fMRI) and analysis of population receptive fields (pRF) can reveal mismatches between clinical perimetry and maps of cortical areas that still respond to visual stimuli after stroke. Here, we demonstrate how information from different brain imaging modalities-visual field maps derived from fMRI, lesion definitions from anatomical scans, and white matter tracts from diffusion weighted MRI data-provides a more complete picture of vision loss. For any given location in the visual field, the combination of anatomical and functional information can help identify whether vision loss is due to absence of gray matter tissue or likely due to white matter disconnection from other cortical areas. We present a combined imaging acquisition and visual stimulus protocol, together with a description of the analysis methodology, and apply it to datasets from four stroke survivors with homonymous field loss (two with hemianopia, two with quadrantanopia). For researchers trying to understand recovery of vision after stroke and clinicians seeking to stratify patients into different treatment pathways, this approach combines multiple, convergent sources of data to characterize the extent of the stroke damage. We show that such an approach gives a more comprehensive measure of residual visual capacity-in two particular respects: which locations in the visual field should be targeted and what kind of visual attributes are most suited for rehabilitation.

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.

The Effect of Perceptual Learning on Face Recognition in Individuals with Central Vision Loss.

Purpose: To examine whether perceptual learning can improve face discrimination and recognition in older adults with central vision loss. Methods: Ten participants with age-related macular degeneration (ARMD) received 5 days of training on a face discrimination task (mean age, 78 ± 10 years). We measured the magnitude of improvements (i.e., a reduction in threshold size at which faces were able to be discriminated) and whether they generalized to an untrained face recognition task. Measurements of visual acuity, fixation stability, and preferred retinal locus were taken before and after training to contextualize learning-related effects. The performance of the ARMD training group was compared to nine untrained age-matched controls (8 = ARMD, 1 = juvenile macular degeneration; mean age, 77 ± 10 years). Results: Perceptual learning on the face discrimination task reduced the threshold size for face discrimination performance in the trained group, with a mean change (SD) of -32.7% (+15.9%). The threshold for performance on the face recognition task was also reduced, with a mean change (SD) of -22.4% (+2.31%). These changes were independent of changes in visual acuity, fixation stability, or preferred retinal locus. Untrained participants showed no statistically significant reduction in threshold size for face discrimination, with a mean change (SD) of -8.3% (+10.1%), or face recognition, with a mean change (SD) of +2.36% (-5.12%). Conclusions: This study shows that face discrimination and recognition can be reliably improved in ARMD using perceptual learning. The benefits point to considerable perceptual plasticity in higher-level cortical areas involved in face-processing. This novel finding highlights that a key visual difficulty in those suffering from ARMD is readily amenable to rehabilitation.

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 to global structure induces spatially remote distortions of perceived orientation.

Adapting neurons at early stages of cortical visual processing to an oriented grating pattern suppresses responses to nearby orientations, causing a shift in the population response to subsequently viewed stimuli and an illusory distortion of perceived orientation: the classic tilt aftereffect. Perception of stimuli that are presented at unadapted spatial locations or that have a substantially different spatial frequency to the adapting stimulus remains veridical however by virtue of the fact that they stimulate a population of neurons distinct to those affected by adaptation. Here, we demonstrate that adapting to complex visual patterns containing circular or radial structure induces distortions of perceived orientation with markedly different characteristics to those traditionally reported. These novel effects occur in remote, unadapted spatial regions and are immune to manipulations of spatial frequency, rendering them incompatible with direct orientation-specific adaptation in early visual cortex. Rather, our results are most readily explained by a recurrent network in which global form encoding mechanisms that combine orientation information across space actively inhibit the local orientation detectors over which they pool.

Masking exposes multiple global form mechanisms.

Previous work suggests there are mechanisms at intermediate levels of visual processing specialized for the detection of radial and circular form. The evidence in favor of specialized global form mechanisms is derived from structure detection experiments that have told us very little about their bandwidth or number. To address these related questions, we examined the effects of configural backward masking on human observers' ability to detect global structure in arrays with different spiral forms. Each array consisted of 100 Gabors randomly positioned within a circular annular window. Observers judged which of two sequentially presented Gabor arrays contained global structure. One array contained Gabors with random orientations; the other contained Gabors with a variable proportion of orientations coherent with a randomly chosen spiral pitch. At its offset, each array was immediately followed by a backward masking Gabor array with a fixed spiral pitch angle. When mask and test had the same spiral pitch, we found an approximately three-fold elevation of structure detection thresholds that was not explained by local orientation masking. The magnitude and breadth of tuning around each masking angle was predicted by a simple model consisting of at least eight detectors broadly tuned for different spiral forms.

