A model for the neuronal implementation of selective visual attention based on temporal correlation among neurons

We propose a model for the neuronal implementation of selective visual attention based on temporal correlation among groups of neurons. Neurons in primary visual cortex respond to visual stimuli with a Poisson distributed spike train with an appropriate, stimulus-dependent mean firing rate. The spike trains of neurons whose receptive fields donot overlap with the focus of attention are distributed according to homogeneous (time-independent) Poisson process with no correlation between action potentials of different neurons. In contrast, spike trains of neurons with receptive fields within the focus of attention are distributed according to non-homogeneous (time-dependent) Poisson processes. Since the short-term average spike rates of all neurons with receptive fields in the focus of attention covary, correlations between these spike trains are introduced which are detected by inhibitory interneurons in V4. These cells, modeled as modified integrate-and-fire neurons, function as coincidence detectors and suppress the response of V4 cells associated with non-attended visual stimuli. The model reproduces quantitatively experimental data obtained in cortical area V4 of monkey by Moran and Desimone (1985).     

Visual cortex: the continuing puzzle of area V2

Surprisingly little is known about the role of V2 in visual processing. A recent study found that the responses of V2 neurons to pairs of angled lines could be predicted from their responses to the individual line components. A simple analysis shows how these neurons may simply sum the responses from one or more orientation selective V1 neurons.     

Suppression without inhibition in visual cortex

Neurons in primary visual cortex (V1) are thought to receive inhibition from other V1 neurons selective for a variety of orientations. Evidence for this inhibition is commonly found in cross-orientation suppression: responses of a V1 neuron to optimally oriented bars are suppressed by superimposed mask bars of different orientation. We show, however, that suppression is unlikely to result from intracortical inhibition. First, suppression can be obtained with masks drifting too rapidly to elicit much of a response in cortex. Second, suppression is immune to hyperpolarization (through visual adaptation) of cortical neurons responding to the mask. Signals mediating suppression might originate in thalamus, rather than in cortex. Thalamic neurons exhibit some suppression; additional suppression might arise from depression at thalamocortical synapses. The mechanisms of suppression are subcortical and possibly include the very first synapse into cortex.     

Coding of stimulus movement parameters in cat visual system

The principal component analysis of matrices composed of spike numbers generated by visual neurons of cats in response to motion of simple and complex stimuli revealed vector encoding. Responses of detectors of moving dot direction and detectors of oblique line orientation are encoded independently in V1 and V2 cortices by excitation of two cardinal neurons. Each pair of these neurons generates sine and cosine functions. Responses of detectors in the association cortex selective to specific orientation of moving stripes depend on the activity of four cardinal neurons which sum up the excitation incoming from the direction and orientation channels.     

Attention, adaptation, and the motion aftereffect

Activation of the human visual motion area V5/MT was previously thought to be the basis of the motion aftereffect. New findings suggest that previous observations were confounded by attention and arousal, providing evidence that adaptation of directionally selective neurons in area V5/MT represents the fundamental substrate for the motion aftereffect.     

Attentional Stimulus Selection through Selective Synchronization between Monkey Visual Areas

A central motif in neuronal networks is convergence, linking several input neurons to one target neuron. In visual cortex, convergence renders target neurons responsive to complex stimuli. Yet, convergence typically sends multiple stimuli to a target, and the behaviorally relevant stimulus must be selected. We used two stimuli, activating separate electrocorticographic V1 sites, and both activating an electrocorticographic V4 site equally strongly. When one of those stimuli activated one V1 site, it gamma synchronized (60-80?Hz) to V4. When the two stimuli activated two V1 sites, primarily the relevant one gamma synchronized to V4. Frequency bands of gamma activities showed substantial overlap containing the band of interareal coherence. The relevant V1 site had its gamma peak frequency 2-3?Hz higher than the irrelevant V1 site and 4-6?Hz higher than V4. Gamma-mediated interareal influences were predominantly directed from V1 to V4. We propose that selective synchronization renders relevant input effective, thereby modulating effective connectivity.     

