Single units and conscious vision.

Figures that can be seen in more than one way are invaluable tools for the study of the neural basis of visual awareness, because such stimuli permit the dissociation of the neural responses that underlie what we perceive at any given time from those forming the sensory representation of a visual pattern. To study the former type of responses, monkeys were subjected to binocular rivalry, and the response of neurons in a number of different visual areas was studied while the animals reported their alternating percepts by pulling levers. Perception-related modulations of neural activity were found to occur to different extents in different cortical visual areas. The cells that were affected by suppression were almost exclusively binocular, and their proportion was found to increase in the higher processing stages of the visual system. The strongest correlations between neural activity and perception were observed in the visual areas of the temporal lobe. A strikingly large number of neurons in the early visual areas remained active during the perceptual suppression of the stimulus, a finding suggesting that conscious visual perception might be mediated by only a subset of the cells exhibiting stimulus selective responses. These physiological findings, together with a number of recent psychophysical studies, offer a new explanation of the phenomenon of binocular rivalry. Indeed, rivalry has long been considered to be closely linked with binocular fusion and stereopsis, and the sequences of dominance and suppression have been viewed as the result of competition between the two monocular channels. The physiological data presented here are incompatible with this interpretation. Rather than reflecting interocular competition, the rivalry is most probably between the two different central neural representations generated by the dichoptically presented stimuli. The mechanisms of rivalry are probably the same as, or very similar to, those underlying multistable perception in general, and further physiological studies might reveal much about the neural mechanisms of our perceptual organization.     

Perceptual organization of local elements into global shapes in the human visual cortex

The question of how local image features on the retina are integrated into perceived global shapes is central to our understanding of human visual perception. Psychophysical investigations have suggested that the emergence of a coherent visual percept, or a "good-Gestalt", is mediated by the perceptual organization of local features based on their similarity. However, the neural mechanisms that mediate unified shape perception in the human brain remain largely unknown. Using human fMRI, we demonstrate that not only higher occipitotemporal but also early retinotopic areas are involved in the perceptual organization and detection of global shapes. Specifically, these areas showed stronger fMRI responses to global contours consisting of collinear elements than to patterns of randomly oriented local elements. More importantly, decreased detection performance and fMRI activations were observed when misalignment of the contour elements disturbed the perceptual coherence of the contours. However, grouping of the misaligned contour elements by disparity resulted in increased performance and fMRI activations, suggesting that similar neural mechanisms may underlie grouping of local elements to global shapes by different visual features (orientation or disparity). Thus, these findings provide novel evidence for the role of both early feature integration processes and higher stages of visual analysis in coherent visual perception.     

Psychophysiological and neurophysiological features of the organization of visuo-spatial performance in right-handed and left-handed six- to seven-year-old children

The developmental features of individual components of the visual perception and brain functional organization during visuo-spatial activity of different complexity were studied in right-handed and left-handed 6–7-year-old children. The results of psychophysiological testing of their visual perception testify to the underdevelopment of the mechanisms of integrative brain activity. Some specific features of the brain functional organization were revealed in the left-handed children during visuo-spatial performance. More autonomous functioning of the cerebral hemispheres and the duplication of the activation processes in the right and left hemisphere during visuo-spaital performance of different complexity are characteristic of these children. This is probably associated with the involvement of compensatory mechanisms, which enable the performance reliability.     

Linking visual perception with human brain activity.

The past year has seen great advances in the use of functional magnetic resonance imaging (fMRI) to study the functional organization of the human visual cortex, to measure the neuronal correlates of visual perception, and to test computational theories of vision. Activity in particular visual brain areas, as measured with fMRI, has been found to correlate with psychophysical performance, with visual attention, and with subjective perceptual experience.     

A map for horizontal disparity in monkey V2

The perception of visual depth is determined by integration of spatial disparities of inputs from the two eyes. Single cells in visual cortex of monkeys are known to respond to specific binocular disparities; however, little is known about their functional organization. We now show, using intrinsic signal optical imaging and single-unit physiology, that, in the thick stripe compartments of the second visual area (V2), there is a clustered organization of Near cells and Far cells, and moreover, there are topographic maps for Near to Far disparities within V2. Our findings suggest that maps for visual disparity are calculated in V2, and demonstrate parallels in functional organization between the thin, pale, and thick stripes of V2.     

