Crossmodal processing of object features in human anterior intraparietal cortex: an fMRI study implies equivalencies between humans and monkeys

The organization of macaque posterior parietal cortex (PPC) reflects its functional specialization in integrating polymodal sensory information for object recognition and manipulation. Neuropsychological and recent human imaging studies imply equivalencies between human and macaque PPC, and in particular, the cortex buried in the intraparietal sulcus (IPS). Using functional MRI, we tested the hypothesis that an area in human anterior intraparietal cortex is activated when healthy subjects perform a crossmodal visuo-tactile delayed matching-to-sample task with objects. Tactile or visual object presentation (encoding and recognition) both significantly activated anterior intraparietal cortex. As hypothesized, neural activity in this area was further enhanced when subjects transferred object information between modalities (crossmodal matching). Based on both the observed functional properties and the anatomical location, we suggest that this area in anterior IPS is the human equivalent of macaque area AIP.     

基于视觉搜索的外源易化和返回抑制脑机制研究

采用事件相关电位技术研究了在视觉搜索过程中的外源易化和返回抑制（inhibition of return,IOR）的相互关系。当外源注意保持在序列搜索过的位置上时,有一个延时反应(即IOR）,伴随其产生的相关脑电成分有：分布在后顶的潜伏期为200 ms 的正差异、分布在前额叶内侧靠左的潜伏期为240 毫秒的负差异,以及分布在两侧颞顶联合区的潜伏期为280 ms 的负差异。而当外源注意保持在平行搜索的位置上时,则出现了明显的易化效应,伴随其产生的脑电成分仅为分布在枕顶区域的潜伏期为280 ms 的负差异。这些结果表明,外源易化和IOR 涉及了不同的脑区和神经过程,从而支持两者在机制上是可分离性的观点。     

Spatiotemporal Dynamics of High-Gamma Activities during a 3-Stimulus Visual Oddball Task

Although many studies have investigated the neural basis of top-down and bottom-up attention, it still requires refinement in both temporal and spatial terms. We used magnetoencephalography to investigate the spatiotemporal dynamics of high-gamma (52–100 Hz) activities during top-down and bottom-up visual attentional processes, aiming to extend the findings from functional magnetic resonance imaging and event-related potential studies. Fourteen participants performed a 3-stimulus visual oddball task, in which both infrequent non-target and target stimuli were presented. We identified high-gamma event-related synchronization in the left middle frontal gyrus, the left intraparietal sulcus, the left thalamus, and the visual areas in different time windows for the target and non-target conditions. We also found elevated imaginary coherence between the left intraparietal sulcus and the right middle frontal gyrus in the high-gamma band from 300 to 400 ms in the target condition, and between the left thalamus and the left middle frontal gyrus in theta band from 150 to 450 ms. In addition, the strength of high-gamma imaginary coherence between the left middle frontal gyrus and left intraparietal sulcus, between the left middle frontal gyrus and the right middle frontal gyrus, and the high-gamma power in the left thalamus predicted inter-subject variation in target detection response time. This source-level electrophysiological evidence enriches our understanding of bi-directional attention processes: stimulus-driven bottom-up attention orientation to a salient, but irrelevant stimulus; and top-down allocation of attentional resources to stimulus evaluation.     

Maps of space in human frontoparietal cortex

Prefrontal cortex (PFC) and posterior parietal cortex (PPC) are neural substrates for spatial cognition. We here review studies in which we tested the hypothesis that human frontoparietal cortex may function as a priority map. According to priority map theory, objects or locations in the visual world are represented by neural activity that is proportional to their attentional priority. Using functional magnetic resonance imaging (fMRI), we first identified topographic maps in PFC and PPC as candidate priority maps of space. We then measured fMRI activity in candidate priority maps during the delay periods of a covert attention task, a spatial working memory task, and a motor planning task to test whether the activity depended on the particular spatial cognition. Our hypothesis was that some, but not all, candidate priority maps in PFC and PPC would be agnostic with regard to what was being prioritized, in that their activity would reflect the location in space across tasks rather than a particular kind of spatial cognition (e.g., covert attention). To test whether patterns of delay period activity were interchangeable during the spatial cognitive tasks, we used multivariate classifiers. We found that decoders trained to predict the locations on one task (e.g., working memory) cross-predicted the locations on the other tasks (e.g., covert attention and motor planning) in superior precentral sulcus (sPCS) and in a region of intraparietal sulcus (IPS2), suggesting that these patterns of maintenance activity may be interchangeable across the tasks. Such properties make sPCS in frontal cortex and IPS2 in parietal cortex viable priority map candidates, and suggest that these areas may be the human homologs of the monkey frontal eye field (FEF) and lateral intraparietal area (LIP).     

