Development of contrast normalization mechanisms during childhood and adolescence.

Contrast sensitivity is regulated by neural mechanisms that flexibly adjust responsiveness to optimize stimulus encoding across different environments. Here we studied the developmental status of gain control mechanisms in school-age children (5-17years) and adults using a visual masking paradigm. A variable contrast, spatially random 2-D noise test pattern was masked by the presence of a superimposed independent noise pattern presented at 0, 12 and 40% contrast. Frequency-tagged steady state visual evoked potentials were used to separately record responses to the test (5.14Hz) and the mask (7.2Hz). By incrementally increasing the test contrast we measured contrast response functions for each mask contrast. The unmasked contrast response functions were largely similar in shape across age, but peak amplitude was higher in the children. Masking shifted the contrast response function rightward on the contrast axis in both the adults and older children, elevating contrast thresholds by a similar factor across age. However, in younger children, masking resulted in a change in the slope of the contrast response function. These findings suggest that immaturity in the contrast normalization process persists until approximately 11years of age.

Electrophysiological measures of low-level vision reveal spatial processing deficits and hemispheric asymmetry in autism spectrum disorder.

There is accumulating evidence from electrophysiological studies that low-level visual processing is atypical in individuals with autism spectrum disorders (ASDs). Abnormalities in early stages of sensory processing are of interest because they could lead to downstream functional deficits in social or cognitive domains. Using steady-state visual evoked potentials (SSVEPs), we studied how well spatial information is transmitted over a wide range of spatial frequencies (2-30 cycles/deg), including those at the limit of visibility (visual acuity). SSVEPs were recorded over 128 channels in 16 ASD participants between 5 and 17 years old and 17 age-matched, neurotypical (NT) participants. We observed a selective reduction of the amplitude of the SSVEP second harmonic pattern reversal response between 5 and 17 cycles/deg. Responses measured at the fourth harmonic were normal at all spatial frequencies tested, as were responses at the lowest and highest spatial frequencies at the second harmonic. The reduction of second harmonic responses occurred preferentially over right occipital electrodes. Because response abnormalities are restricted to a specific response harmonic and to specific ranges of spatial frequency, we can rule out nonspecific differences between the ASD participants and the NT controls. This particular pattern of loss, combined with the observed exaggeration of the loss over the right hemisphere, suggests that a highly specific neural substrate early in the visual pathway is compromised in ASD.

The assessment of subjective visual vertical: comparison of two psychophysical paradigms and age-related performance.

Perception of the subjective visual vertical (SVV) is usually assessed by asking to subjects, in complete darkness, to adjust the position of a luminous rod that is variably tilted (i.e., by the method of adjustment [ADJ]). Conversely, the two-alternative forced choice (2AFC) method requires subjects to categorize, as tilted either clockwise (CW) or counterclockwise (CCW), stimuli that are presented on a computer screen and are variably tilted from vertical. In this study, we aimed to compare the results of these two methods and investigate age-related effects on the SVV. SVV was assessed in 102 healthy individuals, 50 women and 52 men, with a mean age of 45.7 (range 20-91), using both ADJ (ten trials, initial 1°, 2°, 4°, 8°, or 12° bar tilts both CW and CCW) and 2AFC (120 stimuli with a 1°-32° variable tilt). Also, 50 of the subjects performed the ADJ test twice, with different bar lengths. We estimated bias and threshold for the two methods, and found that neither measure differed across the methods. Age was a significant predictor of threshold (2AFC, R (2) = .141; ADJ, R (2) = .190; p < .001), implying lower sensitivity with increasing age. Moreover, the ADJ method showed a significant increase of bias when the initial tilt was farthest from vertical, whereas the rod length was irrelevant. SVV measures obtained with the ADJ and 2AFC methods were comparable, but the latter measures were more resistant to artifacts that might affect the measurement. The lower sensitivity found in older persons may have an influence on their ability to interact with the environment and may contribute to impairment of postural control.

Spatiotemporal dynamics of perisaccadic remapping in humans revealed by classification images.

