Paola Binda

Spatiotemporal Distortions of Visual Perception at the Time of Saccades

Both space and time are grossly distorted during saccades. Here we show that the two distortions are strongly linked, and that both could be a consequence of the transient remapping mechanisms that affect visual neurons perisaccadically. We measured perisaccadic spatial and temporal distortions simultaneously by asking subjects to report both the perceived spatial location of a perisaccadic vertical bar (relative to a remembered ruler), and its perceived timing (relative to two sounds straddling the bar). During fixation and well before or after saccades, bars were localized veridically in space and in time. In different epochs of the perisaccadic interval, temporal perception was subject to different biases. At about the time of the saccadic onset, bars were temporally mislocalized 50–100 ms later than their actual presentation and spatially mislocalized toward the saccadic target. Importantly, the magnitude of the temporal distortions co-varied with the spatial localization bias and the two phenomena had similar dynamics. Within a brief period about 50 ms before saccadic onset, stimuli were perceived with shorter latencies than at other delays relative to saccadic onset, suggesting that the perceived passage of time transiently inverted its direction. Based on this result we could predict the inversion of perceived temporal order for two briefly flashed visual stimuli. We developed a model that simulates the perisaccadic transient change of neuronal receptive fields predicting well the reported temporal distortions. The key aspects of the model are the dynamics of the “remapped” activity and the use of decoder operators that are optimal during fixation, but are not updated perisaccadically.

Saccades Compress Space, Time, and Number

It has been suggested that space, time and number are represented on a common subjective scale. Saccadic eye movements provide a fascinating test. Saccades compress the perceived magnitude of spatial separations and temporal intervals to approximately half of their true value. The question arises as to whether saccades also compress number. They do, and compression follows a very similar time course for all three attributes: it is maximal at saccadic onset and decreases to veridicality within a window of approximately 50ms. These results reinforce the suggestion of a common perceptual metric, which is probably mediated by the intraparietal cortex; they further suggest that before each saccade the common metric for all three is reset, possibly to pave the way for a fresh analysis of the post-saccadic situation.

Attention to Bright Surfaces Enhances the Pupillary Light Reflex

One longstanding question is how early in the visual system attention exerts its influence. Here we show that an effect of attention can be measured at the earliest possible stage of visual information processing, as a change in the optics of the eye. We tested human subjects and found that covertly attending to bright surfaces results in an enhanced pupillary light reflex (PLR)—the pupillary constriction that occurs in response to light increments. The PLR optimizes the optical quality of the retinal image across illumination conditions, increasing sensitivity by modulating retinal illumination, and improving acuity by reducing spherical aberrations. The attentional modulation of the PLR that we describe constitutes a new mechanism through which vision is affected by attention; we discuss three alternatives for the neural substrates of this effect, including the possibility that attention might act indirectly, via its well established effects in early visual cortex.

Keeping a large-pupilled eye on high-level visual processing

The pupillary light response has long been considered an elementary reflex. However, evidence now shows that it integrates information from such complex phenomena as attention, contextual processing, and imagery. These discoveries make pupillometry a promising tool for an entirely new application: the study of high-level vision.

Fusion of Visual and Auditory Stimuli during Saccades: A Bayesian Explanation for Perisaccadic Distortions

Brief stimuli presented near the onset of saccades are grossly mislocalized in space. In this study, we investigated whether the Bayesian hypothesis of optimal sensory fusion could account for the mislocalization. We required subjects to localize visual, auditory, and audiovisual stimuli at the time of saccades (compared with an earlier presented target). During fixation, vision dominates and spatially “captures” the auditory stimulus (the ventriloquist effect). But for perisaccadic presentations, auditory localization becomes more important, so the mislocalized visual stimulus is seen closer to its veridical position. The precision of the bimodal localization (as measured by localization thresholds or just-noticeable difference) was better than either the visual or acoustic stimulus presented in isolation. Both the perceived position of the bimodal stimuli and the improved precision were well predicted by assuming statistically optimal Bayesian-like combination of visual and auditory signals. Furthermore, the time course of localization was well predicted by the Bayesian approach. We present a detailed model that simulates the time-course data, assuming that perceived position is given by the sum of retinal position and a sluggish noisy eye-position signal, obtained by integrating optimally the output of two populations of neural activity: one centered at the current point of gaze, the other centered at the future point of gaze.

