Lora T. Likova

Predominantly extra-retinotopic cortical response to pattern symmetry

Symmetry along one or more axes is a key property of objects and biological organisms. We report on a bilateral visual region of occipital cortex that responds strongly to the presence of multiple symmetries in the viewed image. The stimuli consisted of random dots organized in fourfold and onefold mirror-symmetric patterns, against random control stimuli. The contrast between symmetric and random patterns produced negligible or inconsistent activation of the primary visual projection area V1 or of other medial occipital projection areas. However, there was strong symmetry-specific activation in extra-retinotopic lateral occipital cortex. The high level of activation in this region of cortex may represent part of a general class of computations that require integration of information across a large span of the visual field.

The specificity of cortical region KO to depth structure

Functional MRI studies have identified a cortical region designated as KO between retinotopic areas V3A/B and motion area V5 in human cortex as particularly responsive to motion-defined or kinetic borders. To determine the response of the KO region to more general aspects of structure, we used stereoscopic depth borders and disparate planes with no borders, together with three stimulus types that evoked no depth percept: luminance borders, line contours and illusory phase borders. Responses to these stimuli in the KO region were compared with the responses in retinotopically defined areas that have been variously associated with disparity processing in neurophysiological and fMRI studies. The strongest responses in the KO region were to stimuli evoking perceived depth structure from either disparity or motion cues, but it showed negligible responses either to luminance-based contour stimuli or to edgeless disparity stimuli. We conclude that the region designated as KO is best regarded as a primary center for the generic representation of depth structure rather than any kind of contour specificity.

Extended Concepts of Occipital Retinotopy

Retinotopic mapping is a key property of organization of occipital cortex, predominantly on the medial surface but increasingly being identified in lateral and ventral regions. The retinotopic organization of early visual areas V1-3 is well established, although anatomical landmarks can help to resolve ambiguities in poorly-defined functional maps. New morphing techniques are now available to define the metric mappings quantitatively within each retinotopic area. In the dorsal occipital regions, there is fair agreement that area V3A should be split into separate V3A and V3B maps, and that beyond them lies a further area, V7. We specify the eccentricity mapping of both V3B and V7 for the first time, showing how the latter is roughly parallel to the meridional mapping and offering formal accounts of such paradoxical behavior. In ventral occipital cortex, we support the analysis of Zeki and Bartels (1) and Wade et al. (2) that V4 maps the full hemifield, and show the existence of two more areas, a ventromedial map of the lower quadrant, emphasizing the upper vertical meridian, and an adjacent area with a dominant foveal representation. In lateral cortex, the motion area defined by a motion localizer shows pronounced retinotopy, particularly in the eccentricity parameter. A dorsolateral map between the motion area and V3B, which represents the lower quadrant with an emphasis the foveal part of the lower vertical meridian, may be a counterpart to the ventromedial map.

Stereomotion processing in the human occipital cortex.

Although a high proportion of the motion selective cells in primate motion area, MT, are disparity-selective, there is no convincing evidence for cells in this area specific to stereomotion-in-depth and the neural basis of stereomotion processing remains obscure. With functional magnetic resonance imaging (fMRI), we identify an occipito-temporal region activated by cyclopean stereomotion-in-depth stimulation, centered anterior to the human motion complex hMT+. This discovery suggests a reconceptualization of the architecture of the motion complex to incorporate the processing of motion in the third dimension.

Occipital network for figure/ground organization

To study the cortical mechanism of figure/ground categorization in the human brain, we employed fMRI and the temporal-asynchrony paradigm. This paradigm is able to eliminate any differential activation for local stimulus features, and thus to identify only global perceptual interactions. Strong segmentation of the image into different spatial configurations was generated solely from temporal asynchronies between zones of homogeneous dynamic noise. The figure/ground configuration was a single geometric figure enclosed in a larger surround region. In a control condition, the figure/ground organization was eliminated by segmenting the noise field into many identical temporal-asynchrony stripes. The manipulation of the type of perceptual organization triggered dramatic reorganization in the cortical activation pattern. The figure/ground configuration generated suppression of the ground representation (limited to early retinotopic visual cortex, V1 and V2) and strong activation in the motion complex hMT+/V5+; conversely, both responses were abolished when the figure/ground organization was eliminated. These results suggest that figure/ground processing is mediated by top-down suppression of the ground representation in the earliest visual areas V1/V2 through a signal arising in the motion complex. We propose a model of a recurrent cortical architecture incorporating suppressive feedback that operates in a topographic manner, forming a figure/ground categorization network distinct from that for “pure” scene segmentation and thus underlying the perceptual organization of dynamic scenes into cognitively relevant components.

Crowding: a neuroanalytic approach.

The mechanisms underlying crowding are analyzed in terms of explicit neural processes mediating its perceptual characteristics as originally described by W. Korte (1923). A full understanding of crowding in letter recognition requires a detailed conceptualization of the process of recognition among large numbers of alternatives. The observed masking properties suggest the operation of recursive inhibition from V3 to V1 as a component of the crowding effect. The plausibility of six accounts of the neural basis of crowding (the template matching, feature integrator, attentional feature conjunction, propositional enumeration, attentional tracking, and relaxation network concepts) is then assessed in relation to the task of encoding the spatial structure of the letter forms. We conclude that the relaxation network approach is the most plausible hypothesis to account for the full-spectrum letter recognition performance.