Brain Activation of Identity Switching in Multiple Identity Tracking Task

When different objects switch identities in the multiple identity tracking (MIT) task, viewers need to rebind objects’ identity and location, which requires attention. This rebinding helps people identify the regions targets are in (where they need to focus their attention) and inhibit unimportant regions (where distractors are). This study investigated the processing of attentional tracking after identity switching in an adapted MIT task. This experiment used three identity-switching conditions: a target-switching condition (where the target objects switched identities), a distractor-switching condition (where the distractor objects switched identities), and a no-switching condition. Compared to the distractor-switching condition, the target-switching condition elicited greater activation in the frontal eye fields (FEF), intraparietal sulcus (IPS), and visual cortex. Compared to the no-switching condition, the target-switching condition elicited greater activation in the FEF, inferior frontal gyrus (pars orbitalis) (IFG-Orb), IPS, visual cortex, middle temporal lobule, and anterior cingulate cortex. Finally, the distractor-switching condition showed greater activation in the IFG-Orb compared to the no-switching condition. These results suggest that, in the target-switching condition, the FEF and IPS (the dorsal attention network) might be involved in goal-driven attention to targets during attentional tracking. In addition, in the distractor-switching condition, the activation of the IFG-Orb may indicate salient change that pulls attention away automatically.     

不同注意任务类型及难度对猕猴微眼动的影响

微眼动是视觉注视过程中幅度最大、速度最快的眼动,可以消除由于神经系统适应性而产生的视觉衰退现象,在视觉信息处理过程中发挥着重要作用.基于微眼动与视觉感知功能的相关性,设计实验研究猕猴完成显性、隐性注意任务以及不同难度显性注意任务时,视觉注视情况下微眼动的差异.通过对不同难度显性注意任务下微眼动的参数进行比较,发现随着任务难度的增加,微眼动的幅度、速率和频率都被抑制.另一方面,对比不同类型的视觉感知任务(显性注意和隐性注意),发现在相似的实验范式下,隐性注意对微眼动的频率有明显的抑制作用,但幅度和频率没有得到一致的结果,这表明视觉注意任务类型的不同或将导致猕猴完成任务的策略不同.这些工作将为今后进一步研究微眼动产生的神经机制以及视觉注意过程中眼动的作用机制奠定良好的基础.     

Common Cortical Loci Are Activated during Visuospatial Interpolation and Orientation Discrimination Judgements

There is a wealth of literature on the role of short-range interactions between low-level orientation-tuned filters in the perception of discontinuous contours. However, little is known about how spatial information is integrated across more distant regions of the visual field in the absence of explicit local orientation cues, a process referred to here as visuospatial interpolation (VSI). To examine the neural correlates of VSI high field functional magnetic resonance imaging was used to study brain activity while observers either judged the alignment of three Gabor patches by a process of interpolation or discriminated the local orientation of the individual patches. Relative to a fixation baseline the two tasks activated a largely over-lapping network of regions within the occipito-temporal, occipito-parietal and frontal cortices. Activated clusters specific to the orientation task (orientation>interpolation) included the caudal intraparietal sulcus, an area whose role in orientation encoding per se has been hotly disputed. Surprisingly, there were few task-specific activations associated with visuospatial interpolation (VSI>orientation) suggesting that largely common cortical loci were activated by the two experimental tasks. These data are consistent with previous studies that suggest higher level grouping processes -putatively involved in VSI- are automatically engaged when the spatial properties of a stimulus (e.g. size, orientation or relative position) are used to make a judgement.     

