A microcircuit model involving parvalbumin, somatostatin, and vasoactive intestinal polypeptide inhibitory interneurons for the modulation of neuronal oscillation during visual processing.

Various subtypes of inhibitory interneurons contact one another to organize cortical networks. Most cortical inhibitory interneurons express 1 of 3 genes: parvalbumin (PV), somatostatin (SOM), or vasoactive intestinal polypeptide (VIP). This diversity of inhibition allows the flexible regulation of neuronal responses within and between cortical areas. However, the exact roles of these interneuron subtypes and of excitatory pyramidal (Pyr) neurons in regulating neuronal network activity and establishing perception (via interactions between feedforward sensory and feedback attentional signals) remain largely unknown. To explore the regulatory roles of distinct neuronal types in cortical computation, we developed a computational microcircuit model with biologically plausible visual cortex layers 2/3 that combined Pyr neurons and the 3 inhibitory interneuron subtypes to generate network activity. In simulations with our model, inhibitory signals from PV and SOM neurons preferentially induced neuronal firing at gamma (30-80 Hz) and beta (20-30 Hz) frequencies, respectively, in agreement with observed physiological results. Furthermore, our model indicated that rapid inhibition from VIP to SOM subtypes underlies marked attentional modulation for low-gamma frequency (30-50 Hz) in Pyr neuron responses. Our results suggest the distinct but cooperative roles of inhibitory interneuron subtypes in the establishment of visual perception.

Disinhibitory circuit mediated by connections from vasoactive intestinal polypeptide to somatostatin interneurons underlies the paradoxical decrease in spike synchrony with increased border ownership selective neuron firing rate.

The activity of border ownership selective (BOS) neurons in intermediate-level visual areas indicates which side of a contour owns a border relative to its classical receptive field and provides a fundamental component of figure-ground segregation. A physiological study reported that selective attention facilitates the activity of BOS neurons with a consistent border ownership preference, defined as two neurons tuned to respond to the same visual object. However, spike synchrony between this pair is significantly suppressed by selective attention. These neurophysiological findings are derived from a biologically-plausible microcircuit model consisting of spiking neurons including two subtypes of inhibitory interneurons, somatostatin (SOM) and vasoactive intestinal polypeptide (VIP) interneurons, and excitatory BOS model neurons. In our proposed model, BOS neurons and SOM interneurons cooperate and interact with each other. VIP interneurons not only suppress SOM interneuron responses but also are activated by feedback signals mediating selective attention, which leads to disinhibition of BOS neurons when they are directing selective attention toward an object. Our results suggest that disinhibition arising from the synaptic connections from VIP to SOM interneurons plays a critical role in attentional modulation of neurons in intermediate-level visual areas.

Analysis based on neural representation of natural object surfaces to elucidate the mechanisms of a trained AlexNet model.

Analysis and understanding of trained deep neural networks (DNNs) can deepen our understanding of the visual mechanisms involved in primate visual perception. However, due to the limited availability of neural activity data recorded from various cortical areas, the correspondence between the characteristics of artificial and biological neural responses for visually recognizing objects remains unclear at the layer level of DNNs. In the current study, we investigated the relationships between the artificial representations in each layer of a trained AlexNet model (based on a DNN) for object classification and the neural representations in various levels of visual cortices such as the primary visual (V1), intermediate visual (V4), and inferior temporal cortices. Furthermore, we analyzed the profiles of the artificial representations at a single channel level for each layer of the AlexNet model. We found that the artificial representations in the lower-level layers of the trained AlexNet model were strongly correlated with the neural representation in V1, whereas the responses of model neurons in layers at the intermediate and higher-intermediate levels of the trained object classification model exhibited characteristics similar to those of neural activity in V4 neurons. These results suggest that the trained AlexNet model may gradually establish artificial representations for object classification through the hierarchy of its network, in a similar manner to the neural mechanisms by which afferent transmission beginning in the low-level features gradually establishes object recognition as signals progress through the hierarchy of the ventral visual pathway.

