Large-scale gamma-band phase synchronization and selective attention

Explaining the emergence of a coherent conscious percept and an intentional agent from the activity of distributed neurons is key to understanding how the brain produces higher cognitive processes. Gamma-band synchronization has been proposed to be a mechanism for the functional integration of neural populations that together form a transitory, large-scale, task- and/or percept-specific network. The operation of this mechanism in the context of attention orienting entails that cortical regions representing attended locations should show more gamma-band synchronization with other cortical areas than would those representing unattended locations. This increased synchronization should be apparent in the same time frame as that of the deployment of attention to a particular location. In order to observe this effect, we made electroencephalogram recordings while subjects attended to one side or the other of the visual field (which we confirmed by event-related potential analysis) and calculated phase-locking statistics between the signals recorded at relevant electrode pairs. We observed increased gamma-band phase synchronization between visual cortex contralateral to the attended location and other, widespread, cortical areas approximately 240-380 ms after the directional cue was presented, confirming the prediction of a large-scale gamma synchronous network oriented to the cued location.     

Visual areas in the lateral temporal cortex of the ferret (Mustela putorius)

Using systematic electrophysiological mapping, architectonics and the global pattern of interhemispheric connectivity, we have identified three visual areas in the lateral most part of the posterior suprasylvian gyrus. The most posterior and largest area we call area 20a and anterior to this we defined a smaller area, area 20b. These areas lie lateral to the visual areas 18, 19 and 21 and posterior to a third, but incompletely defined, visual area, area PS. Areas 20a and 20b, emphasize the representation of the upper hemifield. Their interhemispheric connections conform to the so called 'midline rule' in that they are abundant in regions representing central portions of the visual field, scarce or absent elsewhere. These areas are probably homologous to the homonymous areas of the cat and might be indicative of a Bauplan from which the temporal areas of primates may have evolved.     

Attention to form or surface properties modulates different regions of human occipitotemporal cortex

We carried out 2 functional magnetic resonance imaging experiments to investigate the cortical mechanisms underlying the contribution of form and surface properties to object recognition. In experiment 1, participants performed same-different judgments in separate blocks of trials on pairs of unfamiliar "nonsense" objects on the basis of their form, surface properties (i.e., both color and texture), or orientation. Attention to form activated the lateral occipital (LO) area, whereas attention to surface properties activated the collateral sulcus (CoS) and the inferior occipital gyrus (IOG). In experiment 2, participants were required to make same-different judgments on the basis of texture, color, or form. Again attention to form activated area LO, whereas attention to texture activated regions in the IOG and the CoS, as well as regions in the lingual sulcus and the inferior temporal sulcus. Within these last 4 regions, activation associated with texture was higher than activation associated with color. No color-specific cortical areas were identified in these regions, although parts of V1 and the cuneus yielded higher activation for color as opposed to texture. These results suggest that there are separate form and surface-property pathways in extrastriate cortex. The extraction of information about an object's color seems to occur relatively early in visual analysis as compared with the extraction of surface texture, perhaps because the latter requires more complex computations.     

Impaired filtering of distracter stimuli by TE neurons following V4 and TEO lesions in macaques

Directing attention to a behaviorally relevant visual stimulus can overcome the distracting effects of other nearby stimuli. Correspondingly, physiological studies indicate that attention serves to filter distracting stimuli from receptive fields (RFs) in several extrastriate areas. Moreover, a recent study demonstrated that lesions of extrastriate areas V4 and TEO produce impairments in attentional filtering. A critical remaining question concerns why lesions of ventral stream areas cause attentional filtering impairments. To address this question, we tested the effects of restricted area V4 and TEO lesions on both behavioral performance and the responses of downstream neurons in area TE. The lesions impaired behavioral discrimination thresholds and altered neuronal selectivity for target stimuli in the presence of distracters. With attention to the target, but in the absence of V4 and/or TEO inputs, TE neurons responded as though attentional inputs could no longer be used to filter distracters from their RFs. This presumably occurred because top-down attentional signals were no longer able to filter distracters from the RFs of the cells that provide TE with major input. Consistent with this interpretation, increasing the spatial separation between targets and distracters, such that they no longer fell within a typical V4 RF dimension, restored both behavioral performance and neuronal selectivity in the portion of TE RFs affected by the V4 lesion.     