Signal transformation with pairing of sensory stimuli

Rotation of the isolated nervous system of Hermissenda in a caudal orientation causes a synaptic hyperpolarization accompanied by elimination of impulse activity during the steady-state phase of type A but not type B photoreceptors' responses to light. Rotation of the isolated nervous system in a cephalic orientation causes a synaptic depolarization with increase of impulse activity during the steady-state phase of both type A and type B photoreceptors' responses to light. These effects of rotation on photorecptors are explained by known synaptic interactions. Sufficient redundancy is found to be provided by the neural organization of the visual system and its interaction with the statocyst to preserve much of the visual information in spite of signal transformation in specific photorecptors resulting from pairing of rotation with light.     

Bioelectric brain activity during visual perception of verbalized and nonverbalized stimuli in seven-to eight-year-old schoolchildren

Neurophysiological mechanisms of visual perception of verbalized and nonverbalized stimuli have been studied in girls (n = 89) and boys (n = 109) aged seven to eight years. It has been shown that, in children of this age, neurophysiological mechanisms of the visual perception system during the selection of verbalized and nonverbalized stimuli have similar organizations of functional rearrangements and are characterized by the general involvement of cortical areas and by the lack of clear specialization of the cerebral hemispheres. However, the choice of nonverbalized visual stimuli is characterized by a more pronounced generalization of the functional interaction between cortical areas and by the increased spectral power of the EEG’s high-frequency components in anterior associative areas. The increased spatial synchronization in the EEG θ- and β bands in the frontal areas emphasizes the contribution of these structures to the analysis and processing of visual stimuli at this stage of development. Differences in the neurophysiological mechanisms of visual perception in boys and girls have been detected. Spatial and temporal organizations of bioelectric activity (BEA) of the brain in girls performing visual tasks is characterized by a stronger and more varied ipsi- and contralateral statistical relationships in the fronto-occipital direction and displacement of interaction centers to the left hemisphere. In boys perceiving verbalized and nonverbalized visual stimuli, a greater contribution of posterocentral cortical areas has been observed. The detected differences are considered to reflect a greater maturity of identification mechanisms in girls aged seven to eight years as compared to boys.     

A genetic approach to visualization of multisynaptic neural pathways using plant lectin transgene

The wiring patterns among various types of neurons via specific synaptic connections are the basis of functional logic employed by the brain for information processing. This study introduces a powerful method of analyzing the neuronal connectivity patterns by delivering a tracer selectively to specific types of neurons while simultaneously transsynaptically labeling their target neurons. We developed a novel genetic approach introducing cDNA for a plant lectin, wheat germ agglutinin (WGA), as a transgene under the control of specific promoter elements. Using this method, we demonstrate three examples of visualization of specific transsynaptic neural pathways: the mouse cerebellar efferent pathways, the mouse olfactory pathways, and the Drosophila visual pathways. This strategy should greatly facilitate studies on the anatomical and functional organization of the developing and mature nervous system.     

Visibility reflects dynamic changes of effective connectivity between V1 and fusiform cortex

Identifying the neural basis of visibility is central to understanding conscious visual perception. Visibility of basic features such as brightness is often thought to reflect activity in just early visual cortex. But here we show under metacontrast masking that fMRI activity in stimulus-driven areas of early visual cortex did not reflect parametric changes in the visibility of a brightness stimulus. The psychometric visibility function was instead correlated with activity in later visual regions plus parieto-frontal areas, and surprisingly, in representations of the unstimulated stimulus surround for primary visual cortex. Critically, decreased stimulus visibility was associated with a regionally-specific decoupling between early visual cortex and higher visual areas. This provides evidence that dynamic changes in effective connectivity can closely reflect visual perception.     