Effect of steady-state response versus excitatory/inhibitory balance on spiking synchronization in neural networks with log-normal synaptic weight distribution.

Synchronization of neural activity, especially at the gamma band, contributes to perceptual functions. In several psychiatric disorders, deficits of perceptual functions are reflected in synchronization abnormalities. Plausible cause of this impairment is an alteration in the balance between excitation and inhibition (E/I balance); a disruption in the E/I balance leads to abnormal neural interactions reminiscent of pathological states. Moreover, the local lateral excitatory-excitatory synaptic connections in the cortex exhibit excitatory postsynaptic potentials (EPSPs) that follow a log-normal amplitude distribution. This long-tailed distribution is considered an important factor for the emergence of spatiotemporal neural activity. In this context, we hypothesized that manipulating the EPSP distribution under abnormal E/I balance conditions would provide insights into psychiatric disorders characterized by deficits in perceptual functions, potentially revealing the mechanisms underlying pathological neural behaviors. In this study, we evaluated the synchronization of neural activity with external periodic stimuli in spiking neural networks in cases of both E/I balance and imbalance with or without a long-tailed EPSP amplitude distribution. The results showed that external stimuli of a high frequency lead to a decrease in the degree of synchronization with an increasing ratio of excitatory to inhibitory neurons in the presence, but not in the absence, of high-amplitude EPSPs. This monotonic reduction can be interpreted as an autonomous, strong-EPSP-dependent spiking activity selectively interfering with the responses to external stimuli. This observation is consistent with pathological findings. Thus, our modeling approach has potential to improve the understanding of the steady-state response in both healthy and pathological states.

The Effects of Driving Experience on the P300 Event-Related Potential during the Perception of Traffic Scenes.

The incidence of human-error-related traffic collisions is markedly reduced among drivers who have few years of driving experience compared with those with little driving experience or fewer driving opportunities, even if they have a driver's license. This study analyzes the effect of driving experience on the perception of the traffic scenes through electroencephalograms (EEGs). Primarily, we focused on visual attention during driving, the essential visual function in the visual search and human gaze, and evaluated the P300, which is involved in attention, to explore the effect of driving experience on the visual attention of traffic scenes, not for improving visual ability. In the results, the P300 response was observed in both experienced and beginner drivers when they paid visual attention to the visual target. Furthermore, the latency for the peak amplitude of the P300 response among experienced drivers was markedly faster than that in beginner drivers, suggesting that the P300 latency is a piece of crucial information for driving experience on visual attention.

An Approach for Stabilizing Abnormal Neural Activity in ADHD Using Chaotic Resonance.

Reduced integrity of neural pathways from frontal to sensory cortices has been suggested as a potential neurobiological basis of attention-deficit hyperactivity disorder. Neurofeedback has been widely applied to enhance reduced neural pathways in attention-deficit hyperactivity disorder by repeated training on a daily temporal scale. Clinical and model-based studies have demonstrated that fluctuations in neural activity underpin sustained attention deficits in attention-deficit hyperactivity disorder. These aberrant neural fluctuations may be caused by the chaos-chaos intermittency state in frontal-sensory neural systems. Therefore, shifting the neural state from an aberrant chaos-chaos intermittency state to a normal stable state with an optimal external sensory stimulus, termed chaotic resonance, may be applied in neurofeedback for attention-deficit hyperactivity disorder. In this study, we applied a neurofeedback method based on chaotic resonance induced by "reduced region of orbit" feedback signals in the Baghdadi model for attention-deficit hyperactivity disorder. We evaluated the stabilizing effect of reduced region of orbit feedback and its robustness against noise from errors in estimation of neural activity. The effect of chaotic resonance successfully shifted the abnormal chaos-chaos intermittency of neural activity to the intended stable activity. Additionally, evaluation of the influence of noise due to measurement errors revealed that the efficiency of chaotic resonance induced by reduced region of orbit feedback signals was maintained over a range of certain noise strengths. In conclusion, applying chaotic resonance induced by reduced region of orbit feedback signals to neurofeedback methods may provide a promising treatment option for attention-deficit hyperactivity disorder.