Pupil constrictions to photographs of the sun

The pupil constricts in response to light increments and dilates with light decrements. Here we show that a picture of the sun, introducing a small overall decrease in light level across the field of view, results in a pupillary constriction. Thus, the pictorial representation of a high-luminance object (the sun) can override the normal pupillary dilation elicited by a light decrement. In a series of experiments that control for a variety of factors known to modulate pupil size, we show that the effect (a) does not depend on the retinal position of the images and (b) is modulated by attention. It has long been known that cognitive factors can affect pupil diameter by producing pupillary dilations. Our results indicate that high-level visual analysis (beyond the simple subcortical system mediating the pupillary response to light) can also induce pupillary constriction, with an effect size of about 0.1 mm.

Spatiotemporal profile of peri-saccadic contrast sensitivity

Sensitivity to luminance contrast is reduced just before and during saccades (saccadic suppression), whereas sensitivity to color contrast is unimpaired peri-saccadically and enhanced post-saccadically. The exact spatiotemporal map of these perceptual effects is as yet unknown. Here, we measured detection thresholds for briefly flashed Gaussian blobs modulated in either luminance or chromatic contrast, displayed at a range of eccentricities. Sensitivity to luminance contrast was reduced peri-saccadically by a scaling factor, which was almost constant across retinal space. Saccadic suppression followed a similar time course across all tested eccentricities and was maximal shortly after the saccade onset. Sensitivity to chromatic contrast was enhanced post-saccadically at all tested locations. The enhancement was not specifically linked to the execution of saccades, as it was also observed following a displacement of retinal images comparable to that caused by a saccade. We conclude that luminance and chromatic contrast sensitivities are subject to distinct modulations at the time of saccades, resulting from independent neural processes.

Resting-state retinotopic organization in the absence of retinal input and visual experience

Early visual areas have neuronal receptive fields that form a sampling mosaic of visual space, resulting in a series of retinotopic maps in which the same region of space is represented in multiple visual areas. It is not clear to what extent the development and maintenance of this retinotopic organization in humans depend on retinal waves and/or visual experience. We examined the corticocortical receptive field organization of resting-state BOLD data in normally sighted, early blind, and anophthalmic (in which both eyes fail to develop) individuals and found that resting-state correlations between V1 and V2/V3 were retinotopically organized for all subject groups. These results show that the gross retinotopic pattern of resting-state connectivity across V1-V3 requires neither retinal waves nor visual experience to develop and persist into adulthood. SIGNIFICANCE STATEMENT Evidence from resting-state BOLD data suggests that the connections between early visual areas develop and are maintained even in the absence of retinal waves and visual experience.

Pupil size reflects the focus of feature-based attention

We measured pupil size in adult human subjects while they selectively attended to one of two surfaces, bright and dark, defined by coherently moving dots. The two surfaces were presented at the same location; therefore, subjects could select the cued surface only on the basis of its features. With no luminance change in the stimulus, we find that pupil size was smaller when the bright surface was attended and larger when the dark surface was attended: an effect of feature-based (or surface-based) attention. With the same surfaces at nonoverlapping locations, we find a similar effect of spatial attention. The pupil size modulation cannot be accounted for by differences in eye position and by other variables known to affect pupil size such as task difficulty, accommodation, or the mere anticipation (imagery) of bright/dark stimuli. We conclude that pupil size reflects not just luminance or cognitive state, but the interaction between the two: it reflects which luminance level in the visual scene is relevant for the task at hand.

Renewed Attention on the Pupil Light Reflex

In a recent study, Ebitz and Moore described how subthreshold electrical microstimulation of the macaque frontal eye fields (FEF) modulates the pupillary light reflex. This elegant study suggests that the influence of the FEF and prefrontal cortex on attentional modulation of cortical visual processing extends to the subcortical circuit that mediates a very basic reflex, the pupillary light reflex.