Romi Nijhawan

Changing objects lead briefly flashed ones

Continuous, predictable events and spontaneous events may coincide in the visual environment. For a continuously moving object, the brain compensates for delays in transmission between a retinal event and neural responses in higher visual areas. Here we show that it similarly compensated for other smoothly changing features. A disk was flashed briefly during the presentation of another disk of continuously changing color, and observers compared the colors of the disks at the moment of flash. We also tested luminance, spatial frequency and pattern entropy; for all features, the continuously changing item led the flashed item in feature space. Thus the visual systems ability to compensate for delays in information about a continuously changing stimulus may extend to all features. We propose a model based on backward masking and priming to explain the phenomenon.

The Role of Attention in Motion Extrapolation: Are Moving Objects ‘Corrected’ or Flashed Objects Attentionally Delayed?

Objects flashed in alignment with moving objects appear to lag behind [Nijhawan, 1994 Nature (London) 370 256–257], Could this ‘flash-lag’ effect be due to attentional delays in bringing flashed items to perceptual awareness [Titchener, 1908/1973 Lectures on the Elementary Psychology of Feeling and Attention first published 1908 (New York: Macmillan); reprinted 1973 (New York: Arno Press)]? We overtly manipulated attentional allocation in three experiments to address the following questions: Is the flash-lag effect affected when attention is (a) focused on a single event in the presence of multiple events, (b) distributed over multiple events, and (c) diverted from the flashed object? To address the first two questions, five rings, moving along a circular path, were presented while observers attentively tracked one or multiple rings under four conditions: the ring in which the disk was flashed was (i) known or (ii) unknown (randomly selected from the set of five); location of the flashed disk was (i) known or (ii) unknown (randomly selected from ten locations), The third question was investigated by using two moving objects in a cost – benefit cueing paradigm, An arrow cued, with 70% or 80% validity, the position of the flashed object, Observers performed two tasks: (a) reacted as quickly as possible to flash onset; (b) reported the flash-lag effect, We obtained a significant and unaltered flash-lag effect under all the attentional conditions we employed, Furthermore, though reaction times were significantly shorter for validly cued flashes, the flash-lag effect remained uninfluenced by cue validity, indicating that quicker responses to validly cued locations may be due to the shortening of post-perceptual delays in motor responses rather than the perceptual facilitation, We conclude that the computations that give rise to the flash-lag effect are independent of attentional deployment.

Grouping Based on Phenomenal similarity of achromatic color

It is widely acknowledged that a precondition for the perception of the world of objects and events is an early process of organization, and it has generally been assumed that such organization is based on the Gestalt laws of grouping. However, the stage at which such grouping occurs, whether early or late, is an empirical question. It is demonstrated in two experiments that grouping by similarity of neutral color is based not on similarity of absolute luminance at the level of the proximal stimulus, but on phenomenal similarity of lightness resulting from the achievement of lightness constancy. An alternative explanation of such grouping based on the equivalence of luminance ratios between elements and background is ruled out by appropriate control conditions.

Analogous Mechanisms Compensate for Neural Delays in the Sensory and the Motor Pathways: Evidence from Motor Flash-Lag

Motor behaviors require animals to coordinate neural activity across different areas within their motor system. In particular, the significant processing delays within the motor system must somehow be compensated for. Internal models of the motor system, in particular the forward model, have emerged as important potential mechanisms for compensation. For motor responses directed at moving visual objects, there is, additionally, a problem of delays within the sensory pathways carrying crucial position information. The visual phenomenon known as the flash-lag effect has led to a motion-extrapolation model for compensation of sensory delays. In the flash-lag effect, observers see a flashed item colocalized with a moving item as lagging behind the moving item. Here, we explore the possibility that the internal forward model and the motion-extrapolation model are analogous mechanisms compensating for neural delays in the motor and the visual system, respectively. In total darkness, observers moved their right hand gripping a rod while a visual flash was presented at various positions in relation to the rod. When the flash was aligned with the rod, observers perceived it in a position lagging behind the instantaneous felt position of the invisible rod. These results suggest that compensation of neural delays for time-varying motor behavior parallels compensation of delays for time-varying visual stimulation.

The Flash-Lag Phenomenon: Object Motion and Eye Movements

An object flashed briefly in a given location, the moment another moving object arrives in the same location, is perceived by observers as lagging behind the moving object (flash-lag effect). Does the flash-lag effect occur if the retinal image of the moving object is rendered stationary by smooth pursuit of the moving object? Does the flash-lag effect occur if the retinal image of a stationary object is caused to move by smooth-pursuit eye movements? A disk was briefly flashed in the center of a moving ring such that the ring center was completely ‘filled’ by the disk. In this display, observers perceived the flashed disk to lag such that it appeared only to partially ‘fill’ the ring center. The ‘unfilled’ portion (perceived void) of the moving ring was seen in the color of the background. With smooth pursuit of the ring, the flash-lag effect was eliminated, and observers saw the flashed disk centered on the moving ring. A strong flash-lag effect was observed when observers smoothly pursued a moving point target past a continuously visible stationary ring. Once again, the flashed disk appeared to only partially fill the center of the continuously visible stationary ring, yielding a vivid ‘perceived void’. These results are discussed in terms of neural delays and their compensation.

