Richard J. A. van Wezel

Adaptation: from single cells to BOLD signals

Functional magnetic resonance imaging adaptation (fMRIa) is an increasingly popular method that aims to provide insight into the functional properties of subpopulations of neurons within an imaging voxel. The technique relies on the assumption that neural adaptation reduces activity when two successive stimuli activate the same subpopulation but not when they stimulate different subpopulations. Here, we assess the validity of fMRIa by comparing single-cell recordings with functional imaging of orientation, motion and face processing. We find that fMRIa provides novel insight into neural representations in the human brain. However, network responses in general and adaptation in particular are more complex than is often assumed, and an unequivocal interpretation of fMRIa results can be achieved only with great care.

General validity of Levelt's propositions reveals common computational mechanisms for visual rivalry

The mechanisms underlying conscious visual perception are often studied with either binocular rivalry or perceptual rivalry stimuli. Despite existing research into both types of rivalry, it remains unclear to what extent their underlying mechanisms involve common computational rules. Computational models of binocular rivalry mechanisms are generally tested against Levelts four propositions, describing the psychophysical relation between stimulus strength and alternation dynamics in binocular rivalry. Here we use a bistable rotating structure-from-motion sphere, a generally studied form of perceptual rivalry, to demonstrate that Levelts propositions also apply to the alternation dynamics of perceptual rivalry. Importantly, these findings suggest that bistability in structure-from-motion results from active cross-inhibition between neural populations with computational principles similar to those present in binocular rivalry. Thus, although the neural input to the computational mechanism of rivalry may stem from different cortical neurons and different cognitive levels the computational principles just prior to the production of visual awareness appear to be common to the two types of rivalry.

Interactions between Speed and Contrast Tuning in the Middle Temporal Area: Implications for the Neural Code for Speed

A car driving through the fog appears to move more slowly than one driving on a clear and sunny day. In the laboratory, this observation has been confirmed as a pronounced reduction of perceived speed caused by a reduction in contrast. We measured the influence of contrast on cells in the middle temporal area (MT) of the macaque, which has been hypothesized to underlie the perception of speed. The influence of contrast on the responsiveness and speed tuning of these cells was pervasive and highly regular. As expected, most cells responded less at low contrast. More importantly, the preferred speed of most cells shifted to lower speeds at lower contrasts. Moreover, approximately one-third of cells surprisingly responded more strongly to slow low-contrast stimuli than to slow high-contrast stimuli. Current models of speed perception suggest that each MT cell votes for its preferred speed, with a vote determined by its firing rate. We tested a number of these labeled-line models by entering the neural responses we recorded from MT and comparing the predictions of the models with the perceptual reports of human subjects and monkeys. Contrary to the perceptual reports, the labeled-line models predicted that perceived speed should increase when contrast is decreased. We therefore conclude that perceived speed is not based on a labeled-line interpretation of MT cells.

Inhibition of return is not a foraging facilitator in saccadic search and free viewing

The ability to search and scan the environment effectively is a prerequisite for spatial behavior. A longstanding theory proposes that inhibition of previously attended loci (Inhibition of return; IOR) serves to facilitate exploration by increasing the likelihood to inspect new areas instead of returning to locations that have been inspected before. In this eye movement study we tested whether we could find evidence in favor of this hypothesis. Here we report that IOR does occur during search and free viewing, because we found increased fixation times preceding return saccades (eye movements that return to previously fixated locations). Meanwhile we observed no influence of IOR on the search strategy. Rather than the predicted low number we found many return saccades. Therefore, IOR does not serve as a foraging facilitator in saccadic search and free viewing. We hypothesize that IOR is an intrinsic aspect of shifting attention and gaze direction and furthermore that it is not always advantageous to prevent return saccades.

Linking form and motion in the primate brain

Understanding dynamic events entails the integration of information about form and motion that is crucial for fast and successful interactions in complex environments. A striking example of our sensitivity to dynamic information is our ability to recognize animate figures by the way they move and infer motion from still images. Accumulating evidence for form and motion interactions contrasts with the traditional dissociation between shape and motion-related processes in the ventral and dorsal visual pathways. By combining findings from physiology and brain imaging it can be demonstrated that the primate brain converts information about spatiotemporal sequences into meaningful actions through interactions between early and higher visual areas processing form and motion and frontal-parietal circuits involved in the understanding of actions.