Analysis of spiking synchrony in visual cortex reveals distinct types of top-down modulation signals for spatial and object-based attention.

The activity of a border ownership selective (BOS) neuron indicates where a foreground object is located relative to its (classical) receptive field (RF). A population of BOS neurons thus provides an important component of perceptual grouping, the organization of the visual scene into objects. In previous theoretical work, it has been suggested that this grouping mechanism is implemented by a population of dedicated grouping ("G") cells that integrate the activity of the distributed feature cells representing an object and, by feedback, modulate the same cells, thus making them border ownership selective. The feedback modulation by G cells is thought to also provide the mechanism for object-based attention. A recent modeling study showed that modulatory common feedback, implemented by synapses with N-methyl-D-aspartate (NMDA)-type glutamate receptors, accounts for the experimentally observed synchrony in spike trains of BOS neurons and the shape of cross-correlations between them, including its dependence on the attentional state. However, that study was limited to pairs of BOS neurons with consistent border ownership preferences, defined as two neurons tuned to respond to the same visual object, in which attention decreases synchrony. But attention has also been shown to increase synchrony in neurons with inconsistent border ownership selectivity. Here we extend the computational model from the previous study to fully understand these effects of attention. We postulate the existence of a second type of G-cell that represents spatial attention by modulating the activity of all BOS cells in a spatially defined area. Simulations of this model show that a combination of spatial and object-based mechanisms fully accounts for the observed pattern of synchrony between BOS neurons. Our results suggest that modulatory feedback from G-cells may underlie both spatial and object-based attention.

Long-Tailed Characteristic of Spiking Pattern Alternation Induced by Log-Normal Excitatory Synaptic Distribution.

Studies of structural connectivity at the synaptic level show that in synaptic connections of the cerebral cortex, the excitatory postsynaptic potential (EPSP) in most synapses exhibits sub-mV values, while a small number of synapses exhibit large EPSPs ( >~1.0 [mV]). This means that the distribution of EPSP fits a log-normal distribution. While not restricting structural connectivity, skewed and long-tailed distributions have been widely observed in neural activities, such as the occurrences of spiking rates and the size of a synchronously spiking population. Many studies have been modeled this long-tailed EPSP neural activity distribution; however, its causal factors remain controversial. This study focused on the long-tailed EPSP distributions and interlateral synaptic connections primarily observed in the cortical network structures, thereby having constructed a spiking neural network consistent with these features. Especially, we constructed two coupled modules of spiking neural networks with excitatory and inhibitory neural populations with a log-normal EPSP distribution. We evaluated the spiking activities for different input frequencies and with/without strong synaptic connections. These coupled modules exhibited intermittent intermodule-alternative behavior, given moderate input frequency and the existence of strong synaptic and intermodule connections. Moreover, the power analysis, multiscale entropy analysis, and surrogate data analysis revealed that the long-tailed EPSP distribution and intermodule connections enhanced the complexity of spiking activity at large temporal scales and induced nonlinear dynamics and neural activity that followed the long-tailed distribution.

Correspondence between Monkey Visual Cortices and Layers of a Saliency Map Model Based on a Deep Convolutional Neural Network for Representations of Natural Images.

