Timing and sequence of brain activity in top-down control of visual-spatial attention

Recent brain imaging studies using functional magnetic resonance imaging (fMRI) have implicated a frontal-parietal network in the top-down control of attention. However, little is known about the timing and sequence of activations within this network. To investigate these timing questions, we used event-related electrical brain potentials (ERPs) and a specially designed visual-spatial attentional-cueing paradigm, which were applied as part of a multi-methodological approach that included a closely corresponding event-related fMRI study using an identical paradigm. In the first 400 ms post cue, attention-directing and control cues elicited similar general cue-processing activity, corresponding to the more lateral subregions of the frontal-parietal network identified with the fMRI. Following this, the attention-directing cues elicited a sustained negative-polarity brain wave that was absent for control cues. This activity could be linked to the more medial frontal–parietal subregions similarly identified in the fMRI as specifically involved in attentional orienting. Critically, both the scalp ERPs and the fMRI-seeded source modeling for this orienting-related activity indicated an earlier onset of frontal versus parietal contribution (∼400 versus ∼700 ms). This was then followed (∼800–900 ms) by pretarget biasing activity in the region-specific visual-sensory occipital cortex. These results indicate an activation sequence of key components of the attentional-control brain network, providing insight into their functional roles. More specifically, these results suggest that voluntary attentional orienting is initiated by medial portions of frontal cortex, which then recruit medial parietal areas. Together, these areas then implement biasing of region-specific visual-sensory cortex to facilitate the processing of upcoming visual stimuli.     

宽周边视视网膜皮层映射的核磁共振研究

目的:人类视觉皮层的组织方式是视网膜皮层映射组织,先前研究已经证实视觉皮层在中心视采用这种组织方式,本文主要研究宽周边视的视觉皮层组织方式.方法:本文采用一种可以在核磁共振室中使用的光纤设备,设计了30度、40度、50度、60度的类圆环block刺激,使用1.5T的功能性核磁共振仪器,T1高分辨率图像分辨率为1*1*5.5mm,T2加权图像分辨率为4*4*5.5mm,TR反应时间为60,矩阵大小为64*64.核磁共振数据分析使用了SPM2和Brain voyager软件.结果:通过对试验者的数据处理分析,周边视的刺激的反应区域在枕叶上,主要分布在枕叶的前部,刺激反应区域随着偏心率的增大而沿着距状沟从距状沟的后部向前部移动.结论:周边视的视网膜皮层映射组织特性和中心视的特性非常相似.     

Functionally independent components of early event-related potentials in a visual spatial attention task.

Spatial visual attention modulates the first negative-going deflection in the human averaged event-related potential (ERP) in response to visual target and non-target stimuli (the N1 complex). Here we demonstrate a decomposition of N1 into functionally independent subcomponents with functionally distinct relations to task and stimulus conditions. ERPs were collected from 20 subjects in response to visual target and non-target stimuli presented at five attended and non-attended screen locations. Independent component analysis, a new method for blind source separation, was trained simultaneously on 500 ms grand average responses from all 25 stimulus-attention conditions and decomposed the non-target N1 complexes into five spatially fixed, temporally independent and physiologically plausible components. Activity of an early, laterally symmetrical component pair (N1aR and N1aL) was evoked by the left and right visual field stimuli, respectively. Component N1aR peaked ca. 9 ms earlier than N1aL. Central stimuli evoked both components with the same peak latency difference, producing a bilateral scalp distribution. The amplitudes of these components were no reliably augmented by spatial attention. Stimuli in the right visual field evoked activity in a spatio-temporally overlapping bilateral component (N1b) that peaked at ca. 180 ms and was strongly enhanced by attention. Stimuli presented at unattended locations evoked a fourth component (P2a) peaking near 240 ms. A fifth component (P3f) was evoked only by targets presented in either visual field. The distinct response patterns of these components across the array of stimulus and attention conditions suggest that they reflect activity in functionally independent brain systems involved in processing attended and unattended visuospatial events.     