Shifts in perceived position of flashed stimuli by illusory object motion

Moving stimuli cause the position of flashed stimuli to appear shifted in the direction of motion (position capture). To examine whether position capture depends on low-level motion interactions or perception of integrated object motion, we employed a slit-view display. Two line-drawn diamonds translated horizontally in opposite directions, one above and one below the fixation cross, either behind an occluding surface with a narrow slit or without occluding surface. When the diamonds were in vertical alignment, two vertical bars were flashed, one in the center of each diamond. In the slit-view condition, the diamonds were visible through a 4-, 2-, or 1-pixel vertical slit; the width of the flashed bars always matched the width of the slit. Even though the horizontal component of physical motion was greatly reduced or absent in the slit-view conditions, observers perceived diamonds moving behind the occluding surface. Furthermore, the position of the flashed bar was captured by the moving diamonds such that each bar appeared shifted in the direction of perceived motion. We conclude that the position capture reported here has a component based on high-level motion processing that is responsible for dynamically integrating object motion and shape.

Space and time in perception and action

What is the instantaneous position of a moving object from the point of view of the observer? How does a tennis player know when and where to place the racket in order to return a 120 mph serve? Does time stop sometimes and go faster at others? Space, time, and motion have played a fundamental role in extending the foundations of nineteenthand twentieth-century physics. Key breakthroughs resulted from scientists who focused not just on measurements based on rulers and clocks, but also on the role of the observer. Research targeted on the observer’s capabilities and limitations raises a promising new approach that is likely to forward our understanding of neuroscience and psychophysics. Space and Time in Perception and Action brings together theory and empirical findings from world-class experts and is written for advanced students and neuroscientists with a particular interest in the psychophysics of space, time, and motion.

Compensating time delays with neural predictions: are predictions sensory or motor?

Neural delays are a general property of computations carried out by neural circuits. Delays are a natural consequence of temporal summation and coding used by the nervous system to integrate information from multiple resources. For adaptive behaviour, however, these delays must be compensated. In order to sense and interact with moving objects, for example, the visual system must predict the future position of the object to compensate for delays. In this paper, we address two critical questions concerning the implementation of the compensation mechanisms in the brain, namely, where does compensation occur and how is it realized. We present evidence showing that compensation can happen in both the motor and sensory systems, and that compensation using ‘diagonal neural pathways’ is a suitable strategy for implementing compensation in the visual system. In this strategy, neural signals in the early stage of information processing are sent to the future cortical positions that correspond to the distance the object will travel in the period of transmission delay. We propose a computational model to elucidate this using the retinal visual information pathway.

The Perceived Position of Moving Objects: Transcranial Magnetic Stimulation of Area MT+ Reduces the Flash-Lag Effect

How does the visual system assign the perceived position of a moving object? This question is surprisingly complex, since sluggish responses of photoreceptors and transmission delays along the visual pathway mean that visual cortex does not have immediate information about a moving objects position. In the flash-lag effect (FLE), a moving object is perceived ahead of an aligned flash. Psychophysical work on this illusion has inspired models for visual localization of moving objects. However, little is known about the underlying neural mechanisms. Here, we investigated the role of neural activity in areas MT+ and V1/V2 in localizing moving objects. Using short trains of repetitive Transcranial Magnetic Stimulation (TMS) or single pulses at different time points, we measured the influence of TMS on the perceived location of a moving object. We found that TMS delivered to MT+ significantly reduced the FLE; single pulse timings revealed a broad temporal tuning with maximum effect for TMS pulses, 200 ms after the flash. Stimulation of V1/V2 did not significantly influence perceived position. Our results demonstrate that area MT+ contributes to the perceptual localization of moving objects and is involved in the integration of position information over a long time window.

Going, going, gone: Localizing abrupt offsets of moving objects

When a moving object abruptly disappears, this profoundly influences its localization by the visual system. In Experiment 1, 2 aligned objects moved across the screen, and 1 of them abruptly disappeared. Observers reported seeing the objects misaligned at the time of the offset, with the continuing object leading. Experiment 2 showed that the perceived forward displacement of the moving object depended on speed and that offsets were localized accurately. Two competing representations of position for moving objects are proposed: 1 based on a spatially extrapolated internal model, and the other based on transient signals elicited by sudden changes in the object trajectory that can correct the forward-shifted position. Experiment 3 measured forward displacements for moving objects that disappeared only for a short time or abruptly reduced contrast by various amounts. Manipulating the relative strength of the 2 position representations in this way resulted in intermediate positions being perceived, with weaker motion signals or stronger transients leading to less forward displacement. This 2-process mechanism is advantageous because it uses available information about object position to maximally reduce spatio-temporal localization errors.