We actively scan our environment with fast ballistic movements called saccades, which create large and rapid displacements of the image on the retina. At the time of saccades, vision becomes transiently distorted in many ways: Briefly flashed stimuli are displaced in space and in time, and spatial and temporal intervals appear compressed. Here we apply the psychophysical technique of classification images to study the spatiotemporal dynamics of visual mechanisms during saccades. We show that saccades cause gross distortions of the classification images. Before the onset of saccadic eye movements, the positive lobes of the images become enlarged in both space and in time and also shifted in a systematic manner toward the pre-saccadic fixation (in space) and anticipated in time by about 50 ms. The transient reorganization creates a spatiotemporal organization oriented in the direction of saccadic-induced motion at the time of saccades, providing a potential mechanism for integrating stimuli across saccades, facilitating stable and continuous vision in the face of constant eye movements.

Spatio-temporal templates of transient attention revealed by classification images.

Visual attention is captured by transient signals in the periphery of the visual field, allowing enhanced perceptual representations in spatial tasks. However, it has been reported that the same cues impair performance in temporal tasks (e.g., Yeshurun, 2004; Yeshurun & Levy, 2003). This findings suggest that transient attention enhances the activity of slow, high-resolution channels, like parvocellular neurons, and/or shuts off faster channels better sensitive to low spatial frequencies, such as the ones of the magnocellular system. To test this idea, we have measured the spatio-temporal perceptive fields for transiently cued signals at various eccentricities using the classification images (CI) technique. At near eccentricities transient attention caused the perceptual templates to be sharper in space and characterized by much stronger high spatial frequency components. At the same time, they show a consistently larger temporal integration window. These effects of attention on perceptual filters are strongly reduced at far eccentricities and disappear when using longer target-cue lags. These data provide evidence in support of the parvocellular model of transient, exogenous attention, showing that in the presence of a well timed spatial cue observers rely on noisy evidence lasting longer and with finer spatial configurations.

The subjective visual vertical in patients with pusher behaviour: a pilot study with a psychophysical approach.

Pusher behaviour (PB) reflects some misrepresentation of verticality. However, its neural mechanisms are still unclear. The aim of this pilot study is to assess the perception of the subjective visual vertical in patients with PB using an orientation discrimination task relying on a psychophysical forced-choice procedure. A sample of eight patients with post-stroke hemiplegia, three of whom with (PB+ group) and five without (PB- group) a clear PB, and 10 matched healthy subjects, was selected. All participants were assessed with an orientation discrimination task based on the objective Two-Alternatives Forced Choice (2AFC) procedure, in which observers are forced to report whether an oriented stimulus that was tilted off-vertical by a varying amount (but never vertical) was tilted clockwise or counterclockwise from vertical. Participants' ability in detecting the visual vertical was estimated by calculation of bias and threshold, which represent deviations of the subjective vertical from the physical vertical and the sensitivity to orientation offsets away from vertical, respectively. While there were no significant differences between groups in terms of bias, statistical analysis showed a significantly higher threshold in the PB+ group compared to both PB- and control groups. Results suggest that vertical misrepresentation might be due to the presence, in patients with PB, of a lower signal-to-noise ratio in coding systems. Implications for clinical practice are discussed.

Acoustic cues to visual detection: a classification image study.

A non-informative sound is known to improve contrast detection thresholds for a synchronous visual target (M. Lippert, N. K. Logothetis, & C. Kayser, 2007). We investigated the spatio-temporal characteristics of the mechanisms underlying this crossmodal effect by using a classification image paradigm specifically suited to investigate perceptual templates across both space and time (P. Neri & D. J. Heeger, 2002). A bright bar was embedded in 2D (space-time) dynamic noise and observers were asked to detect its presence in both unimodal (only visual) and bimodal (audio-visual) conditions. Classification image analysis was performed and the 1st and 2nd order kernels were derived. Our results show that the cross-modal facilitation of detection consists in a reduction of activity of the early mechanisms elicited by the onset of the stimulation and not directly involved in the identification of the target. In fact, the sound sharpens the 2nd order kernels (involved in target detection) by suppressing the activation preceding the target, whereas it does not influence the 1st order kernels. These data suggest that the sound affects some non-linear process involved with the detection of a visual stimulus by, decreasing the activity of contrast energy filters temporally uncorrelated with the target, hence reducing temporal uncertainty.