A supramodal number representation in human intraparietal cortex

The triple-code theory of numerical processing postulates an abstract-semantic "number sense." Neuropsychology points to intraparietal cortex as a potential substrate, but previous functional neuroimaging studies did not dissociate the representation of numerical magnitude from task-driven effects on intraparietal activation. In an event-related fMRI study, we presented numbers, letters, and colors in the visual and auditory modality, asking subjects to respond to target items within each category. In the absence of explicit magnitude processing, numbers compared with letters and colors across modalities activated a bilateral region in the horizontal intraparietal sulcus. This stimulus-driven number-specific intraparietal response supports the idea of a supramodal number representation that is automatically accessed by presentation of numbers and may code magnitude information.     

Single neurons in posterior parietal cortex of monkeys encode cognitive set

The primate posterior parietal cortex (PPC), part of the dorsal visual pathway, is best known for its role in encoding salient spatial information. Yet there are indications that neural activity in the PPC can also be modulated by nonspatial task-related information. In this study, we tested whether neurons in the PPC encode signals related to cognitive set, that is, the preparation to perform a particular task. Cognitive set has previously been associated with the frontal cortex but not the PPC. In this study, monkeys performed a cognitive set shifting paradigm in which they were cued in advance to apply one of two different task rules to the subsequent stimulus on every trial. Here we show that a subset of neurons in the PPC, concentrated in the lateral bank of the intraparietal sulcus and on the angular gyrus, responds selectively to cues for different task rules.     

Decoding successive computational stages of saliency processing

An important requirement for vision is to identify interesting and relevant regions of the environment for further processing. Some models assume that salient locations from a visual scene are encoded in a dedicated spatial saliency map [1, 2]. Then, a winner-take-all (WTA) mechanism [1, 2] is often believed to threshold the graded saliency representation and identify the most salient position in the visual field. Here we aimed to assess whether neural representations of graded saliency and the subsequent WTA mechanism can be dissociated. We presented images of natural scenes while subjects were in a scanner performing a demanding fixation task, and thus their attention was directed away. Signals in early visual cortex and posterior intraparietal sulcus (IPS) correlated with graded saliency as defined by a computational saliency model. Multivariate pattern classification [3, 4] revealed that the most salient position in the visual field was encoded in anterior IPS and frontal eye fields (FEF), thus reflecting a potential WTA stage. Our results thus confirm that graded saliency and WTA-thresholded saliency are encoded in distinct neural structures. This could provide the neural representation required for rapid and automatic orientation toward salient events in natural environments.     

Neuronal correlates of the set-size effect in monkey lateral intraparietal area

It has long been known that the brain is limited in the amount of sensory information that it can process at any given time. A well-known form of capacity limitation in vision is the set-size effect, whereby the time needed to find a target increases in the presence of distractors. The set-size effect implies that inputs from multiple objects interfere with each other, but the loci and mechanisms of this interference are unknown. Here we show that the set-size effect has a neural correlate in competitive visuo-visual interactions in the lateral intraparietal area, an area related to spatial attention and eye movements. Monkeys performed a covert visual search task in which they discriminated the orientation of a visual target surrounded by distractors. Neurons encoded target location, but responses associated with both target and distractors declined as a function of distractor number (set size). Firing rates associated with the target in the receptive field correlated with reaction time both within and across set sizes. The findings suggest that competitive visuo-visual interactions in areas related to spatial attention contribute to capacity limitations in visual searches.     

Neural activities in v1 create a bottom-up saliency map

The bottom-up contribution to the allocation of exogenous attention is a saliency map, whose neural substrate is hard to identify because of possible contamination by top-down signals. We obviated this possibility using stimuli that observers could not perceive, but that nevertheless, through orientation contrast between foreground and background regions, attracted attention to improve a localized visual discrimination. When orientation contrast increased, so did the degree of attraction, and two physiological measures: the amplitude of the earliest (C1) component of the ERP, which is associated with primary visual cortex, and fMRI BOLD signals in areas V1-V4 (but not the intraparietal sulcus). Significantly, across observers, the degree of attraction correlated with the C1 amplitude and just the V1 BOLD signal. These findings strongly support the proposal that a bottom-up saliency map is created in V1, challenging the dominant view that the saliency map is generated in the parietal cortex.     