A preference for contralateral stimuli in human object- and face-selective cortex

Visual input from the left and right visual fields is processed predominantly in the contralateral hemisphere. Here we investigated whether this preference for contralateral over ipsilateral stimuli is also found in high-level visual areas that are important for the recognition of objects and faces. Human subjects were scanned with functional magnetic resonance imaging (fMRI) while they viewed and attended faces, objects, scenes, and scrambled images in the left or right visual field. With our stimulation protocol, primary visual cortex responded only to contralateral stimuli. The contralateral preference was smaller in object- and face-selective regions, and it was smallest in the fusiform gyrus. Nevertheless, each region showed a significant preference for contralateral stimuli. These results indicate that sensitivity to stimulus position is present even in high-level ventral visual cortex.     

Attention Narrows Position Tuning of Population Responses in V1

When attention is directed to a region of space, visual resolution at that location flexibly adapts, becoming sharper to resolve fine-scale details or coarser to reflect large-scale texture and surface properties [1]. By what mechanism does attention improve spatial resolution? An improved signal-to-noise ratio (SNR) at the attended location contributes [2], because of retinotopically specific signal gain [3], [4], [5], [6], [7], [8], [9] and [10]. Additionally, attention could sharpen position tuning at the neural population level, so that adjacent objects activate more distinct regions of the visual cortex. A dual mechanism involving both signal gain and sharpened position tuning would be highly efficient at improving visual resolution, but there is no direct evidence that attention can narrow the position tuning of population responses. Here, we compared the spatial spread of the fMRI BOLD response for attended versus ignored stimuli. The activity produced by adjacent stimuli overlapped less when subjects were attending at their locations versus attending elsewhere, despite a stronger peak response with attention. Our results show that even as early as primary visual cortex (V1), spatially directed attention narrows the tuning of population-coded position representations.     

Mood modulates auditory laterality of hemodynamic mismatch responses during dichotic listening

Hemodynamic mismatch responses can be elicited by deviant stimuli in a sequence of standard stimuli even during cognitive demanding tasks. Emotional context is known to modulate lateralized processing. Right-hemispheric negative emotion processing may bias attention to the right and enhance processing of right-ear stimuli. The present study examined the influence of induced mood on lateralized pre-attentive auditory processing of dichotic stimuli using functional magnetic resonance imaging (fMRI). Faces expressing emotions (sad/happy/neutral) were presented in a blocked design while a dichotic oddball sequence with consonant-vowel (CV) syllables in an event-related design was simultaneously administered. Twenty healthy participants were instructed to feel the emotion perceived on the images and to ignore the syllables. Deviant sounds reliably activated bilateral auditory cortices and confirmed attention effects by modulation of visual activity. Sad mood induction activated visual, limbic and right prefrontal areas. A lateralization effect of emotion-attention interaction was reflected in a stronger response to right-ear deviants in the right auditory cortex during sad mood. This imbalance of resources may be a neurophysiological correlate of laterality in sad mood and depression. Conceivably, the compensatory right-hemispheric enhancement of resources elicits increased ipsilateral processing.     

Localization of human brain areas activated for chaotic and ordered pattern perception

The aim of our work was to localize cortical areas involved in the processing of incomplete figures using functional MRI (fMRI) for 8 healthy volunteers (18-30 year old) with the did of anatomical and fMRI fast imaging technique: echo planar imaging (EPI), whole brain scan (36 slices) matrix 64 x 64, 3.7 second. We used 1.5 T MR-scanner and BOLD-method (Blood Oxygenation Level Dependent), based on distinctions of magnetic properties of hemoglobin. Fast imaging technique on modern MR-scanners with > or = 1.5 T provides precise statistical maps of oxygenation increase with high spatial resolution. For test stimuli we used matrix of Gabor grating. We used two types of 10 x 10 matrices with chaotic and ordered orientation of Gabor gratings. The size, brightness and contrast of the stimuli were identical. The chaotic and ordered patterns activated different brain areas. We establish that ordered patterns activated only primary visual cortex - V1 and V2, (BA17-18), wheareas chaotic patterns activated in addition primary visual cortex, the V3,V4,V5 (BA19) of the occipital cortex and the area 7 of parietal area (BA7) classification. Decision making for that task is localized in prefrontal and frontal cortex, including (BA 6, 9, 10).     