Reward sharpens orientation coding independently of attention.

It has long been known that rewarding improves performance. However it is unclear whether this is due to high level modulations in the output modules of associated neural systems or due to low level mechanisms favoring more "generous" inputs? Some recent studies suggest that primary sensory areas, including V1 and A1, may form part of the circuitry of reward-based modulations, but there is no data indicating whether reward can be dissociated from attention or cross-trial forms of perceptual learning. Here we address this issue with a psychophysical dual task, to control attention, while perceptual performance on oriented targets associated with different levels of reward is assessed by measuring both orientation discrimination thresholds and behavioral tuning functions for tilt values near threshold. We found that reward, at any rate, improved performance. However, higher reward rates showed an improvement of orientation discrimination thresholds by about 50% across conditions and sharpened behavioral tuning functions. Data were unaffected by changing the attentional load and by dissociating the feature of the reward cue from the task-relevant feature. These results suggest that reward may act within the span of a single trial independently of attention by modulating the activity of early sensory stages through a improvement of the signal-to-noise ratio of task-relevant channels.

Alpha waves: a neural signature of visual suppression.

Alpha waves are traditionally considered a passive consequence of the lack of stimulation of sensory areas. However, recent results have challenged this view by showing a modulation of alpha activity in cortical areas representing unattended information during active tasks. These data have led us to think that alpha waves would support a 'gating function' on sensorial stimulation that actively inhibits unattended information in attentional tasks. Visual suppression occurring during a saccade and blink entails an inhibition of incoming visual information, and it seems to occur at an early processing stage. In this study, we hypothesized that the neural mechanism through which the visual system exerts this inhibition is the active imposition of alpha oscillations in the occipital cortex, which in turn predicts an increment of alpha amplitude during a visual suppression phenomena. We measured visual suppression occurring during short closures of the eyelids, a situation well suited for EEG recordings and stimulated the retinae with an intra-oral light administered through the palate. In the behavioral experiment, detection thresholds were measured with eyes steady open and steady closed, showing a reduction of sensitivity in the latter case. In the EEG recordings performed under identical conditions we found stronger alpha activity with closed eyes. Since the stimulation does not depend on whether the eyes were open or closed, we reasoned that this should be a central effect, probably due to a functional role of alpha oscillation in agreement with the 'gating function' theory.

Reported quality of randomized controlled trials in neglect rehabilitation.

The aim of this study is to assess the reported quality of randomized controlled trials (RCTs) on the effectiveness of neglect rehabilitation using a standardized scale. A search of seven electronic databases was carried out. Selected articles were scored using the PEDro scale and classified as high or low quality study both with the original cut off of 6 and a modified cut off of 5. A linear regression analysis between year of publication and quality rate was used to test whether the quality of the studies improved with time. A total of 18 RCTs were selected. Six articles (33.3%) and 10 articles (55.56%) were classified as having high quality when the original cut off or the modified cut off of the PEDro scale were used, respectively. Analysis shows no time-related changes in PEDro scores. The results show that reported quality is moderate for RCTs in neglect rehabilitation.

Neural correlates of texture and contour integration in children with autism spectrum disorders.