The effect of gaze angle and fixation distance on the responses of neurons in V1, V2, and V4.

What we see depends on where we look. This paper characterizes the modulatory effects of point of regard in three-dimensional space on responsiveness of visual cortical neurons in areas V1, V2, and V4. Such modulatory effects are both common, affecting 85% of cells, and strong, frequently producing changes of mean firing rate by a factor of 10. The prevalence of neurons in area V4 showing a preference for near distances may be indicative of the involvement of this area in close scrutiny during object recognition. We propose that eye-position signals can be exploited by visual cortex as classical conditioning stimuli, enabling the perceptual learning of systematic relationships between point of regard and the structure of the visual environment.     

Learning the selectivity of V2 and V4 neurons using non-linear multi-layer wavelet networks

We investigate a non-linear network with two processing stages optimized to reduce the statistical dependencies in natural images. This network serves as a model for the neural information processing in the higher visual areas of primates (visual cortices V2-V4). The resulting population is analyzed with regard to non-linear selectivity and invariance properties. We find units that are very selective with respect to the space spanned by all possible input signals and units that are invariant with respect to certain stimulus classes. In comparison to the measured distribution of selectivity in V2 neurons, the selectivity histogram of the network units shows an even more pronounced tendency towards higher selectivities. A special property of the system is the emergence of non-linear interactions between coefficients from different scales and orientations, which are necessary for the exploitation of higher-order statistical redundancies of natural images. We extend the concept to multi-layer systems and present some simulation results.     

Adaptation of oriented and unoriented color-selective neurons in human visual areas

Primary visual cortex contains at least two distinct populations of color-selective cells: neurons in one have circularly symmetric receptive fields and respond best to reddish and greenish light, while neurons in another have oriented receptive fields and a variety of color preferences. The relative prevalence and perceptual roles of the two kinds of neurons remain controversial, however. We used fMRI and a selective adaptation technique to measure responses attributable to these two populations. The technique revealed evidence of adaptation in both populations and indicated that they each produced strong signals in V1 and other human visual areas. The activity of both sets of neurons was also reflected in color appearance measurements made with the same stimuli. Thus, both oriented and unoriented color-selective cells in V1 are important components of the neural pathways that underlie perception of color.     

The effect of learning on the function of monkey extrastriate visual cortex

One of the most remarkable capabilities of the adult brain is its ability to learn and continuously adapt to an ever-changing environment. While many studies have documented how learning improves the perception and identification of visual stimuli, relatively little is known about how it modifies the underlying neural mechanisms. We trained monkeys to identify natural images that were degraded by interpolation with visual noise. We found that learning led to an improvement in monkeys' ability to identify these indeterminate visual stimuli. We link this behavioral improvement to a learning-dependent increase in the amount of information communicated by V4 neurons. This increase was mediated by a specific enhancement in neural activity. Our results reveal a mechanism by which learning increases the amount of information that V4 neurons are able to extract from the visual environment. This suggests that V4 plays a key role in resolving indeterminate visual inputs by coordinated interaction between bottom-up and top-down processing streams.     

Specialized circuits from primary visual cortex to V2 and area MT

Primary visual cortex recombines inputs from magnocellular (M) and parvocellular (P) streams to create functionally specialized outputs. Understanding these input-output relationships is complicated by the fact that layer 4B, which provides outputs to dorsal visual areas, contains multiple cell types. Using a modified rabies virus that expresses green fluorescent protein, we show that layer 4B neurons projecting to MT are a majority spiny stellate, whereas those projecting to V2 are overwhelmingly pyramidal. Regardless of cell type, MT-projecting neurons have larger cell bodies, more dendritic length, and are deeper within layer 4B. Furthermore, MT-projecting pyramidal neurons are located preferentially underneath cytochrome oxidase blobs, indicating that MT-projecting neurons of both types restrict their dendrites to M-recipient zones. We conclude that MT-projecting layer 4B neurons are specialized for the fast transmission of information from the M pathway, while V2-projecting neurons are likely to mediate slower computations involving mixed M and P signals.     

