Cortical thickness of primary visual cortex correlates with motion deficits in periventricular leukomalacia.

Impairments of visual motion perception and, in particular, of flow motion have been consistently observed in premature and very low birth weight subjects during infancy. Flow motion information is analyzed at various cortical levels along the dorsal pathways, with information mainly provided by primary and early visual cortex (V1, V2 and V3). We investigated the cortical stage of the visual processing that underlies these motion impairments, measuring Grey Matter Volume and Cortical Thickness in 13 children with Periventricular Leukomalacia (PVL). The cortical thickness, but not the grey matter volume of area V1, correlates negatively with motion coherence sensitivity, indicating that the thinner the cortex, the better the performance among the patients. However, we did not find any such association with either the thickness or volume of area MT, MST and areas of the IPS, suggesting damage at the level of primary visual cortex or along the optic radiation.

Supramodal agnosia for oblique mirror orientation in patients with periventricular leukomalacia.

Periventricular leukomalacia (PVL) is characterized by focal necrosis at the level of the periventricular white matter, often observed in preterm infants. PVL is frequently associated with motor impairment and with visual deficits affecting primary stages of visual processes as well as higher visual cognitive abilities. Here we describe six PVL subjects, with normal verbal IQ, showing orientation perception deficits in both the haptic and visual domains. Subjects were asked to compare the orientation of two stimuli presented simultaneously or sequentially, using both a two alternative forced choice (2AFC) orientation-discrimination and a matching procedure. Visual stimuli were oriented gratings or bars or collinear short lines embedded within a random pattern. Haptic stimuli comprised two rotatable wooden sticks. PVL patients performed at chance in discriminating the oblique orientation, both for visual and haptic stimuli. Moreover when asked to reproduce the oblique orientation, they often oriented the stimulus along the symmetric mirror orientation. The deficit generalized to stimuli varying in many low level features, was invariant for spatiotopic object orientation, and also occurred for sequential presentations. The deficit was specific to oblique orientations, and not for horizontal or vertical stimuli. These findings show that PVL can affect a specific network involved with the supramodal perception of mirror symmetry orientation.

Rhythmic modulation of visual contrast discrimination triggered by action.

Recent evidence suggests that ongoing brain oscillations may be instrumental in binding and integrating multisensory signals. In this experiment, we investigated the temporal dynamics of visual-motor integration processes. We show that action modulates sensitivity to visual contrast discrimination in a rhythmic fashion at frequencies of about 5 Hz (in the theta range), for up to 1 s after execution of action. To understand the origin of the oscillations, we measured oscillations in contrast sensitivity at different levels of luminance, which is known to affect the endogenous brain rhythms, boosting the power of alpha-frequencies. We found that the frequency of oscillation in sensitivity increased at low luminance, probably reflecting the shift in mean endogenous brain rhythm towards higher frequencies. Importantly, both at high and at low luminance, contrast discrimination showed a rhythmic motor-induced suppression effect, with the suppression occurring earlier at low luminance. We suggest that oscillations play a key role in sensory-motor integration, and that the motor-induced suppression may reflect the first manifestation of a rhythmic oscillation.

Short-term monocular deprivation alters early components of visual evoked potentials.

KEY POINTS: Short-term monocular deprivation in adult humans produces a perceptual boost of the deprived eye reflecting homeostatic plasticity. Visual evoked potentials (VEPs) to transient stimuli change after 150 min of monocular deprivation in adult humans. The amplitude of the C1 component of the VEP at a latency of about 100 ms increases for the deprived eye and decreases for the non-deprived eye after deprivation, the two effects being highly negatively correlated. Similarly, the evoked alpha rhythm increases after deprivation for the deprived eye and decreases for the non-deprived eye. The data demonstrate that primary visual cortex excitability is altered by a short period of monocular deprivation, reflecting homeostatic plasticity. ABSTRACT: Very little is known about plasticity in the adult visual cortex. In recent years psychophysical studies have shown that short-term monocular deprivation alters visual perception in adult humans. Specifically, after 150 min of monocular deprivation the deprived eye strongly dominates the dynamics of binocular rivalry, reflecting homeostatic plasticity. Here we investigate the neural mechanisms underlying this form of short-term visual cortical plasticity by measuring visual evoked potentials (VEPs) on the scalp of adult humans during monocular stimulation before and after 150 min of monocular deprivation. We found that monocular deprivation had opposite effects on the amplitude of the earliest component of the VEP (C1) for the deprived and non-deprived eye stimulation. C1 amplitude increased (+66%) for the deprived eye, while it decreased (-29%) for the non-deprived eye. Source localization analysis confirmed that the C1 originates in the primary visual cortex. We further report that following monocular deprivation, the amplitude of the peak of the evoked alpha spectrum increased on average by 23% for the deprived eye and decreased on average by 10% for the non-deprived eye, indicating a change in cortical excitability. These results indicate that a brief period of monocular deprivation alters interocular balance in the primary visual cortex of adult humans by both boosting the activity of the deprived eye and reducing the activity of the non-deprived eye. This indicates a high level of residual homeostatic plasticity in the adult human primary visual cortex, probably mediated by a change in cortical excitability.