In this study, we have used an electrophysiological paradigm to investigate the neural correlates of the visual integration of local signals across space to generate global percepts in a group of low functioning autistic kids. We have analyzed the amplitude of key harmonics of the Visual Evoked Potentials (VEPs) recorded while participants observed orientation-based texture and contour stimuli, forming coherent global patterns, alternating with visual patterns in which the same number of local elements were randomly oriented in order to loose any globally organized feature. Comparing the results of the clinical sample with those obtained in an age-matched control group, we have observed that in the texture conditions the 1st and 3rd harmonics, containing signature of global form processing (Norcia, Pei, Bonneh, Hou, Sampath, & Pettet, 2005), were present in the control group, while in the experimental group only the 1st harmonic was present. In the Contour condition the 1st harmonic was not present for both groups while the 3rd harmonic was significantly present in the control group but absent in the group with autism. Moreover, the amount of organization required to elicit significant 1st harmonic response in the texture condition was higher in the clinical group. The present results bring additional support to the idea that texture and contour processing are supported by independent mechanisms in normal vision. Autistic vision would thus be characterized by a preserved, perhaps weaker texture mechanism, possibly mediated by feedback interactions between visual areas, and by a disfunction of the mechanism supporting contour processing, possibly mediated by long-range intra-cortical connections. Within this framework, the residual ability to detect contours shown in psychophysical studies could be due to the contribution of the texture mechanism to contour processing.

Search superiority in autism within, but not outside the crowding regime.

Visual cognition of observers with autism spectrum disorder (ASD) seems to show an unbalance between the complementary functions of integration and segregation. This study uses visual search and crowding paradigms to probe the relative ability of children with autism, compared to normal developments children, to extract individual targets from cluttered backgrounds both within and outside the crowding regime. The data show that standard search follows the same pattern in the ASD and control groups with a strong effect of the set size that is substantially weakened by cueing the target location with a synchronous spatial cue. On the other hand, the crowding effect of eight flankers surrounding a small peripheral target is virtually absent in the clinical sample, indicating a superior ability to segregate cluttered visual items. This data, along with evidence of an impairment to the neural system for binding contours in ASD, bring additional support to the general idea of a shift of the trade-off between integration and segregation toward the latter. More specifically, they show that when discriminability is balanced across conditions, an advantage in odd-man out tasks is evident in ASD observers only within the crowding regime, when binding mechanism might get compulsorily triggered in normal observers.

Pooling and segmenting motion signals.

Humans are extremely sensitive to visual motion, largely because local motion signals can be integrated over a large spatial region. On the other hand, summation is often not advantageous, for example when segmenting a moving stimulus against a stationary or oppositely moving background. In this study we show that the spatial extent of motion integration is not compulsory, but is subject to voluntary attentional control. Measurements of motion coherence sensitivity with summation and search paradigms showed that human observers can combine motion signals from cued regions or patches in an optimal manner, even when the regions are quite distinct and remote from each other. Further measurements of contrast sensitivity reinforce previous studies showing that motion integration is preceded by a local analysis akin to contrast thresholding (or intrinsic uncertainty). The results were well modelled by two standard signal-detection-theory models.

Non-linear integration of crowded orientation signals.

Crowding of oriented signals has been explained as linear, compulsory averaging of the signals from target and flankers [Parkes, L., Lund, J., Angelucci, A., Solomon, J. A., & Morgan, M. (2001). Compulsory averaging of crowded orientation signals in human vision. Nature Neuroscience, 4(7), 739-744]. On the other hand, a comparable search task with sparse stimuli is well modeled by a 'Signed-Max' rule that integrates non-linearly local tilt estimates [Baldassi, S., & Verghese, P. (2002). Comparing integration rules in visual search. Journal of Vision, 2(8), 559-570], as reflected by the bimodality of the distributions of reported tilts in a magnitude matching task [Baldassi, S., Megna, N., & Burr, D. C. (2006). Visual clutter causes high-magnitude errors. PLoS Biology, 4(3), e56]. This study compares the two models in the context of crowding by using a magnitude matching task, to measure distributions of perceived target angles and a localization task, to probe the degree of access to local information. Response distributions were bimodal, implying uncertainty, only in the presence of abutting flankers. Localization of the target is relatively preserved but it quantitatively falls in between the predictions of the two models, possibly suggesting local averaging followed by a max operation. This challenges the notion of global averaging and suggests some conscious access to local orientation estimates.