Attentional selection is a function that allocates the brain's computational resources to the most important part of a visual scene at a specific moment. Saliency map models have been proposed as computational models to predict attentional selection within a spatial location. Recent saliency map models based on deep convolutional neural networks (DCNNs) exhibit the highest performance for predicting the location of attentional selection and human gaze, which reflect overt attention. Trained DCNNs potentially provide insight into the perceptual mechanisms of biological visual systems. However, the relationship between artificial and neural representations used for determining attentional selection and gaze location remains unknown. To understand the mechanism underlying saliency map models based on DCNNs and the neural system of attentional selection, we investigated the correspondence between layers of a DCNN saliency map model and monkey visual areas for natural image representations. We compared the characteristics of the responses in each layer of the model with those of the neural representation in the primary visual (V1), intermediate visual (V4), and inferior temporal (IT) cortices. Regardless of the DCNN layer level, the characteristics of the responses were consistent with that of the neural representation in V1. We found marked peaks of correspondence between V1 and the early level and higher-intermediate-level layers of the model. These results provide insight into the mechanism of the trained DCNN saliency map model and suggest that the neural representations in V1 play an important role in computing the saliency that mediates attentional selection, which supports the V1 saliency hypothesis.

Deterministic characteristics of spontaneous activity detected by multi-fractal analysis in a spiking neural network with long-tailed distributions of synaptic weights.

Cortical neural networks maintain autonomous electrical activity called spontaneous activity that represents the brain's dynamic internal state even in the absence of sensory stimuli. The spatio-temporal complexity of spontaneous activity is strongly related to perceptual, learning, and cognitive brain functions; multi-fractal analysis can be utilized to evaluate the complexity of spontaneous activity. Recent studies have shown that the deterministic dynamic behavior of spontaneous activity especially reflects the topological neural network characteristics and changes of neural network structures. However, it remains unclear whether multi-fractal analysis, recently widely utilized for neural activity, is effective for detecting the complexity of the deterministic dynamic process. To verify this point, we focused on the log-normal distribution of excitatory postsynaptic potentials (EPSPs) to evaluate the multi-fractality of spontaneous activity in a spiking neural network with a log-normal distribution of EPSPs. We found that the spiking activities exhibited multi-fractal characteristics. Moreover, to investigate the presence of a deterministic process in the spiking activity, we conducted a surrogate data analysis against the time-series of spiking activity. The results showed that the spontaneous spiking activity included the deterministic dynamic behavior. Overall, the combination of multi-fractal analysis and surrogate data analysis can detect deterministic complex neural activity. The multi-fractal analysis of neural activity used in this study could be widely utilized for brain modeling and evaluation methods for signals obtained by neuroimaging modalities.

Transition of Neural Activity From the Chaotic Bipolar-Disorder State to the Periodic Healthy State Using External Feedback Signals.

Chronotherapy is a treatment for mood disorders, including major depressive disorder, mania, and bipolar disorder (BD). Neurotransmitters associated with the pathology of mood disorders exhibit circadian rhythms. A functional deficit in the neural circuits related to mood disorders disturbs the circadian rhythm; chronotherapy is an intervention that helps resynchronize the patient's biological clock with the periodic daily cycle, leading to amelioration of symptoms. In previous reports, Hadaeghi et al. proposed a non-linear dynamic model composed of the frontal and sensory cortical neural networks and the hypothalamus to explain the relationship between deficits in neural function in the frontal cortex and the disturbed circadian rhythm/mood transitions in BD (hereinafter referred to as the Hadaeghi model). In this model, neural activity in the frontal and sensory lobes exhibits periodic behavior in the healthy state; while in BD, this neural activity is in a state of chaos-chaos intermittency; this temporal departure from the healthy periodic state disturbs the circadian pacemaker in the hypothalamus. In this study, we propose an intervention based on a feedback method called the "reduced region of orbit" (RRO) method to facilitate the transition of the disturbed frontal cortical neural activity underlying BD to healthy periodic activity. Our simulation was based on the Hadaeghi model. We used an RRO feedback signal based on the return-map structure of the simulated frontal and sensory lobes to induce synchronization with a relatively weak periodic signal corresponding to the healthy condition by applying feedback of appropriate strength. The RRO feedback signal induces chaotic resonance, which facilitates the transition to healthy, periodic frontal neural activity, although this synchronization is restricted to a relatively low frequency of the periodic input signal. Additionally, applying an appropriate strength of the RRO feedback signal lowered the amplitude of the periodic input signal required to induce a synchronous state compared with the periodic signal applied alone. In conclusion, through a chaotic-resonance effect induced by the RRO feedback method, the state of the disturbed frontal neural activity characteristic of BD was transformed into a state close to healthy periodic activity by relatively weak periodic perturbations. Thus, RRO feedback-modulated chronotherapy might be an innovative new type of minimally invasive chronotherapy.