Feature-based attention modulates orientation-selective responses in human visual cortex

How does feature-based attention modulate neural responses? We used adaptation to quantify the effect of feature-based attention on orientation-selective responses in human visual cortex. Observers were adapted to two superimposed oblique gratings while attending to one grating only. We measured the magnitude of attention-induced orientation-selective adaptation both psychophysically, by the behavioral tilt aftereffect, and physiologically, using fMRI response adaptation. We found evidence for orientation-selective attentional modulation of neuronal responses-a lower fMRI response for the attended than the unattended orientation-in multiple visual areas, and a significant correlation between the magnitude of the tilt aftereffect and that of fMRI response adaptation in V1, the earliest site of orientation coding. These results show that feature-based attention can selectively increase the response of neuronal subpopulations that prefer the attended feature, even when the attended and unattended features are coded in the same visual areas and share the same retinotopic location.     

Increased activity in human visual cortex during directed attention in the absence of visual stimulation

When subjects direct attention to a particular location in a visual scene, responses in the visual cortex to stimuli presented at that location are enhanced, and the suppressive influences of nearby distractors are reduced. What is the top-down signal that modulates the response to an attended versus an unattended stimulus? Here, we demonstrate increased activity related to attention in the absence of visual stimulation in extrastriate cortex when subjects covertly directed attention to a peripheral location expecting the onset of visual stimuli. Frontal and parietal areas showed a stronger signal increase during this expectation than did visual areas. The increased activity in visual cortex in the absence of visual stimulation may reflect a top-down bias of neural signals in favor of the attended location, which derives from a fronto-parietal network.     

Neuromagnetic evidence that the P100 component of the pattern reversal visual evoked response originates in the bottom of the calcarine fissure

Visual evoked magnetic fields due to pattern reversal stimuli were measured in 5 normal subjects using a helmet-shaped 66 channel magnetoencephalography system linked to MRI. The magnetic topography of the prominent 100 ms response (P100m) evoked by full-field visual showed a double-dipole pattern in the occipital areas of all subjects. Right or left half-field stimuli and upper or lower quadrant-field stimuli evoked a single-dipole pattern in the contralateral occipital area. The P100m sources were when localized using a current dipole model and superimposed on MRI images of each subject. The visual cortex was morphologically variable among the subjects, but the P100m dipoles were all localized at the lateral bottom of the calcarine fissure. Moreover, these P100m dipoles had similar orientations for both half- or quadrant-field stimuli. These results suggest that the P100m is located in a smaller part of the striate cortex than previously reported.     

Long-range, pattern-dependent contextual effects in early human visual cortex

The standard view of neurons in early visual cortex is that they behave like localized feature detectors. Here we demonstrate that processing in early visual areas goes beyond feature detection by showing that neural responses are greater when a feature deviates from its context compared to when it does not deviate from its context. Using psychophysics, fMRI, and electroencephalography methodologies, we measured neural responses to an oriented Gabor ("target") embedded in various visual patterns as defined by the relative orientation of flanking stimuli. We first show using psychophysical contrast adaptation and fMRI that a target that differs from its context results in more neural activity compared to a target that is contained within an alternating sequence, suggesting that neurons in early visual cortex are sensitive to large-scale orientation patterns. Next, we use event-related potentials to show that orientation deviations affect the earliest sensory components of the target response. Finally, we use forced-choice classification of "noise" stimuli to show that we are more likely to "see" orientations that deviate from the context. Our results suggest that early visual cortex is sensitive to global patterns in images in a way that is markedly different from the predictions of standard models of cortical visual processing.     