Analysis of human vergence dynamics

Disparity vergence is commonly viewed as being controlled by at least two mechanisms, an open-loop vergence-specific burst mechanism analogous to the ballistic drive of saccades, and a closed-loop feedback mechanism controlled by the disparity error. We show that human vergence dynamics for disparity jumps of a large textured field have a typical time course consistent with predominant control by the open-loop vergence-specific burst mechanism, although various subgroups of the population show radically different vergence behaviors. Some individuals show markedly slow divergence responses, others slow convergence responses, others slow responses in both vergence directions, implying that the two vergence directions have separate control mechanisms. The faster time courses usually had time-symmetric velocity waveforms implying open-loop burst control, while the slow response waveforms were usually time-asymmetric implying closed-loop feedback control. A further type of behavior seen in a distinct subpopulation was a compound anomalous divergence response consisting of an initial convergence movement followed by a large corrective divergence movement with time courses implying closed-loop feedback control. This analysis of the variety of human vergence responses thus contributes substantially to the understanding of the oculomotor control mechanisms underlying the generation of vergence movements [corrected].

Drawing enhances cross-modal memory plasticity in the human brain: a case study in a totally blind adult

In a memory-guided drawing task under blindfolded conditions, we have recently used functional Magnetic Resonance Imaging (fMRI) to demonstrate that the primary visual cortex (V1) may operate as the visuo-spatial buffer, or “sketchpad,” for working memory. The results implied, however, a modality-independent or amodal form of its operation. In the present study, to validate the role of V1 in non-visual memory, we eliminated not only the visual input but all levels of visual processing by replicating the paradigm in a congenitally blind individual. Our novel Cognitive-Kinesthetic method was used to train this totally blind subject to draw complex images guided solely by tactile memory. Control tasks of tactile exploration and memorization of the image to be drawn, and memory-free scribbling were also included. FMRI was run before training and after training. Remarkably, V1 of this congenitally blind individual, which before training exhibited noisy, immature, and non-specific responses, after training produced full-fledged response time-courses specific to the tactile-memory drawing task. The results reveal the operation of a rapid training-based plasticity mechanism that recruits the resources of V1 in the process of learning to draw. The learning paradigm allowed us to investigate for the first time the evolution of plastic re-assignment in V1 in a congenitally blind subject. These findings are consistent with a non-visual memory involvement of V1, and specifically imply that the observed cortical reorganization can be empowered by the process of learning to draw.

Peak localization of sparsely sampled luminance patterns is based on interpolated 3D surface representation.

Objects in the world are typically defined by contours and local features separated by extended featureless regions. Sparsely sampled profiles were therefore used to evaluate the cues involved in localizing objects defined by such separated features (as opposed to typical Vernier acuity or other line-based localization tasks). Objects, in the form of Gaussian blobs, were defined at the sample positions by luminance cues, binocular disparity cues or both together. Remarkably, the luminance information in the sampled profiles was unable to support localization for objects requiring interpolation when the perceived depth from the luminance cue was cancelled by a disparity cue. Disparity cues, on the other hand, improved localization substantially over that for luminance cues alone. These data indicate that it is only through the interpolated depth representation that the position of the sampled object can be recognized. The dominance of a depth representation in the performance of such tasks shows that the depth information is not just an overlay to the 2D sketch of the positional information, but a core process that must be completed before the position of the object can be recognized.

Drawing in the blind and the sighted as a probe of cortical reorganization

In contrast to other arts, such as music, there is a very little neuroimaging research on visual art and in particular - on drawing. Drawing - from artistic to technical - involves diverse aspects of spatial cognition, precise sensorimotor planning and control as well as a rich set of higher cognitive functions. A new method for learning the drawing skill in the blind that we have developed, and the technological advances of a multisensory MR-compatible drawing system, allowed us to run for the first time a comparative fMRI study on drawing in the blind and the sighted. In each population, we identified widely distributed cortical networks, extending from the occipital and temporal cortices, through the parietal to the frontal lobe. This is the first neuroimaging study of drawing in blind novices, as well as the first study on the learning to draw in either population. We sought to determine the cortical reorganization taking place as a result of learning to draw, despite the lack of visual input to the brains of the blind. Remarkably, we found massive recruitment of the visual cortex on learning to draw, although our subjects had no previous experience, but only a short training with our new drawing method. This finding implies a rapid, learning-based plasticity mechanism. We further proposed that the functional level of the brain reorganization in the blind may still differ from that in the sighted even in areas that overlap between the two populations, such as in the visual cortex. We tested this idea in the framework of saccadic suppression. A methodological innovation allowed us to estimate the retinotopic regions locations in the blind brain. Although the visual cortex of both groups was greatly recruited, only the sighted experienced dramatic suppression in hMT+ and V1, while there was no sign of an analogous process in the blind. This finding has important implications and suggests that the recruitment of the visual cortex in the blind does not assure a full functional parallel.