Experience-Driven Plasticity in Binocular Vision

Experience-driven neuronal plasticity allows the brain to adapt its functional connectivity to recent sensory input. Here we use binocular rivalry, an experimental paradigm in which conflicting images are presented to the individual eyes, to demonstrate plasticity in the neuronal mechanisms that convert visual information from two separated retinas into single perceptual experiences. Perception during binocular rivalry tended to initially consist of alternations between exclusive representations of monocularly defined images, but upon prolonged exposure, mixture percepts became more prevalent. The completeness of suppression, reflected in the incidence of mixture percepts, plausibly reflects the strength of inhibition that likely plays a role in binocular rivalry. Recovery of exclusivity was possible but required highly specific binocular stimulation. Documenting the prerequisites for these observed changes in perceptual exclusivity, our experiments suggest experience-driven plasticity at interocular inhibitory synapses, driven by the correlated activity (and also the lack thereof) of neurons representing the conflicting stimuli. This form of plasticity is consistent with a previously proposed but largely untested anti-Hebbian learning mechanism for inhibitory synapses in vision. Our results implicate experience-driven plasticity as one governing principle in the neuronal organization of binocular vision.

Delayed Response to Animate Implied Motion in Human Motion Processing Areas

Viewing static photographs of objects in motion evokes higher fMRI activation in the human medial temporal complex (MT+) than looking at similar photographs without this implied motion. As MT+ is traditionally thought to be involved in motion perception (and not in form perception), this finding suggests feedback from object-recognition areas onto MT+. To investigate this hypothesis, we recorded extracranial potentials evoked by the sight of photographs of biological agents with and without implied motion. The difference in potential between responses to pictures with and without implied motion was maximal between 260 and 400 msec after stimulus onset. Source analysis of this difference revealed one bilateral, symmetrical dipole pair in the occipital lobe. This area also showed a response to real motion, but approximately 100 msec earlier than the implied motion response. The longer latency of the implied motion response in comparison to the real motion response is consistent with a feedback projection onto MT+ following object recognition in higher-level temporal areas.

Recovery from adaptation for dynamic and static motion aftereffects: Evidence for two mechanisms

The motion aftereffect (MAE) is an illusory drift of a physically stationary pattern induced by prolonged viewing of a moving pattern. Depending on the nature of the test pattern the MAE can be phenomenally different. This difference in appearance has led to the suggestion that different underlying mechanisms may be responsible and several reports show that this might be the case. Here, we tested whether differences in MAE duration obtained with stationary test patterns and dynamic test patterns can be explained by a single underlying mechanism. We find the results support the existence of (at least) two mechanisms. The two mechanisms show different characteristics: the static MAE (i.e. the MAE tested with a static test pattern) is almost completely stored when the static test is preceded by a dynamic test; in contradistinction, the dynamic MAE is not stored when dynamic testing is preceded by a static test pattern.

Patterns of resting state connectivity in human primary visual cortical areas: A 7T fMRI study

The nature and origin of fMRI resting state fluctuations and connectivity are still not fully known. More detailed knowledge on the relationship between resting state patterns and brain function may help to elucidate this matter. We therefore performed an in depth study of how resting state fluctuations map to the well known architecture of the visual system. We investigated resting state connectivity at both a fine and large scale within and across visual areas V1, V2 and V3 in ten human subjects using a 7Tesla scanner. We found evidence for several coexisting and overlapping connectivity structures at different spatial scales. At the fine-scale level we found enhanced connectivity between the same topographic locations in the fieldmaps of V1, V2 and V3, enhanced connectivity to the contralateral functional homologue, and to a lesser extent enhanced connectivity between iso-eccentric locations within the same visual area. However, by far the largest proportion of the resting state fluctuations occurred within large-scale bilateral networks. These large-scale networks mapped to some extent onto the architecture of the visual system and could thereby obscure fine-scale connectivity. In fact, most of the fine-scale connectivity only became apparent after the large-scale network fluctuations were filtered from the timeseries. We conclude that fMRI resting state fluctuations in the visual cortex may in fact be a composite signal of different overlapping sources. Isolating the different sources could enhance correlations between BOLD and electrophysiological correlates of resting state activity.

The time course of hemispheric differences in categorical and coordinate spatial processing

Spatial relations between objects can be represented either categorically or coordinately. The metric, coordinate representation is associated with predominant right hemisphere activity, while the abstract, qualitative categorical representation is thought to be processed more in the left hemisphere [Kosslyn, S. M. (1987). Seeing and imagining in the cerebral hemispheres: A computational analysis. Psychological Review, 94, 148-175]. This hypothesized lateralization effect has been found in a number of studies, along with indications that specific task demands can be crucial for these outcomes. In the current experiment a new visual half field task was used which explores these hemispheric differences and their time course by means of a match-to-sample design. Within retention intervals that were brief (500 ms), intermediate (2000 ms), or long (5000 ms), the processing of categorical and coordinate representations was studied. In the 500 ms interval, the hemispheric effect suggested by Kosslyn (1987) was found, but in the longer intervals it was absent. This pattern of the lateralization effect is proposed to be caused by the differential effect the retention interval has on coordinate and categorical representations. Coordinate spatial relations appear susceptible to changes in retention interval and decay very quickly over time, congruent with previous findings about accurate location memory. The processing of categorical spatial relations showed less decay and only between 2000 ms and 5000 ms. Qualitative self reports suggest that the decay found for categorical relations might be caused by a switch from a visual to a more verbal memorization strategy.