Studying modulation on simultaneously activated SSVEP neural networks by a cognitive task

Since the discovery of steady-state visually evoked potential (SSVEP), it has been used in many fields. Numerous studies suggest that there exist three SSVEP neural networks in different frequency bands. An obvious phenomenon has been observed, that the amplitude and phase of SSVEP can be modulated by a cognitive task. Previous works have studied this modulation on separately activated SSVEP neural networks by a cognitive task. If two or more SSVEP neural networks are activated simultaneously in the process of a cognitive task, is the modulation on different SSVEP neural networks the same? In this study, two different SSVEP neural networks were activated simultaneously by two different frequency flickers, with a working memory task irrelevant to the flickers being conducted at the same time. The modulated SSVEP waves were compared with each other and to those only under one flicker in previous studies. The comparison results show that the cognitive task can modulate different SSVEP neural networks with a similar style.     

Evidence for differential top-down and bottom-up suppression in posterior parietal cortex

When searching for an object, we usually avoid items that are visually different from the target and objects or places that have been searched already. Previous studies have shown that neural activity in the lateral intraparietal area (LIP) can be used to guide this behaviour; responses to task irrelevant stimuli or to stimuli that have been fixated previously in the trial are reduced compared with responses to potential targets. Here, we test the hypothesis that these reduced responses have a different genesis. Two animals were trained on a visual foraging task, in which they had to find a target among a number of physically identical potential targets (T) and task irrelevant distractors. We recorded neural activity and local field potentials (LFPs) in LIP while the animals performed the task. We found that LFP power was similar for potential targets and distractors but was greater in the alpha and low beta bands when a previously fixated T was in the response field. We interpret these data to suggest that the reduced single-unit response to distractors is a bottom-up feed-forward result of processing in earlier areas and the reduced response to previously fixated Ts is a result of active top-down suppression.     

One-dimensional dynamics of attention and decision making in LIP

Where we allocate our visual spatial attention depends upon a continual competition between internally generated goals and external distractions. Recently it was shown that single neurons in the macaque lateral intraparietal area (LIP) can predict the amount of time a distractor can shift the locus of spatial attention away from a goal. We propose that this remarkable dynamical correspondence between single neurons and attention can be explained by a network model in which generically high-dimensional firing-rate vectors rapidly decay to a single mode. We find direct experimental evidence for this model, not only in the original attentional task, but also in a very different task involving perceptual decision making. These results confirm a theoretical prediction that slowly varying activity patterns are proportional to spontaneous activity, pose constraints on models of persistent activity, and suggest a network mechanism for the emergence of robust behavioral timing from heterogeneous neuronal populations.     

The role of low and high spatial frequencies in exogenous attention to biologically salient stimuli

Exogenous attention can be understood as an adaptive tool that permits the detection and processing of biologically salient events even when the individual is engaged in a resource-consuming task. Indirect data suggest that the spatial frequency of stimulation may be a crucial element in this process. Behavioral and neural data (both functional and structural) were analyzed for 36 participants engaged in a digit categorization task in which distracters were presented. Distracters were biologically salient or anodyne images, and had three spatial frequency formats: intact, low spatial frequencies only, and high spatial frequencies only. Behavior confirmed enhanced exogenous attention to biologically salient distracters. The activity in the right and left intraparietal sulci and the right middle frontal gyrus was associated with this behavioral pattern and was greater in response to salient than to neutral distracters, the three areas presenting strong correlations to each other. Importantly, the enhanced response of this network to biologically salient distracters with respect to neutral distracters relied on low spatial frequencies to a significantly greater extent than on high spatial frequencies. Structural analyses suggested the involvement of internal capsule, superior longitudinal fasciculus and corpus callosum in this network. Results confirm that exogenous attention is preferentially captured by biologically salient information, and suggest that the architecture and function underlying this process are low spatial frequency-biased.     