Playing the electric light orchestra—how electrical stimulation of visual cortex elucidates the neural basis of perception

Vision research has the potential to reveal fundamental mechanisms underlying sensory experience. Causal experimental approaches, such as electrical microstimulation, provide a unique opportunity to test the direct contributions of visual cortical neurons to perception and behaviour. But in spite of their importance, causal methods constitute a minority of the experiments used to investigate the visual cortex to date. We reconsider the function and organization of visual cortex according to results obtained from stimulation techniques, with a special emphasis on electrical stimulation of small groups of cells in awake subjects who can report their visual experience. We compare findings from humans and monkeys, striate and extrastriate cortex, and superficial versus deep cortical layers, and identify a number of revealing gaps in the ‘causal map′ of visual cortex. Integrating results from different methods and species, we provide a critical overview of the ways in which causal approaches have been used to further our understanding of circuitry, plasticity and information integration in visual cortex. Electrical stimulation not only elucidates the contributions of different visual areas to perception, but also contributes to our understanding of neuronal mechanisms underlying memory, attention and decision-making.     

Reentrant Processing in Intuitive Perception

The process of perception requires not only the brain''s receipt of sensory data but also the meaningful organization of that data in relation to the perceptual experience held in memory. Although it typically results in a conscious percept, the process of perception is not fully conscious. Research on the neural substrates of human visual perception has suggested that regions of limbic cortex, including the medial orbital frontal cortex (mOFC), may contribute to intuitive judgments about perceptual events, such as guessing whether an object might be present in a briefly presented fragmented drawing. Examining dense array measures of cortical electrical activity during a modified Waterloo Gestalt Closure Task, results show, as expected, that activity in medial orbital frontal electrical responses (about 250 ms) was associated with intuitive judgments. Activity in the right temporal-parietal-occipital (TPO) region was found to predict mOFC (∼150 ms) activity and, in turn, was subsequently influenced by the mOFC at a later time (∼300 ms). The initial perception of gist or meaning of a visual stimulus in limbic networks may thus yield reentrant input to the visual areas to influence continued development of the percept. Before perception is completed, the initial representation of gist may support intuitive judgments about the ongoing perceptual process.     

Psychophysiological Correlates of Writing and Reading Difficulties in Children of Elementary School Age

A psychophysiological study of the mechanisms underlying writing and reading difficulties at elementary school included assessment of the actual level of speech development, language abilities, intellectual functions, visual memory, visuomotor coordination, and neuropsychological parameters of first- and third- or fourth-grade students. The leading mechanisms of writing and reading difficulties were identified for different stages of skill formation. The role of factors determining the efficiency of learning changed accordingly. From the first to the third (fourth) grade, a decrease was observed in the role of the factors of speech development, formation of motor skills, visual perception, and visual memory, while the voluntary organization of activity, working capacity, and fatigability became the most significant factors. At all stages of teaching writing and reading, the rate of teaching activity must correspond to the functional and adaptive capabilities of schoolchildren.     

Hierarchical organization of macaque and cat cortical sensory systems explored with a novel network processor

Neuroanatomists have described a large number of connections between the various structures of monkey and cat cortical sensory systems. Because of the complexity of the connection data, analysis is required to unravel what principles of organization they imply. To date, analysis of laminar origin and termination connection data to reveal hierarchical relationships between the cortical areas has been the most widely acknowledged approach. We programmed a network processor that searches for optimal hierarchical orderings of cortical areas given known hierarchical constraints and rules for their interpretation. For all cortical systems and all cost functions, the processor found a multitude of equally low-cost hierarchies. Laminar hierarchical constraints that are presently available in the anatomical literature were therefore insufficient to constrain a unique ordering for any of the sensory systems we analysed. Hierarchical orderings of the monkey visual system that have been widely reported, but which were derived by hand, were not among the optimal orderings. All the cortical systems we studied displayed a significant degree of hierarchical organization, and the anatomical constraints from the monkey visual and somato-motor systems were satisfied with very few constraint violations in the optimal hierarchies. The visual and somato-motor systems in that animal were therefore surprisingly strictly hierarchical. Most inconsistencies between the constraints and the hierarchical relationships in the optimal structures for the visual system were related to connections of area FST (fundus of superior temporal sulcus). We found that the hierarchical solutions could be further improved by assuming that FST consists of two areas, which differ in the nature of their projections. Indeed, we found that perfect hierarchical arrangements of the primate visual system, without any violation of anatomical constraints, could be obtained under two reasonable conditions, namely the subdivision of FST into two distinct areas, whose connectivity we predict, and the abolition of at least one of the less reliable rule constraints. Our analyses showed that the future collection of the same type of laminar constraints, or the inclusion of new hierarchical constraints from thalamocortical connections, will not resolve the problem of multiple optimal hierarchical representations for the primate visual system. Further data, however, may help to specify the relative ordering of some more areas. This indeterminacy of the visual hierarchy is in part due to the reported absence of some connections between cortical areas. These absences are consistent with limited cross-talk between differentiated processing streams in the system. Hence, hierarchical representation of the visual system is affected by, and must take into account, other organizational features, such as processing streams.     

