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Neuroimaging in human amblyopia.

Functional magnetic resonance imaging (fMRI), positron emission tomography (PET) and magnetoencephalography (MEG) have been the principal neuroimaging tools used to assess the site and nature of cortical deficits in human amblyopia. A review of this growing body of work is presented here with particular reference to various controversial issues, including whether or not the primary visual cortex is dysfunctional, the involvement of higher-order visual areas, neural differences between strabismic and anisometropic amblyopes, and the effects of modern-day drug treatments. We also present our own recent MEG work in which we used the analysis technique of synthetic aperture magnetometry (SAM) to examine the effects of strabismic amblyopia on cortical function. Our results provide evidence that the neuronal assembly associated with form perception in the extrastriate cortex may be dysfunctional in amblyopia, and that the nature of this dysfunction may relate to a change in the normal temporal pattern of neuronal discharges. Based on these results and existing literature, we conclude that a number of cortical areas show reduced levels of activation in amblyopia, including primary and secondary visual areas and regions within the parieto-occipital cortex and ventral temporal cortex.

Amblyopia↗

Layer IVA of rhesus monkey primary visual cortex.

Layer IVA of rhesus monkey striate cortex contains pyramidal cells arranged in distinct groups. Their cell bodies are in a configuration of flat cones, each with an average diameter of 60 microns, and their apical dendrites aggregate into bundles that ascend toward the pial surface. Nissl-stained sections suggest that these pyramidal cell cones have their bases in layer IVB, with their tops extending into layer IVA. The neurons in the cones are readily apparent in MAP2 antibody-stained material, and in cytochrome oxidase-reacted tissue it is evident that the pyramidal cell cones occupy the pale spaces that are surrounded by the darkly reactive honeycomb lattice. This lattice of neuropil around the cones contains some axons and boutons that are immunoreactive for parvalbumin, and it is within the lattice that other investigators have shown afferents from the parvocellular (P)-layers of the dLGN to terminate. Because of this input, it is likely that the pyramidal cell cones of layer IVA are involved with color and form perception. The relationship between the layer IVA cones of neurons and the underlying system of previously described pyramidal cell modules (Peters and Sethares, 1991) is discussed, as well as the possibility that the pyramidal cell cones might represent aggregations of neurons, which receive input from basic sets of P-like afferents originating from color-responsive ganglion cells of the retina, as described by Schein and de Monasterio (1987).

Animals↗

Representation of the perceived 3-D object shape in the human lateral occipital complex.

We used human functional magnetic resonance imaging (fMRI) to test whether the human lateral occipital complex (LOC), an area known to be involved in the analysis of visual shape, represents the perceived 3-D shape of objects or simply their 2-D contours. We employed an fMRI adaptation paradigm, in which repeated presentation of a stimulus results in decreased responses compared to responses to different stimuli. We found adaptation in the LOC for images of objects with the same perceived 3-D shape structure but different 2-D contours that resulted from small rotations of the objects in the frontal plane or in depth. However, no adaptation was observed in the LOC for images of objects that had the same 2-D contours but differed in their perceived 3-D shape; namely, 2-D silhouettes versus 3-D shaded images of objects, or convex versus concave objects. Differences in the fMRI adaptation responses across subregions in the LOC suggest that different neural populations in the LOC may mediate different mechanisms for the processing of object features.

Depth Perception↗

How are three-dimensional objects represented in the brain?

In this report we discuss a variety of psychophysical experiments that explore different aspects of the problem of object recognition and representation in human vision. In all experiments, subjects were presented with realistically rendered images of computer-generated 3D objects, with tight control over stimulus shape, surface properties, illumination, and viewpoint, as well as subjects' prior exposure to the stimulus objects. Contrary to the predictions of the paradigmatic theory of recognition, which holds that object representations are viewpoint invariant, performance in all experiments was consistently viewpoint dependent, was only partially aided by binocular stereo and other depth information, was specific to viewpoints that were familiar, and was systematically disrupted by rotation in depth more than by deforming the 2D images of the stimuli. The emerging concept of multiple-views representation supported by these results is consistent with recently advanced computational theories of recognition based on view interpolation. Moreover, in several simulated experiments employing the same stimuli used in experiments with human subjects, models based on multiple-views representations replicated many of the psychophysical results concerning the observed pattern of human performance.

Cues↗

View-based models of 3D object recognition: invariance to imaging transformations.

This report describes the main features of a view-based model of object recognition. The model does not attempt to account for specific cortical structures; it tries to capture general properties to be expected in a biological architecture for object recognition. The basic module is a regularization network (RBF-like; see Poggio and Girosi, 1989; Poggio, 1990) in which each of the hidden units is broadly tuned to a specific view of the object to be recognized. The network output, which may be largely view independent, is first described in terms of some simple simulations. The following refinements and details of the basic module are then discussed: (1) some of the units may represent only components of views of the object--the optimal stimulus for the unit, its "center," is effectively a complex feature; (2) the units' properties are consistent with the usual description of cortical neurons as tuned to multidimensional optimal stimuli and may be realized in terms of plausible biophysical mechanisms; (3) in learning to recognize new objects, preexisting centers may be used and modified, but also new centers may be created incrementally so as to provide maximal view invariance; (4) modules are part of a hierarchical structure--the output of a network may be used as one of the inputs to another, in this way synthesizing increasingly complex features and templates; (5) in several recognition tasks, in particular at the basic level, a single center using view-invariant features may be sufficient.