Visual system: how is the retina wired up to the cortex?

A single retinal output neuron transmits to primary visual cortex through multiple pathways with different strengths. A new study in which activity was simultaneously recorded in pairs of retinal and cortical neurons provides evidence that these pathways converge on a single cortical neuron.

Early and late mechanisms of surround suppression in striate cortex of macaque.

The response of a neuron in striate cortex to an optimally configured visual stimulus is generally reduced when the stimulus is enlarged to encroach on a suppressive region that surrounds its classical receptive field (CRF). To characterize the mechanism that gives rise to this suppression, we measured its spatiotemporal tuning, its susceptibility to contrast adaptation, and its capacity for interocular transfer. Responses to an optimally configured grating confined to the CRF were strongly suppressed by annular surrounding gratings drifting at a wide range of temporal and spatial frequencies (including spatially uniform fields) that extended from well below to well above the range that drives most cortical neurons. Suppression from gratings capable of driving cortical CRFs was profoundly reduced by contrast adaptation and showed substantial interocular transfer. Suppression from stimuli that lay outside the spatiotemporal passband of most cortical CRFs was relatively stronger when the stimulus on the CRF was of low contrast, was generally insusceptible to contrast adaptation, and showed little interocular transfer. Our findings point to the existence of two mechanisms of surround suppression: one that is prominent when high-contrast stimuli drive the CRF, is orientation selective, has relatively sharp spatiotemporal tuning, is binocularly driven, and can be substantially desensitized by adaptation; the other is relatively more prominent when low-contrast stimuli drive the CRF, has very broad spatiotemporal tuning, is monocularly driven, and is insusceptible to adaptation. Its character suggests an origin in the input layers of primary visual cortex, or earlier.

The challenges of developing a contrast-based video game for treatment of amblyopia.

Perceptual learning of visual tasks is emerging as a promising treatment for amblyopia, a developmental disorder of vision characterized by poor monocular visual acuity. The tasks tested thus far span the gamut from basic psychophysical discriminations to visually complex video games. One end of the spectrum offers precise control over stimulus parameters, whilst the other delivers the benefits of motivation and reward that sustain practice over long periods. Here, we combined the advantages of both approaches by developing a video game that trains contrast sensitivity, which in psychophysical experiments, is associated with significant improvements in visual acuity in amblyopia. Target contrast was varied adaptively in the game to derive a contrast threshold for each session. We tested the game on 20 amblyopic subjects (10 children and 10 adults), who played at home using their amblyopic eye for an average of 37 sessions (approximately 11 h). Contrast thresholds from the game improved reliably for adults but not for children. However, logMAR acuity improved for both groups (mean = 1.3 lines; range = 0-3.6 lines). We present the rationale leading to the development of the game and describe the challenges of incorporating psychophysical methods into game-like settings.

The effect of aging on crowded letter recognition in the peripheral visual field.

PURPOSE: Crowding describes the increased difficulty in identifying a target object when it is surrounded by nearby objects (flankers). A recent study investigated the effect of age on visual crowding and found equivocal results: Although crowded visual acuity was worse in older participants, crowding expressed as a ratio did not change with age. However, the spatial extent of crowding is a better index of crowding effects and remains unknown. In the present study, we used established psychophysical methods to characterize the effect of age on visual crowding (magnitude and extent) in a letter recognition task. METHODS: Letter recognition thresholds were determined for three different flanker separations in 54 adults (aged 18-76 years) with normal vision. Additionally, the spatial extent of crowding was established by measuring spacing thresholds: the flanker-to-target separation required to produce a given reduction in performance. Uncrowded visual acuity, crowded visual acuity, and spacing thresholds were expressed as a function of age, avoiding arbitrary categorization of young and old participants. RESULTS: Our results showed that uncrowded and crowded visual acuities do not change significantly as a function of age. Furthermore, spacing thresholds did not change with age and approximated Bouma's law (half eccentricity). CONCLUSIONS: These data show that crowding in adults is unaffected by senescence and provide additional evidence for distinct neural mechanisms mediating surround suppression and visual crowding, since the former shows a significant age effect. Finally, our data suggest that the well-documented age-related decline in peripheral reading ability is not due to age-related changes in visual crowding.