The distribution and morphology of calbindin D28K- and calretinin-immunoreactive neurons in the visual cortex of mouse

We studied the distribution and morphology of calbindin D28K- and calretinin-immunoreactive (IR) neurons in the mouse visual cortex with immunocytochemistry. Most of the calbindin D28K-IR neurons were located in layers II/III and V, while calretinin-IR neurons were predominantly located in layers II/III. The labeled neurons showed variations in morphology. The majority of the calbindin D28K-IR neurons were stellate and round or oval cells with multipolar dendrites. The majority of calretinin-IR neurons were vertical fusiform cells with long processes traveling perpendicular to the pial surface. In the mouse visual cortex, 20.2% of calbindin D28K-IR neurons contained calretinin and 27.2% of calretinin-IR neurons contained calbindin D28K. These results indicate that the calcium-binding proteins, calbindin D28K and calretinin are distributed in specific layers and in selective cell types of the mouse visual cortex.     

Gain modulation by nicotine in macaque v1

Acetylcholine is a ubiquitous cortical neuromodulator implicated in cognition. In order to understand the potential for acetylcholine to play a role in visual attention, we studied nicotinic acetylcholine receptor (nAChR) localization and function in area V1 of the macaque. We found nAChRs presynaptically at thalamic synapses onto excitatory, but not inhibitory, neurons in the primary thalamorecipient layer 4c. Furthermore, consistent with the release enhancement suggested by this localization, we discovered that nicotine increases responsiveness and lowers contrast threshold in layer 4c neurons. We also found that nAChRs are expressed by GABAergic interneurons in V1 but rarely by pyramidal neurons, and that nicotine suppresses visual responses outside layer 4c. All sensory systems incorporate gain control mechanisms, or processes which dynamically alter input/output relationships. We demonstrate that at the site of thalamic input to visual cortex, the effect of this nAChR-mediated gain is an enhancement of the detection of visual stimuli.     

Specialized color modules in macaque extrastriate cortex

Imaging studies are consistent with the existence of brain regions specialized for color, but electrophysiological studies have produced conflicting results. Here we address the neural basis for color, using targeted single-unit recording in alert macaque monkeys, guided by functional magnetic resonance imaging (fMRI) of the same subjects. Distributed within posterior inferior temporal cortex, a large region encompassing V4, PITd, and posterior TEO that some have proposed functions as a single visual complex, we found color-biased fMRI hotspots that we call "globs," each several millimeters wide. Almost all cells located in globs showed strong luminance-invariant color tuning and some shape selectivity. Cells in different globs represented distinct visual field locations, consistent with the coarse retinotopy of this brain region. Cells in "interglob" regions were not color tuned, but were more strongly shape selective. Neither population was direction selective. These results suggest that color perception is mediated by specialized neurons that are clustered within the extrastriate brain.     

Localization of calcium-binding protein parvalbumin-immunoreactive neurons in mouse and hamster visual cortex

Calcium-binding proteins are thought to play important roles in regulating intracellular calcium in the central nervous system. In the present study, we investigated the distribution and morphology of neurons containing parvalbumin in the visual cortex of mouse and hamster. The calcium-binding proteins were localized using immunocytochemistry. Parvalbumin-immunoreactive neurons were located in all layers except layer I. The highest density of parvalbumin immunoreactivity was found in layer V of both mouse and hamster. The labeled neurons varied in morphology. The majority of the parvalbumin-immunoreactive neurons both in mouse and hamster visual cortex was stellate and round, or oval with multipolar dendrites. These results indicate that the calcium-binding protein parvalbumin is contained in specific layers and in selective cell types of the mouse and hamster visual cortex. The distribution of parvalbumin in the mouse visual cortex is very similar to that of hamster.     