Resonance phenomena controlled by external feedback signals and additive noise in neural systems.

Chaotic resonance is a phenomenon that can replace the fluctuation source in stochastic resonance from additive noise to chaos. We previously developed a method to control the chaotic state for suitably generating chaotic resonance by external feedback even when the external adjustment of chaos is difficult, establishing a method named reduced region of orbit (RRO) feedback. However, a feedback signal was utilized only for dividing the merged attractor. In addition, the signal sensitivity in chaotic resonance induced by feedback signals and that of stochastic resonance by additive noise have not been compared. To merge the separated attractor, we propose a negative strength of the RRO feedback signal in a discrete neural system which is composed of excitatory and inhibitory neurons. We evaluate the features of chaotic resonance and compare it to stochastic resonance. The RRO feedback signal with negative strength can merge the separated attractor and induce chaotic resonance. We also confirm that additive noise induces stochastic resonance through attractor merging. The comparison of these resonance modalities verifies that chaotic resonance provides more applicability than stochastic resonance given its capability to handle attractor separation and merging.

Invariance to low-level features and partial transfer over space in the tilt aftereffects evoked by symmetrical patterns.

The symmetry axis is the midline that divides a pattern into congruent halves, which is physically nonexistent but evokes tilt aftereffect (TAE). To investigate the cortical correspondence of the symmetry axis, we examined the invariance of symmetry-induced TAE with regard to low-level visual features and the spatial transfer of TAE over visual fields. When the adaptation pattern was rotated and changed sequentially with the orientation of the symmetry axis unchanged, the measured TAE decreased only slightly (18%) compared to stationary patterns. This effect persisted when the adaptation and test patterns were presented in different visual fields. These results indicate that the cortical representation of symmetry is generated independently of low-level features and involves higher-level visual areas.

Saliency model based on a neural population for integrating figure direction and organizing Border Ownership.

Attentional selection is a function of the brain that allocates computational resources momentarily to the most important part of a visual scene. Saliency map models have been used to predict the location of attentional selection and gaze. Border Ownership (BO) indicates the direction of the figure with respect to the border. I here propose a biologically plausible saliency model based on neural population for integrating the activities of intermediate-level visual areas with neurons selective to BO. A variety of BO organizations produces a population of model neurons that represent the grouping structure. In the model I propose, the interactions and the population responses of these model neurons underlie the determination of saliency and the accurate prediction of gaze location. I tested 100 patterns for BO organizations and found that the proposed saliency model not only reproduced the characteristics of perceptual organization but also captured object locations in natural images. Furthermore, the saliency model based on the population responses of the BO organization significantly improved the gaze prediction accuracy compared with previous saliency-based models. These results suggest a crucial role for a wide variety of BO organizations and neural population coding to determine saliency mediating attentional selection and to predict gaze location.

Interactions Elicited by the Contradiction Between Figure Direction Discrimination and Figure-Ground Segregation.

Figure-ground (FG) segregation that separates an object from the rest of the image is a fundamental problem in vision science. A majority of neurons in monkey V2 showed the selectivity to border ownership (BO) that indicates which side of a contour owns the border. Although BO could be a precursor of FG segregation, the contribution of BO to FG segregation has not been clarified. Because FG segregation is the perception of the global region that belongs to an object, whereas BO determination provides the local direction of figure (DOF) along a contour, a spatial integration of BO might be expected for the generation of FG. To understand the mechanisms underlying the perception of figural regions, we investigated the interaction between the local BO determination and the global FG segregation through the quantitative analysis of the visual perception and the spatiotemporal characteristics of eye movements. We generated a set of novel stimuli in which translucency induces local DOF along the contour and global FG independently so that DOF and FG could be either consistent or contradictory. The perceptual responses showed better performance in DOF discrimination than FG segregation, supporting distinct mechanisms for the DOF discrimination and the FG segregation. We examined whether the contradiction between DOF and FG modulates the eye movement while participants judged DOF and FG. The duration of the first eye fixation was modulated by the contradiction during FG segregation but not DOF discrimination, suggesting a sequential processing from the BO determination to the FG segregation. These results of human perception and eye fixation provide important clues for understanding the visual processing for FG segregation.