Visual clutter causes high-magnitude errors.

Perceptual decisions are often made in cluttered environments, where a target may be confounded with competing "distractor" stimuli. Although many studies and theoretical treatments have highlighted the effect of distractors on performance, it remains unclear how they affect the quality of perceptual decisions. Here we show that perceptual clutter leads not only to an increase in judgment errors, but also to an increase in perceived signal strength and decision confidence on erroneous trials. Observers reported simultaneously the direction and magnitude of the tilt of a target grating presented either alone, or together with vertical distractor stimuli. When presented in isolation, observers perceived isolated targets as only slightly tilted on error trials, and had little confidence in their decision. When the target was embedded in distractors, however, they perceived it to be strongly tilted on error trials, and had high confidence of their (erroneous) decisions. The results are well explained by assuming that the observers' internal representation of stimulus orientation arises from a nonlinear combination of the outputs of independent noise-perturbed front-end detectors. The implication that erroneous perceptual decisions in cluttered environments are made with high confidence has many potential practical consequences, and may be extendable to decision-making in general.

Attention to locations and features: different top-down modulation of detector weights.

It is well known that attention improves the visibility of a target. In this study, we examined the effect of attention on the selectivity profile for a target. We used a masking technique to measure the tuning function for detecting a target while cueing either its orientation or its location. In the presence of an orientation mask, uncued thresholds were maximally elevated with a parallel mask and decreased with increasing mask orientation from the target. The presence of a cue reduced the masking effect but the shape of the function was cue-specific: The orientation cue consistently improved thresholds at the target orientation, whereas the location cue typically improved thresholds at all orientations relative to the function measured in the absence of attention. The selective versus overall increase of sensitivity observed in our study may be due to differences in the weighting of individual detectors that determine the behavioral tuning function in the two cueing conditions.

"Pop-out" of targets modulated in luminance or colour: the effect of intrinsic and extrinsic uncertainty.

Targets defined by attributes such as colour or brightness are said to "pop-out" from a cluttered scene, with little or no dependency on the size of the set to be searched, while search for other attributes can depend strongly on set-size. We measured contrast thresholds for increments and decrements in luminance or colour and show that they increase strongly with set-size (as previously observed for orientation). However, in some conditions, where the potential distractors were not salient visual targets, there was no dependency of set-size at all ("pop-out"). All the data can be modelled by assuming two main sources of uncertainty: the intrinsic uncertainty due to the number of detectors monitored during a specific task and the extrinsic uncertainty introduced by increasing the number of items displayed. The strength of the effect is well explained by a simple signal detection theory "signed-max" model suited for two-tailed tasks [Journal of Vision 2 (8), 559]. The results suggest that "pop-out" is not peculiar to luminance or colour, but may occur in conditions when the intrinsic uncertainty is so high as to saturate the effects of further uncertainty sources.

Comparing integration rules in visual search.

Search performance for a target tilted in a known direction among vertical distractors is well explained by signal detection theory models. Typically these models use a maximum-of-outputs rule (Max rule) to predict search performance. The Max rule bases its decision on the largest response from a set of independent noisy detectors. When the target is tilted in either direction from the reference orientation and the task is to identify the sign of tilt, the loss of performance with set size is much greater than predicted by the Max rule. Here we varied the target tilt and measured psychometric functions for identifying the direction of tilt from vertical. Measurements were made at different set sizes in the presence of various levels of orientation jitter. The orientation jitter was set at multiples of the estimated internal noise, which was invariant across set sizes and measurement techniques. We then compared the data to the predictions of two models: a Summation model that integrates both signal and noise from local detectors and a Signed-Max model that first picks the maxima on both sides of vertical and then chooses the value with the highest absolute deviation from the reference. Although the function relating thresholds to set size had a slope consistent with both the Signed-Max and the Summation models, the shape of individual psychometric functions was in the most crucial conditions better predicted by the Signed-Max model, which chooses the largest tilt while keeping track of the direction of tilt.