Dynamics of the morpho-quantitative changes in the central and intermediate regions of the visual analyzer during whole body x-irradiation

Morpho-quantitative estimation of changes in IV and V layers of visual cortex and outer geniculate bodies of the brain of guinea pigs subjected to whole-body X-irradiation with a median lethal doses shows that neurons from the visual analyzer sections under study are differently damaged. The morphogenesis of changes is connected with the original variations in the quantitative structure of a neuron-glial complex and a selective alterative and productive reaction of the perivascular glia.     

Stability of cortical responses and the statistics of natural scenes.

The primary visual cortex (V1) of higher mammals contains maps of stimulus features; how these maps influence vision remains unknown. We have examined the functional significance of an asymmetry in the orientation map in cat V1, i.e., the fact that a larger area of V1 is preferentially activated by vertical and horizontal contours than by contours at oblique orientations. Despite the fact that neurons tuned to cardinal and oblique orientations have indistinguishable tuning characteristics, cardinal neurons remain more stable in their response properties after selective perturbation induced by adaptation. Similarly, human observers report different adaptation-induced changes in orientation tuning between cardinal and oblique axes. We suggest that the larger cortical area devoted to cardinal orientations imposes stability on the processing of cardinal contours during visual perception, by retaining invariant cortical responses along cardinal axes.     

基于初级视觉皮层的图像匹配模型

提出一种基于初级视觉皮层的图像匹配模型。该模型只采用方位选择性细胞和皮层内有限范围水平连接等Ｖ１基本单元,它以链码表示的目标轮廓作为知识,允许该知识以时间脉冲的形式控制Ｖ１区内神经细胞的动态活动,使与知识轮廓形状相符合的轮廓内的细胞,逐步进入并维持在兴奋状态,最终实现对视野中特定目标轮廓的提取     

Implications on visual apperception: Energy, duration, structure and synchronization

Although primary visual cortex (V1 or striate) activity per se is not sufficient for visual apperception (normal conscious visual experiences and conscious functions such as detection, discrimination, and recognition), the same is also true for extrastriate visual areas (such as V2, V3, V4/V8/VO, V5/M5/MST, IT, and GF). In the lack of V1 area, visual signals can still reach several extrastriate parts but appear incapable of generating normal conscious visual experiences. It is scarcely emphasized in the scientific literature that conscious perceptions and representations must have also essential energetic conditions. These energetic conditions are achieved by spatiotemporal networks of dynamic mitochondrial distributions inside neurons. However, the highest density of neurons in neocortex (number of neurons per degree of visual angle) devoted to representing the visual field is found in retinotopic V1. It means that the highest mitochondrial (energetic) activity can be achieved in mitochondrial cytochrome oxidase-rich V1 areas. Thus, V1 bear the highest energy allocation for visual representation.In addition, the conscious perceptions also demand structural conditions, presence of adequate duration of information representation, and synchronized neural processes and/or ‘interactive hierarchical structuralism.’ For visual apperception, various visual areas are involved depending on context such as stimulus characteristics such as color, form/shape, motion, and other features. Here, we focus primarily on V1 where specific mitochondrial-rich retinotopic structures are found; we will concisely discuss V2 where smaller riches of these structures are found. We also point out that residual brain states are not fully reflected in active neural patterns after visual perception. Namely, after visual perception, subliminal residual states are not being reflected in passive neural recording techniques, but require active stimulation to be revealed.     

Attention to stimulus features shifts spectral tuning of V4 neurons during natural vision

Previous neurophysiological studies suggest that attention can alter the baseline or gain of neurons in extrastriate visual areas but that it cannot change tuning. This suggests that neurons in visual cortex function as labeled lines whose meaning does not depend on task demands. To test this common assumption, we used a system identification approach to measure spatial frequency and orientation tuning in area V4 during two attentionally demanding visual search tasks, one that required fixation and one that allowed free viewing during search. We found that spatial attention modulates response baseline and gain but does not alter tuning, consistent with previous reports. In contrast, feature-based attention often shifts neuronal tuning. These tuning shifts are inconsistent with the labeled-line model and tend to enhance responses to stimulus features that distinguish the search target. Our data suggest that V4 neurons behave as matched filters that are dynamically tuned to optimize visual search.