Spike synchrony generated by modulatory common input through NMDA-type synapses.

Common excitatory input to neurons increases their firing rates and the strength of the spike correlation (synchrony) between them. Little is known, however, about the synchronizing effects of modulatory common input. Here, we show that modulatory common input with the slow synaptic kinetics of N-methyl-d-aspartate (NMDA) receptors enhances firing rates and also produces synchrony. Tight synchrony (correlations on the order of milliseconds) always increases with modulatory strength. Unexpectedly, the relationship between strength of modulation and strength of loose synchrony (tens of milliseconds) is not monotonic: The strongest loose synchrony is obtained for intermediate modulatory amplitudes. This finding explains recent neurophysiological results showing that in cortical areas V1 and V2, presumed modulatory top-down input due to contour grouping increases (loose and tight) synchrony but that additional modulatory input due to top-down attention does not change tight synchrony and actually decreases loose synchrony. These neurophysiological findings are understood from our model of integrate-and-fire neurons under the assumption that contour grouping as well as attention lead to additive modulatory common input through NMDA-type synapses. In contrast, circuits with common projections through model α-amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid receptors did not exhibit the paradoxical decrease of synchrony with increased input. Our results suggest that NMDA receptors play a critical role in top-down response modulation in the visual cortex.

Modeling the Time-Course of Responses for the Border Ownership Selectivity Based on the Integration of Feedforward Signals and Visual Cortical Interactions.

Border ownership (BO) indicates which side of a contour owns a border, and it plays a fundamental role in figure-ground segregation. The majority of neurons in V2 and V4 areas of monkeys exhibit BO selectivity. A physiological work reported that the responses of BO-selective cells show a rapid transition when a presented square is flipped along its classical receptive field (CRF) so that the opposite BO is presented, whereas the transition is significantly slower when a square with a clear BO is replaced by an ambiguous edge, e.g., when the square is enlarged greatly. The rapid transition seemed to reflect the influence of feedforward processing on BO selectivity. Herein, we investigated the role of feedforward signals and cortical interactions for time-courses in BO-selective cells by modeling a visual cortical network comprising V1, V2, and posterior parietal (PP) modules. In our computational model, the recurrent pathways among these modules gradually established the visual progress and the BO assignments. Feedforward inputs mainly determined the activities of these modules. Surrounding suppression/facilitation of early-level areas modulates the activities of V2 cells to provide BO signals. Weak feedback signals from the PP module enhanced the contrast gain extracted in V1, which underlies the attentional modulation of BO signals. Model simulations exhibited time-courses depending on the BO ambiguity, which were caused by the integration delay of V1 and V2 cells and the local inhibition therein given the difference in input stimulus. However, our model did not fully explain the characteristics of crucially slow transition: the responses of BO-selective physiological cells indicated the persistent activation several times longer than that of our model after the replacement with the ambiguous edge. Furthermore, the time-course of BO-selective model cells replicated the attentional modulation of response time in human psychophysical experiments. These attentional modulations for time-courses were induced by selective enhancement of early-level features due to interactions between V1 and PP. Our proposed model suggests fundamental roles of surrounding suppression/facilitation based on feedforward inputs as well as the interactions between early and parietal visual areas with respect to the ambiguity dependence of the neural dynamics in intermediate-level vision.

Spatial and feature-based attention in a layered cortical microcircuit model.