The visual component to saccadic compression.

Visual objects presented around the time of saccadic eye movements are strongly mislocalized towards the saccadic target, a phenomenon known as "saccadic compression." Here we show that perisaccadic compression is modulated by the presence of a visual saccadic target. When subjects saccaded to the center of the screen with no visible target, perisaccadic localization was more veridical than when tested with a target. Presenting a saccadic target sometime before saccade initiation was sufficient to induce mislocalization. When we systematically varied the onset of the saccade target, we found that it had to be presented around 100 ms before saccade execution to cause strong mislocalization: saccadic targets presented after this time caused progressively less mislocalization. When subjects made a saccade to screen center with a reference object placed at various positions, mislocalization was focused towards the position of the reference object. The results suggest that saccadic compression is a signature of a mechanism attempting to match objects seen before the saccade with those seen after.

Buildup of spatial information over time and across eye-movements.

To interact rapidly and effectively with our environment, our brain needs access to a neural representation of the spatial layout of the external world. However, the construction of such a map poses major challenges, as the images on our retinae depend on where the eyes are looking, and shift each time we move our eyes, head and body to explore the world. Research from many laboratories including our own suggests that the visual system does compute spatial maps that are anchored to real-world coordinates. However, the construction of these maps takes time (up to 500ms) and also attentional resources. We discuss research investigating how retinotopic reference frames are transformed into spatiotopic reference-frames, and how this transformation takes time to complete. These results have implications for theories about visual space coordinates and particularly for the current debate about the existence of spatiotopic representations.

Spatial position information accumulates steadily over time.

One of the more enduring mysteries of neuroscience is how the visual system constructs robust maps of the world that remain stable in the face of frequent eye movements. Here we show that encoding the position of objects in external space is a relatively slow process, building up over hundreds of milliseconds. We display targets to which human subjects saccade after a variable preview duration. As they saccade, the target is displaced leftwards or rightwards, and subjects report the displacement direction. When subjects saccade to targets without delay, sensitivity is poor; but if the target is viewed for 300-500 ms before saccading, sensitivity is similar to that during fixation with a strong visual mask to dampen transients. These results suggest that the poor displacement thresholds usually observed in the "saccadic suppression of displacement" paradigm are a result of the fact that the target has had insufficient time to be encoded in memory, and not a result of the action of special mechanisms conferring saccadic stability. Under more natural conditions, trans-saccadic displacement detection is as good as in fixation, when the displacement transients are masked.

Early interaction between vision and touch during binocular rivalry.

Multisensory integration is known to occur at high neural levels, but there is also growing evidence that cross-modal signals can be integrated at the first stages of sensory processing. We investigated whether touch specifically affected vision during binocular rivalry, a particular type of visual bistability that engages neural competition in early visual cortices. We found that tactile signals interact with visual signals outside of awareness, when the visual stimulus congruent with the tactile one is perceptually suppressed during binocular rivalry and when the interaction is strictly tuned for matched visuo-tactile spatial frequencies. We also found that voluntary action does not play a leading role in mediating the effect, since the interaction was observed also when tactile stimulation was passively delivered to the finger. However, simultaneous presentation of visual and tactile stimuli is necessary to elicit the interaction, and an asynchronous priming touch stimulus is not affecting the onset of rivalry. These results point to a very early cross-modal interaction site, probably V1. By showing that spatial proximity between visual and tactile stimuli is a necessary condition for the interaction, we also suggest that the two sensory spatial maps are aligned according to retinotopic coordinates, corroborating the hypothesis of a very early interaction between visual and tactile signals during binocular rivalry.