A motion direction map in macaque V2

In mammals, the perception of motion starts with direction-selective neurons in the visual cortex. Despite numerous studies in monkey primary and second visual cortex (V1 and V2), there has been no evidence of direction maps in these areas. In the present study, we used optical imaging methods to study the organization of motion response in macaque V1 and V2. In contrast to the findings in other mammals (e.g., cats and ferrets), we found no direction maps in macaque V1. Robust direction maps, however, were found in V2 thick/pale stripes and avoided thin stripes. In many cases direction maps were located within thick stripes and exhibited pinwheel or linear organizations. The presence of motion maps in V2 points to a newfound prominence of V2 in motion processing, for contributing to motion perception in the dorsal pathway and/or for motion cue-dependent form perception in the ventral pathway.     

Cell type specificity of local cortical connections

The function of the cerebral cortex is dependent on the precise organization of the circuits formed by its component neurons. The connections between neurons are not random, but are specific at multiple levels of organization. For example, each cortical area connects to only a selected subset of other areas and within any given area the axonal and dendritic arbors of individual neurons arborize in precise, layer-specific patterns (for review see Felleman & Van Essen, 1991; Callaway, 1998) . In each layer there are dendrites from multiple cell types including cells with somata both within and outside that layer. Anatomical studies have shown that axons arborizing in a particular cortical layer can connect selectively onto dendrites of some cell types in the layer, while avoiding the dendrites of other cell types (e.g. Freund & Gulyas, 1991; Hornung & Celio, 1992; Staiger et al., 1996). These cell type specific connections are, however, difficult to elucidate with anatomical methods, so the frequency of such specificity has remained elusive. Recent experimental methods combining intracellular recording of single neurons with focal neuronal stimulation by uncaging glutamate with light (“photostimulation”) have made the analysis of cell type specific cortical connections more tractable. These studies show that cell type specificity of connections is prevalent in cortex. Here I review photostimulation-based studies investigating the laminar sources of cortical input to distinct cell types in the visual and somatosensory cortices of rats and the primary visual cortex of monkeys.     

The functional impact of mental imagery on conscious perception

Mental imagery has been proposed to contribute to a variety of high-level cognitive functions, including memory encoding and retrieval, navigation, spatial planning, and even social communication and language comprehension. However, it is debated whether mental imagery relies on the same sensory representations as perception, and if so, what functional consequences such an overlap might have on perception itself. We report novel evidence that single instances of imagery can have a pronounced facilitatory influence on subsequent conscious perception. Either seeing or imagining a specific pattern could strongly bias which of two competing stimuli reach awareness during binocular rivalry. Effects of imagery and perception were location and orientation specific, accumulated in strength over time, and survived an intervening visual task lasting several seconds prior to presentation of the rivalry display. Interestingly, effects of imagery differed from those of feature-based attention. The results demonstrate that imagery, in the absence of any incoming visual signals, leads to the formation of a short-term sensory trace that can bias future perception, suggesting a means by which high-level processes that support imagination and memory retrieval may shape low-level sensory representations.     