Directing attention to the spatial location or the distinguishing feature of a visual object modulates neuronal responses in the visual cortex and the stimulus discriminability of subjects. However, the spatial and feature-based modes of attention differently influence visual processing by changing the tuning properties of neurons. Intriguingly, neurons' tuning curves are modulated similarly across different visual areas under both these modes of attention. Here, we explored the mechanism underlying the effects of these two modes of visual attention on the orientation selectivity of visual cortical neurons. To do this, we developed a layered microcircuit model. This model describes multiple orientation-specific microcircuits sharing their receptive fields and consisting of layers 2/3, 4, 5, and 6. These microcircuits represent a functional grouping of cortical neurons and mutually interact via lateral inhibition and excitatory connections between groups with similar selectivity. The individual microcircuits receive bottom-up visual stimuli and top-down attention in different layers. A crucial assumption of the model is that feature-based attention activates orientation-specific microcircuits for the relevant feature selectively, whereas spatial attention activates all microcircuits homogeneously, irrespective of their orientation selectivity. Consequently, our model simultaneously accounts for the multiplicative scaling of neuronal responses in spatial attention and the additive modulations of orientation tuning curves in feature-based attention, which have been observed widely in various visual cortical areas. Simulations of the model predict contrasting differences between excitatory and inhibitory neurons in the two modes of attentional modulations. Furthermore, the model replicates the modulation of the psychophysical discriminability of visual stimuli in the presence of external noise. Our layered model with a biologically suggested laminar structure describes the basic circuit mechanism underlying the attention-mode specific modulations of neuronal responses and visual perception.

Feature-based attention in early vision for the modulation of figure-ground segregation.

We investigated psychophysically whether feature-based attention modulates the perception of figure-ground (F-G) segregation and, based on the results, we investigated computationally the neural mechanisms underlying attention modulation. In the psychophysical experiments, the attention of participants was drawn to a specific motion direction and they were then asked to judge the side of figure in an ambiguous figure with surfaces consisting of distinct motion directions. The results of these experiments showed that the surface consisting of the attended direction of motion was more frequently observed as figure, with a degree comparable to that of spatial attention (Wagatsuma et al., 2008). These experiments also showed that perception was dependent on the distribution of feature contrast, specifically the motion direction differences. These results led us to hypothesize that feature-based attention functions in a framework similar to that of spatial attention. We proposed a V1-V2 model in which feature-based attention modulates the contrast of low-level feature in V1, and this modulation of contrast changes directly the surround modulation of border-ownership-selective cells in V2; thus, perception of F-G is biased. The model exhibited good agreement with human perception in the magnitude of attention modulation and its invariance among stimuli. These results indicate that early-level features that are modified by feature-based attention alter subsequent processing along afferent pathway, and that such modification could even change the perception of object.

Layer-Dependent Attentional Processing by Top-down Signals in a Visual Cortical Microcircuit Model.

A vast amount of information about the external world continuously flows into the brain, whereas its capacity to process such information is limited. Attention enables the brain to allocate its resources of information processing to selected sensory inputs for reducing its computational load, and effects of attention have been extensively studied in visual information processing. However, how the microcircuit of the visual cortex processes attentional information from higher areas remains largely unknown. Here, we explore the complex interactions between visual inputs and an attentional signal in a computational model of the visual cortical microcircuit. Our model not only successfully accounts for previous experimental observations of attentional effects on visual neuronal responses, but also predicts contrasting differences in the attentional effects of top-down signals between cortical layers: attention to a preferred stimulus of a column enhances neuronal responses of layers 2/3 and 5, the output stations of cortical microcircuits, whereas attention suppresses neuronal responses of layer 4, the input station of cortical microcircuits. We demonstrate that the specific modulation pattern of layer-4 activity, which emerges from inter-laminar synaptic connections, is crucial for a rapid shift of attention to a currently unattended stimulus. Our results suggest that top-down signals act differently on different layers of the cortical microcircuit.