Predictive Coding or Evidence Accumulation? False Inference and Neuronal Fluctuations

Perceptual decisions can be made when sensory input affords an inference about what generated that input. Here, we report findings from two independent perceptual experiments conducted during functional magnetic resonance imaging (fMRI) with a sparse event-related design. The first experiment, in the visual modality, involved forced-choice discrimination of coherence in random dot kinematograms that contained either subliminal or periliminal motion coherence. The second experiment, in the auditory domain, involved free response detection of (non-semantic) near-threshold acoustic stimuli. We analysed fluctuations in ongoing neural activity, as indexed by fMRI, and found that neuronal activity in sensory areas (extrastriate visual and early auditory cortex) biases perceptual decisions towards correct inference and not towards a specific percept. Hits (detection of near-threshold stimuli) were preceded by significantly higher activity than both misses of identical stimuli or false alarms, in which percepts arise in the absence of appropriate sensory input. In accord with predictive coding models and the free-energy principle, this observation suggests that cortical activity in sensory brain areas reflects the precision of prediction errors and not just the sensory evidence or prediction errors per se.     

Contrast invariance in the human lateral occipital complex depends on attention

The human visual system has a remarkable ability to successfully operate under a variety of challenging viewing conditions. For example, our object-recognition capabilities are largely unaffected by low-contrast (e.g., foggy) environments. The basis for this ability appears to be reflected in the neural responses in higher cortical visual areas that have been characterized as being invariant to changes in luminance contrast: neurons in these areas respond nearly equally to low-contrast as compared to high-contrast stimuli. This response pattern is fundamentally different than that observed in earlier visual areas such as primary visual cortex (V1), which is highly dependent on contrast. How this invariance is achieved in higher visual areas is largely unknown. We hypothesized that directed spatial attention is an important prerequisite of the contrast-invariant responses in higher visual areas and tested this with functional MRI (fMRI) while subjects directed their attention either toward or away from contrast-varying shape stimuli. We found that in the lateral occipital complex (LOC), a visual area important for processing shape information, attention changes the form of the contrast response function (CRF). By directing attention away from the shape stimuli, the CRF in the LOC was similar to that measured in V1. We describe a number of mechanisms that could account for this important function of attention.     

Separate modulations of human V1 associated with spatial attention and task structure

Functional magnetic resonance imaging (fMRI) was used while normal human volunteers engaged in simple detection and discrimination tasks, revealing separable modulations of early visual cortex associated with spatial attention and task structure. Both modulations occur even when there is no change in sensory stimulation. The modulation due to spatial attention is present throughout the early visual areas V1, V2, V3, and VP, and varies with the attended location. The task structure activations are strongest in V1 and are greater in regions that represent more peripheral parts of the visual field. Control experiments demonstrate that the task structure activations cannot be attributed to visual, auditory, or somatosensory processing, the motor response for the detection/discrimination judgment, or oculomotor responses such as blinks or saccades. These findings demonstrate that early visual areas are modulated by at least two types of endogenous signals, each with distinct cortical distributions.     

形状识别的功能定位和时间过程:功能磁共振与脑电结合的研究

通过结合具有高空间分辨率的功能磁共振成像（fMRI）和具有高时间分辨率的128导脑电事件相关电位（ERP）两项技术,测量了视皮层腹侧区域对图形形状识别任务反应的空间定位和时间过程。fMRI的实验结果表明,图形的形状和觉引起了腹测GTi／GF皮层区域的兴奋。进一步,基于fMRI兴奋区域的种子偶极子模型拟合的的ERP动态定位分析的结果和自由运动的偶极子模型拟合的ERP定位分析结果表明：GTi／GF区域活动的时间发生在刺激呈现之后132－176ms时间段,峰值150ms左右,相应于ERP的N1成分。这些结果在人类大脑皮层上同时确定了视觉通路中涉及图形形状识别的兴奋区域和兴奋的时间过程。     

Decoding seen and attended motion directions from activity in the human visual cortex

