Neuronal latencies and the position of moving objects.

Neuronal latencies delay the registration of the visual signal from a moving object. By the time the visual input reaches brain structures that encode its position, the object has already moved on. Do we perceive the position of a moving object with a delay because of neuronal latencies? Or is there a brain mechanism that compensates for latencies such that we perceive the true position of a moving object in real time? This question has been intensely debated in the context of the flash-lag illusion: a moving object and an object flashed in alignment with it appear to occupy different positions. The moving object is seen ahead of the flash. Does this show that the visual system extrapolates the position of moving objects into the future to compensate for neuronal latencies? Alternative accounts propose that it simply shows that moving and flashed objects are processed with different delays, or that it reflects temporal integration in brain areas that encode position and motion. The flash-lag illusion and the hypotheses put forward to explain it lead to interesting questions about the encoding of position in the brain. Where is the 'where' pathway and how does it work?

The persistence of position.

I describe a signal coined position persistence that stores information about the last seen position of an object. Position persistence is not the same as visible persistence, although some of its properties are similar. The duration of position persistence is such that objects visible briefly always generate a position signal for at least 180 ms. The signal is not affected by the intensity of the object, nor of the background. Position persistence decreases with increasing speed, but does not depend on retinal eccentricity. Finally, the persisting signal is not tightly bound to the object that causes it. The signal contains no information on the colour of the object, whereas shape information may become represented after approximately 100 ms. The existence of this signal is interpreted as a psychophysical signature of the parallel processing of visual information.

A model of the perceived relative positions of moving objects based upon a slow averaging process.

We extend the local energy model of position detection to cope with temporally varying position signals and the perception of relative position. The extension entails two main components. First, a form of persistence for the position signal based on the temporal impulse response function of the visual system. Secondly, we hypothesise that the perceived relative position of two objects is determined by a slow average of the difference of the objects' position signals. The model explains why briefly flashed static dots are perceived to lag behind continuously visible moving dots, without the need for a motion extrapolation process [Nijhawan, R. (1994). Nature, 370, 256-257]. The dependence of this illusion on parameters such as the velocity, duration, frequency and number of flashes of the motion trajectories is accurately captured by the model. Furthermore, the model makes two predictions. First, briefly flashed dots on a staircase trajectory should lead dots with a long duration. Secondly, it should be possible to abolish the lag-effect between continuously visible and stroboscopically moving objects by halting the continuously visible dots during the interflash interval of the stroboscopic dots. Both predictions are corroborated in experiments.

Postsaccadic visual references generate presaccadic compression of space.

With every rapid gaze shift (saccade), our eyes experience a different view of the world. Stable perception of visual space requires that points in the new image are associated with corresponding points in the previous image. The brain may use an extraretinal eye position signal to compensate for gaze changes, or, alternatively, exploit the image contents to determine associated locations. Support for a uniform extraretinal signal comes from findings that the apparent position of objects briefly flashed around the time of a saccade is often shifted in the direction of the saccade. This view is challenged, however, by observations that the magnitude and direction of the displacement varies across the visual field. Led by the observation that non-uniform displacements typically occurred in studies conducted in slightly illuminated rooms, here we determine the dependence of perisaccadic mislocalization on the availability of visual spatial references at various times around a saccade. We find that presaccadic compression occurs only if visual references are available immediately after, rather than before or during, the saccade. Our findings indicate that the visual processes of transsaccadic spatial localization use mainly postsaccadic visual information.

Temporal recruitment along the trajectory of moving objects and the perception of position.

The trajectory of a moving object provides information about its velocity, direction and position. This information can be used to enhance the visual system's ability to detect changes in these parameters. We show that the visibility of the trajectory of a moving object influences the perception of its position. This form of temporal recruitment builds up on a long timescale of approximately 500 ms. Temporary occlusion of the trajectory during this time period reduces recruitment, but does not abolish it. Moreover, we found no spatial restrictions on recruitment on the scale of 10 degrees of arc. When the position of objects on trajectories with different degrees of visibility are compared, this recruitment effect causes spatial offsets. This leads to a visual illusion in which the position of moving objects is misperceived.

The position of moving objects.

Moving objects occupy a range of positions during the period of integration of the visual system. Nevertheless, a unique position is usually observed. We investigate how the trajectory of a stimulus influences the position at which the object is seen. It has been shown before that moving objects are perceived ahead of static objects shown at the same place and time. We show here that this perceived position difference builds up over the first 500 ms of a visible trajectory. Discontinuities in the visual input reduce this buildup when the presentation frequency of a stimulus with a duration of 42 ms falls below 16 Hz. We interpret this relative mislocalization in terms of a spatiotemporal-filtering model. This model fits well with the data, given two assumptions. First, the position signal persists even though the objects are no longer visible and, second, the perceived distance is a 500 ms average of the difference of these position signals.

Nitric oxide in cortical map formation.

We analyse the possible role of nitric oxide in the early development of cortical maps. A mathematical analysis shows that with the aid of the diffusing messenger NO, cortical maps can develop without the need for lateral-inhibitory interactions. We derive that the maps that NO can set up depend on the stimulus environment in a way that is similar to lateral-inhibitory models of map formation but that they also have properties that depend specifically on the parameters of the diffusion process. These dependencies have not before been extracted and can be used to test the involvement of NO in cortical map formation.