Depth Perception↗

Psychophysical and physiological evidence for viewer-centered object representations in the primate.

A key question concerning the perception of 3D objects is the spatial reference frame used by the brain to represent them. The celerity of the recognition process could be explained by the visual system's ability to quickly transform stored models of familiar 3D objects, or by its ability to specify the relationship among viewpoint-invariant features or volumetric primitives that can be used to accomplish a structural description of an image. Alternatively, viewpoint-invariant recognition could be realized by a system endowed with the ability to perform an interpolation between a set of stored 2D templates, created for each experienced viewpoint. In the present study we set out to examine the nature of object representation in the primate in combined psychophysical-electrophysiological experiments. Monkeys were trained to recognize novel objects from a given viewpoint and subsequently were tested for their ability to generalize recognition for views generated by mathematically rotating the objects around any arbitrary axis. The perception of 3D novel objects was found to be a function of the object's retinal projection at the time of the recognition encounter. Recognition became increasingly difficult for the monkeys as the stimulus was rotated away from its familiar attitude. The generalization field for novel wire-like and spheroidal objects extended to about +/- 40 degrees around an experienced viewpoint. When the animals were trained with as few as three views of the object, 120 degrees apart, they could often interpolate recognition for all views resulting from rotations around the same axis. Recordings from inferotemporal cortex during the psychophysical testing showed a number of neurons with remarkable selectivity for individual views of those objects that the monkey had learned to recognize. Plotting the response of neurons as a function of rotation angle revealed systematic view-tuning curves for rotations in depth. A small percentage of the view-selective cells responded strongly for a particular view and its mirror-symmetrical view. For some of the tested objects, different neurons were found to be tuned to different views of the same object; the peaks of the view-tuning curves were 40-50 degrees apart. Neurons were also found that responded to the sight of unfamiliar objects or distractors. Such cells, however, gave nonspecific responses to a variety of other patterns presented while the monkey performed a simple fixation task.

Animals↗

Neural activity during attention shifts between object features.

To investigate the neural mechanisms involved in shifting attention we used positron emission tomography to examine regional cerebral blood flow (rCBF) during a task that demands shifting attention between color and shape. Significant activation was observed in the right dorsal prefrontal cortex and parieto-occipital cortex at all frequencies of attention shifts. The frequency of shifts between categories correlated significantly with rCBF in the rostral part of the supplementary motor area and the left precuneus, whereas the number of successive correct responses correlated with rCBF in the orbitofrontal cortex and the caudate nucleus. This study suggests that several prefrontal regions may participate in the processes of shifting attention in different ways.

Adult↗

The interpretation of structure from motion.

The interpretation of structure from motion is examined from a computional point of view. The question addressed is how the three dimensional structure and motion of objects can be inferred from the two dimensional transformations of their projected images when no three dimensional information is conveyed by the individual projections. The following scheme is proposed: (i) divide the image into groups of four elements each; (ii) test each group for a rigid interpretation; (iii) combine the results obtained in (ii). It is shown that this scheme will correctly decompose scenes containing arbitrary rigid objects in motion, recovering their three dimensional structure and motion. The analysis is based primarily on the "structure from motion" theorem which states that the structure of four non-coplanar points is recoverable from three orthographic projections. The interpretation scheme is extended to cover perspective projections, and its psychological relevance is discussed.

Computers↗

Psychophysical investigations into the neural basis of synaesthesia.

We studied two otherwise normal, synaesthetic subjects who 'saw' a specific colour every time they saw a specific number or letter. We conducted four experiments in order to show that this was a genuine perceptual experience rather than merely a memory association. (i) The synaesthetically induced colours could lead to perceptual grouping, even though the inducing numerals or letters did not. (ii) Synaesthetically induced colours were not experienced if the graphemes were presented peripherally. (iii) Roman numerals were ineffective: the actual number grapheme was required. (iv) If two graphemes were alternated the induced colours were also seen in alternation. However, colours were no longer experienced if the graphemes were alternated at more than 4 Hz. We propose that grapheme colour synaesthesia arises from 'cross-wiring' between the 'colour centre' (area V4 or V8) and the 'number area', both of which lie in the fusiform gyrus. We also suggest a similar explanation for the representation of metaphors in the brain: hence, the higher incidence of synaesthesia among artists and poets.

Auditory Perception↗

Binocular vision: moving closer to reality.

Vision science is a truly interdisciplinary field of research, lying at the interface between psychology, computer science and neurobiology. It spans a broad range of research questions, from what visual neurons in the brain do, to the design and use of virtual reality technology. Here, I will take one well-researched area of human vision binocular depth perception, as an exemplar of the various ways in which the processes of human perception can be studied: from basic questions about the sensitivity of binocular vision to understanding how having two eyes helps us function in the natural environment.

Animals↗