Contextual effects in interval-duration judgements in vision, audition and touch.

We examined the effect of temporal context on discrimination of intervals marked by auditory, visual and tactile stimuli. Subjects were asked to compare the duration of the interval immediately preceded by an irrelevant "distractor" stimulus with an interval with no distractor. For short interval durations, the presence of the distractor affected greatly the apparent duration of the test stimulus: short distractors caused the test interval to appear shorter and vice versa. For very short reference durations (≤ 100 ms), the contextual effects were large, changing perceived duration by up to a factor of two. The effect of distractors reduced steadily for longer reference durations, to zero effect for durations greater than 500 ms. We found similar results for intervals defined by visual flashes, auditory tones and brief finger vibrations, all falling to zero effect at 500 ms. Under appropriate conditions, there were strong cross-modal interactions, particularly from audition to vision. We also measured the Weber fractions for duration discrimination and showed that under the conditions of this experiment, Weber fractions decreased steadily with duration, following a square-root law, similarly for all three modalities. The magnitude of the effect of the distractors on apparent duration correlated well with Weber fraction, showing that when duration discrimination was relatively more precise, the context dependency was less. The results were well fit by a simple Bayesian model combining noisy estimates of duration with the action of a resonance-like mechanism that tended to regularize the sound sequence intervals.

Long-term effects of monocular deprivation revealed with binocular rivalry gratings modulated in luminance and in color.

During development, within a specific temporal window called the critical period, the mammalian visual cortex is highly plastic and literally shaped by visual experience; to what extent this extraordinary plasticity is retained in the adult brain is still a debated issue. We tested the residual plastic potential of the adult visual cortex for both achromatic and chromatic vision by measuring binocular rivalry in adult humans following 150 minutes of monocular patching. Paradoxically, monocular deprivation resulted in lengthening of the mean phase duration of both luminance-modulated and equiluminant stimuli for the deprived eye and complementary shortening of nondeprived phase durations, suggesting an initial homeostatic compensation for the lack of information following monocular deprivation. When equiluminant gratings were tested, the effect was measurable for at least 180 minutes after reexposure to binocular vision, compared with 90 minutes for achromatic gratings. Our results suggest that chromatic vision shows a high degree of plasticity, retaining the effect for a duration (180 minutes) longer than that of the deprivation period (150 minutes) and twice as long as that found with achromatic gratings. The results are in line with evidence showing a higher vulnerability of the P pathway to the effects of visual deprivation during development and a slower development of chromatic vision in humans.

Spatio-temporal topography of saccadic overestimation of time.

Rapid eye movements (saccades) induce visual misperceptions. A number of studies in recent years have investigated the spatio-temporal profiles of effects like saccadic suppression or perisaccadic mislocalization and revealed substantial functional similarities. Saccade induced chronostasis describes the subjective overestimation of stimulus duration when the stimulus onset falls within a saccade. In this study we aimed to functionally characterize saccade induced chronostasis in greater detail. Specifically we tested if chronostasis is influenced by or functionally related to saccadic suppression. In a first set of experiments, we measured the perceived duration of visual stimuli presented at different spatial positions as a function of presentation time relative to the saccade. We further compared perceived duration during saccades for isoluminant and luminant stimuli. Finally, we investigated whether or not saccade induced chronostasis is dependent on the execution of a saccade itself. We show that chronostasis occurs across the visual field with a clear spatio-temporal tuning. Furthermore, we report chronostasis during simulated saccades, indicating that spurious retinal motion induced by the saccade is a prime origin of the phenomenon.

Transient spatiotopic integration across saccadic eye movements mediates visual stability.

Eye movements pose major problems to the visual system, because each new saccade changes the mapping of external objects on the retina. It is known that stimuli briefly presented around the time of saccades are systematically mislocalized, whereas continuously visible objects are perceived as spatially stable even when they undergo large transsaccadic displacements. In this study we investigated the relationship between these two phenomena and measured how human subjects perceive the position of pairs of bars briefly displayed around the time of large horizontal saccades. We show that they interact strongly, with the perisaccadic bar being drawn toward the other, dramatically altering the pattern of perisaccadic mislocalization. The interaction field extends over a wide range (200 ms and 20°) and is oriented along the retinotopic trajectory of the saccade-induced motion, suggesting a mechanism that integrates pre- and postsaccadic stimuli at different retinal locations but similar external positions. We show how transient changes in spatial integration mechanisms, which are consistent with the present psychophysical results and with the properties of "remapping cells" reported in the literature, can create transient craniotopy by merging the distinct retinal images of the pre- and postsaccadic fixations to signal a single stable object.