地下啮齿动物视觉系统的形态结构与机能进化

感觉系统的适应进化机制一直是动物行为学研究的焦点。生活在特殊环境中的动物,其感觉系统在进化过程中表现出的显著差异更是引人注目。由于适应地下黑暗生活环境,地下啮齿动物感觉系统在各个组织水平都表现出进化和退化镶嵌的形态特征,其视觉系统表现得最为突出：视觉器官退化,有关图象分析结构、由视觉诱导产生行为反应的脑区及视觉投射严重退化,有关感受光周期的“非成像” 视觉通路结构高度发达。本文综述了地下啮齿动物视觉系统的结构、功能、进化与发育等方面的研究进展,旨在阐明地下啮齿动物视觉系统的特点,有助于开展地下啮齿动物视觉系统适应进化机制的研究。     

Cerebral Asymmetry of fMRI-BOLD Responses to Visual Stimulation

Hemispheric asymmetry of a wide range of functions is a hallmark of the human brain. The visual system has traditionally been thought of as symmetrically distributed in the brain, but a growing body of evidence has challenged this view. Some highly specific visual tasks have been shown to depend on hemispheric specialization. However, the possible lateralization of cerebral responses to a simple checkerboard visual stimulation has not been a focus of previous studies. To investigate this, we performed two sessions of blood-oxygenation level dependent (BOLD) functional magnetic resonance imaging (fMRI) in 54 healthy subjects during stimulation with a black and white checkerboard visual stimulus. While carefully excluding possible non-physiological causes of left-to-right bias, we compared the activation of the left and the right cerebral hemispheres and related this to grey matter volume, handedness, age, gender, ocular dominance, interocular difference in visual acuity, as well as line-bisection performance. We found a general lateralization of cerebral activation towards the right hemisphere of early visual cortical areas and areas of higher-level visual processing, involved in visuospatial attention, especially in top-down (i.e., goal-oriented) attentional processing. This right hemisphere lateralization was partly, but not completely, explained by an increased grey matter volume in the right hemisphere of the early visual areas. Difference in activation of the superior parietal lobule was correlated with subject age, suggesting a shift towards the left hemisphere with increasing age. Our findings suggest a right-hemispheric dominance of these areas, which could lend support to the generally observed leftward visual attentional bias and to the left hemifield advantage for some visual perception tasks.     

A universal model of esthetic perception based on the sensory coding of natural stimuli

Philosophers have pointed out that there is a close relation between the esthetics of art and the beauty of natural scenes. Supporting this similarity at the experimental level, we have recently shown that visual art and natural scenes share fractal-like, scale-invariant statistical properties. Moreover, evidence from neurophysiological experiments shows that the visual system uses an efficient (sparse) code to process optimally the statistical properties of natural stimuli. In the present work, a hypothetical model of esthetic perception is described that combines both lines of evidence. Specifically, it is proposed that an artist creates a work of art so that it induces a specific resonant state in the visual system. This resonant state is thought to be based on the adaptation of the visual system to natural scenes. The proposed model is universal and predicts that all human beings share the same general concept of esthetic judgment. The model implies that esthetic perception, like the coding of natural stimuli, depends on stimulus form rather than content, depends on higher-order statistics of the stimuli, and is non-intuitive to cognitive introspection. The model accommodates the central tenet of neuroesthetic theory that esthetic perception reflects fundamental functional properties of the nervous system.     

A multilayer neural network model for perception of rotational motion

A multilayer neural nerwork model for the perception of rotational motion has been developed usingReichardt's motion detector array of correlation type, Kohonen's self-organized feature map and Schuster-Wagner's oscillating neural network. It is shown that the unsupervised learning could make the neurons on the second layer of the network tend to be self-organized in a form resembling columnar organization of selective directions in area MT of the primate's visual cortex. The output layer can interpret rotation information and give the directions and velocities of rotational motion. The computer simulation results are in agreement with some psychophysical observations of rotation-al perception. It is demonstrated that the temporal correlation between the oscillating neurons would be powerful for solving the "binding problem" of shear components of rotational motion.