Functional neuroimaging has successfully identified brain areas that show greater responses to visual motion and adapted responses to repeated motion directions. However, such methods have been thought to lack the sensitivity and spatial resolution to isolate direction-selective responses to individual motion stimuli. Here, we used functional magnetic resonance imaging (fMRI) and pattern classification methods to show that ensemble activity patterns in human visual cortex contain robust direction-selective information, from which it is possible to decode seen and attended motion directions. Ensemble activity in areas V1-V4 and MT+/V5 allowed us to decode which of eight possible motion directions the subject was viewing on individual stimulus blocks. Moreover, ensemble activity evoked by single motion directions could effectively predict which of two overlapping motion directions was the focus of the subject's attention and presumably dominant in perception. Our results indicate that feature-based attention can bias direction-selective population activity in multiple visual areas, including MT+/V5 and early visual areas (V1-V4), consistent with gain-modulation models of feature-based attention and theories of early attentional selection. Our approach for measuring ensemble direction selectivity may provide new opportunities to investigate relationships between attentional selection, conscious perception, and direction-selective responses in the human brain.     

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.     

The temporal resolution of neural codes: does response latency have a unique role?

This article reviews the nature of the neural code in non-human primate cortex and assesses the potential for neurons to carry two or more signals simultaneously. Neurophysiological recordings from visual and motor systems indicate that the evidence for a role for precisely timed spikes relative to other spike times (ca. 1-10 ms resolution) is inconclusive. This indicates that the visual system does not carry a signal that identifies whether the responses were elicited when the stimulus was attended or not. Simulations show that the absence of such a signal reduces, but does not eliminate, the increased discrimination between stimuli that are attended compared with when the stimuli are unattended. The increased accuracy asymptotes with increased gain control, indicating limited benefit from increasing attention. The absence of a signal identifying the attentional state under which stimuli were viewed can produce the greatest discrimination between attended and unattended stimuli. Furthermore, the greatest reduction in discrimination errors occurs for a limited range of gain control, again indicating that attention effects are limited. By contrast to precisely timed patterns of spikes where the timing is relative to other spikes, response latency provides a fine temporal resolution signal (ca. 10 ms resolution) that carries information that is unavailable from coarse temporal response measures. Changes in response latency and changes in response magnitude can give rise to different predictions for the patterns of reaction times. The predictions are verified, and it is shown that the standard method for distinguishing executive and slave processes is only valid if the representations of interest, as evidenced by the neural code, are known. Overall, the data indicate that the signalling evident in neural signals is restricted to the spike count and the precise times of spikes relative to stimulus onset (response latency). These coding issues have implications for our understanding of cognitive models of attention and the roles of executive and slave systems.     

Characteristics of the parietal cortex activation in humans during different attention tasks

The cortical activation was estimated by the event-related potential (ERPs) methods during selection tasks of lateralized visual stimuli requiring different forms of attention: 1) form of stimuli, 2) stimuli position, 3) combined attention of form and position. The ERPs were recorded in 15 young healthy adults in 6 leads P3, P4, T3, T4, T5, T6, and endogenous ERPs components: CNV (contingent negative variation), N1, P3 and the complex [N1-P3]. Differences between the ERPs at "attended" and "non-attended" stimuli were considered as indices of selection attention of particular feature of visual stimuli. Such indices of form and position were revealed selectivity in parietal leads. The most eminent ERPs components, the pronounced activation gradient during increase of attention demands were revealed in parietal regions (vs. temporal ones). In our opinion, parietal cortex has a high priority in selection attention system.     

Transient attention enhances perceptual performance and FMRI response in human visual cortex

When a visual stimulus suddenly appears, it captures attention, producing a transient improvement of performance on basic visual tasks. We investigate the effect of transient attention on stimulus representations in early visual areas using rapid event-related fMRI. Participants discriminated the orientation of one of two gratings preceded or followed by a nonpredictive peripheral cue. Compared to control conditions, precueing the target location improved performance and produced a larger fMRI response in corresponding retinotopic areas. This enhancement progressively increased from striate to extrastriate areas. Control conditions indicated that the enhanced fMRI response was not due to sensory summation of cue and target signals. Thus, an uninformative precue increases both perceptual performance and the concomitant stimulus-evoked activity in early visual areas. These results provide evidence regarding the retinotopically specific neural correlate for the effects of transient attention on early vision.