Saccadic compression of symbolic numerical magnitude.

Stimuli flashed briefly around the time of saccadic eye movements are subject to complex distortions: compression of space and time; underestimate of numerosity. Here we show that saccadic distortions extend to abstract quantities, affecting the representation of symbolic numerical magnitude. Subjects consistently underestimated the results of rapidly computed mental additions and subtractions, when the operands were briefly displayed before a saccade. However, the recognition of the number symbols was unimpaired. These results are consistent with the hypothesis of a common, abstract metric encoding magnitude along multiple dimensions. They suggest that a surprising link exists between the preparation of action and the representation of abstract quantities.

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.

Spatiotopic selectivity of adaptation-based compression of event duration.

A. Bruno, I. Ayhan, and A. Johnston (2010) have recently challenged our report of spatiotopic selectivity for adaptation of event time (D. Burr, A. Tozzi, & M. C. Morrone, 2007) and also our claim that retinotopic adaptation of event time depends on perceived speed. To assist the reader judge this issue, we present here a mass of data accumulated in our laboratories over the last few years, all confirming our original conclusions. We also point out that where Bruno et al. made experimental measurements (rather than relying on theoretical reasoning), they too find clearly significant spatiotopically tuned adaptation-based compression of event time but of lower magnitude to ours. We speculate on the reasons for the differences in magnitude.

Underestimation of perceived number at the time of saccades.

Saccadic eye movements produce transient distortions in both space and time. Mounting evidence suggests that space and time perception are linked, and associated with the perception of another important perceptual attribute, numerosity. Here we investigate the effect of saccades on the perceived numerosity of briefly presented arrays of visual elements. We report a systematic underestimation of numerosity for stimuli flashed just before or during saccades, of about 35% of the reference numerosity. The bias is observed only for relatively large arrays of visual elements, in line with the notion that a distinct perceptual mechanism is involved with enumeration of small numerosities in the 'subitizing' range. This study provides further evidence for the notion that space, time and number share common neural representations, all affected by saccades.

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.

Compression of time during smooth pursuit eye movements.

Humans have a clear sense for the passage of time, but while implicit motor timing is quite accurate, explicit timing is prone to distortions particularly during action (Wenke & Haggard, 2009) and saccadic eye movements (Morrone, Ross, & Burr, 2005). Here, we investigated whether perceived duration is also affected by the execution of smooth pursuit eye movements, showing a compression of apparent duration similar to that observed during saccades. To this end, we presented two brief bars that marked intervals between 100 and 300 ms and asked subjects to judge their duration during fixation and pursuit. We found a compression of perceived duration for bars modulated in luminance contrast of about 32% and for bars modulated in chromatic contrast of 14% during pursuit compared to fixation. Interestingly, Weber ratios were similar for fixation and pursuit, if they are expressed as ratio between JND and perceived duration. This compression was constant for pursuit speeds from 7 to 14 deg/s and did not occur for intervals marked by auditory events. These results argue for a modality-specific component in the processing of temporal information.

Temporal auditory capture does not affect the time course of saccadic mislocalization of visual stimuli.

Irrelevant sounds can "capture" visual stimuli to change their apparent timing, a phenomenon sometimes termed "temporal ventriloquism". Here we ask whether this auditory capture can alter the time course of spatial mislocalization of visual stimuli during saccades. We first show that during saccades, sounds affect the apparent timing of visual flashes, even more strongly than during fixation. However, this capture does not affect the dynamics of perisaccadic visual distortions. Sounds presented 50 ms before or after a visual bar (that change perceived timing of the bars by more than 40 ms) had no measurable effect on the time courses of spatial mislocalization of the bars, in four subjects. Control studies showed that with barely visible, low-contrast stimuli, leading, but not trailing, sounds can have a small effect on mislocalization, most likely attributable to attentional effects rather than auditory capture. These findings support previous studies showing that integration of multisensory information occurs at a relatively late stage of sensory processing, after visual representations have undergone the distortions induced by saccades.