Two cortical systems for reaching in central and peripheral vision

Parietal lesions in humans can produce a specific disruption of visually guided hand movement, termed optic ataxia. The fact that the deficit mainly occurs in peripheral vision suggests that reaching in foveal and extrafoveal vision rely on two different neural substrates. In the present study, we have directly tested this hypothesis by event-related fMRI in healthy subjects. Brain activity was measured when participants reached toward central or peripheral visual targets. Our results confirm the existence of two systems, differently modulated by the two conditions. Reaching in central vision involved a restricted network including the medial intraparietal sulcus (mIPS) and the caudal part of the dorsal premotor cortex (PMd). Reaching in peripheral vision activated in addition the parieto-occipital junction (POJ) and a more rostral part of PMd. These results show that reaching to the peripheral visual field engages a more extensive cortical network than reaching to the central visual field.     

Desynchronization α frequency event-related in visual selective attention requiring tasks

Comparative analysis of the EEG amplitude depression in the α band has been performed in two paradigms varying in the degree of involvement of functionally different attention processes: visual search for the relevant stimulus (RS) among many irrelevant stimuli (iRS) and oddball. Simple visual examination of several identical stimuli was used as a control for the visual search task. The method of videooculography was used for verification of gaze direction during RS search. The EEG dynamics in the α band (desynchronization, D) was considered to be a correlate of attention processes. The visual search task performance revealed considerably higher D after RS finding compared to the control. The higher degree of D during the search seems to be due to the higher complexity of the task and complexity of visual environment. The D in the frontal regions, which has the greatest amplitude, supposedly reflects the performance of an adequate motor performance program under the control of voluntary (top-down) attention. At the same time, the D in the occipital and parietal areas seems to reflect the processes of involuntary attention activated due to the change in visual information (the finding of the only RS among numerous iRS). In the oddball task, presentation of both RS and iRS also induced D, which proved to be more marked in response to RS and maximal in the visual areas. We suppose that D under oddball reflects the involvement of involuntary attention.     

The graded and dichotomous nature of visual awareness

Is our visual experience of the world graded or dichotomous? Opposite pre-theoretical intuitions apply in different cases. For instance, when looking at a scene, one has a distinct sense that our experience has a graded character: one cannot say that there is no experience of contents that fall outside the focus of attention, but one cannot say that there is full awareness of such contents either. By contrast, when performing a visual detection task, our sense of having perceived the stimulus or not exhibits a more dichotomous character. Such issues have recently been the object of intense debate because different theoretical frameworks make different predictions about the graded versus dichotomous character of consciousness. Here, we review both relevant empirical findings as well as the associated theories (i.e. local recurrent processing versus global neural workspace theory). Next, we attempt to reconcile such contradictory theories by suggesting that level of processing is an often-ignored but highly relevant dimension through which we can cast a novel look at existing empirical findings. Thus, using a range of different stimuli, tasks and subjective scales, we show that processing low-level, non-semantic content results in graded visual experience, whereas processing high-level semantic content is experienced in a more dichotomous manner. We close by comparing our perspective with existing proposals, focusing in particular on the partial awareness hypothesis.     

Functional Anatomy of Writing with the Dominant Hand

While writing performed by any body part is similar in style, indicating a common program, writing with the dominant hand is particularly skilled. We hypothesized that this skill utilizes a special motor network supplementing the motor equivalence areas. Using functional magnetic resonance imaging in 13 normal subjects, we studied nine conditions: writing, zigzagging and tapping, each with the right hand, left hand and right foot. We identified brain regions activated with the right (dominant) hand writing task, exceeding the activation common to right-hand use and the writing program, both identified without right-hand writing itself. Right-hand writing significantly differed from the other tasks. First, we observed stronger activations in the left dorsal prefrontal cortex, left intraparietal sulcus and right cerebellum. Second, the left anterior putamen was required to initiate all the tested tasks, but only showed sustained activation during the right-hand writing condition. Lastly, an exploratory analysis showed clusters in the left ventral premotor cortex and inferior and superior parietal cortices were only significantly active for right-hand writing. The increased activation with right-hand writing cannot be ascribed to increased effort, since this is a well-practiced task much easier to perform than some of the other tasks studied. Because parietal-premotor connections code for particular skills, it would seem that the parietal and premotor regions, together with basal ganglia-sustained activation likely underlie the special skill of handwriting with the dominant hand.