Functional connectivity of frontal cortex in healthy and ADHD children reflected in EEG coherence

Abnormal functional brain connectivity is a candidate factor in developmental brain disorders associated with cognitive dysfunction. We analyzed a substantial (10 min per subject) record of dense array electroencephalography with spectral power and coherence methods in attention-deficit hyperactivity disorder (ADHD) (n = 42) and control (n = 21) 10- to 13-year-old children. We found topographically distinct narrow band coherence differences between subject groups: ADHD subjects showed elevated coherence in the lower alpha (8 Hz) band and reduced coherence in the upper alpha (10-11 Hz) band. The 8-Hz ADHD elevation and a 2- to 6-Hz control group coherence elevation were independent of stimulus presentation. In response to visual stimulation, the ADHD group exhibited reduced evoked potential power and elevated frontal coherence. Only the upper alpha band control group coherence elevation discriminated according to ADHD group medication status. The findings suggest a static state of deficient connectivity in ADHD and a stimulus-induced state of overconnectivity within and between frontal hemispheres.     

Rapid cortical oscillations and early motor activity in premature human neonate

Delta-brush is the dominant pattern of rapid oscillatory activity (8-25 Hz) in the human cortex during the third trimester of gestation. Here, we studied the relationship between delta-brushes in the somatosensory cortex and spontaneous movements of premature human neonates of 29-31 weeks postconceptional age using a combination of scalp electroencephalography and monitoring of motor activity. We found that sporadic hand and foot movements heralded the appearance of delta-brushes in the corresponding areas of the cortex (lateral and medial regions of the contralateral central cortex, respectively). Direct hand and foot stimulation also reliably evoked delta-brushes in the same areas. These results suggest that sensory feedback from spontaneous fetal movements triggers delta-brush oscillations in the central cortex in a somatotopic manner. We propose that in the human fetus in utero, before the brain starts to receive elaborated sensory input from the external world, spontaneous fetal movements provide sensory stimulation and drive delta-brush oscillations in the developing somatosensory cortex contributing to the formation of cortical body maps.     

Control of object-based attention in human cortex

Visual attention is a mechanism by which observers select relevant or important information from the current visual array. Previous investigations have focused primarily on the ability to select a region of space for further visual analysis. These studies have revealed a distributed frontoparietal circuit that is responsible for the control of spatial attention. However, vision must ultimately represent objects and in real scenes objects often overlap spatially; thus attention must be capable of selecting objects and their properties nonspatially. Little is known about the neural basis of object-based attentional control. In two experiments, human observers shifted attention between spatially superimposed faces and houses. Event-related functional magnetic resonance imaging (fMRI) revealed attentional modulation of activity in face- and house-selective cortical regions. Posterior parietal and frontal regions were transiently active when attention was shifted between spatially superimposed perceptual objects. The timecourse of activity provides insight into the functional role that these brain regions play in attentional control processes.     

Beyond retinotopic mapping: the spatial representation of objects in the human lateral occipital complex

The spatial representation in the human ventral object-related areas (i.e., the lateral occipital complex [LOC]) is currently unknown. It seems plausible, however, that it would diverge from the strict retinotopic mapping (characteristic of V1) to a more invariant coordinate frame, thereby allowing for reliable object recognition in the face of eye, head, or body movement. To study this, we compared the fMRI activation in LOC when object displacement was limited to either the retina or the screen by manipulating eye position and object locations. We found clear adaptation in LOC when the object's screen position was fixed, regardless of the object's retinal position. Furthermore, we found significantly greater activation in LOC in the hemisphere contralateral to the object's screen position, although the visual task was constructed in a way that the objects were present equally often on each of the 2 retinal hemifields. Together, these results indicate that a sizeable fraction of the neurons in LOC may have head-based receptive fields. Such an extraretinal representation may be useful for maintenance of object coherence across saccadic eye movements, which are an integral part of natural vision.     

A contralateral preference in the lateral occipital area: sensory and attentional mechanisms

Here we examined the level of the lateral occipital (LO) area within the processing stream of the ventral visual cortex. An important determinant of an area's level of processing is whether it codes visual elements on both sides of the visual field, as do higher visual areas, or prefers those in the contralateral visual field, as do early visual areas. The former would suggest that LO, on one side, combines bilateral visual elements into a whole, while the latter suggests that it codes only the parts of forms. We showed that LO has a relative preference for visual objects in the contralateral visual field. LO responses were influenced by attention. However, relative changes in LO activity caused by changes in object location were preserved even when attention was shifted away from the objects to moving random dot patterns on the opposite side. Our data offer a new view on LO as an intermediate, but not a high-level, visual area in which neurons are driven by visual input and spatial attention in a multiplicative fashion.     

Spatially selective representations of voluntary and stimulus-driven attentional priority in human occipital, parietal, and frontal cortex

When multiple objects are present in a visual scene, they compete for cortical processing in the visual system; selective attention biases this competition so that representations of behaviorally relevant objects enter awareness and irrelevant objects do not. Deployments of selective attention can be voluntary (e.g., shift or attention to a target's expected spatial location) or stimulus driven (e.g., capture of attention by a target-defining feature such as color). Here we use functional magnetic resonance imaging to show that both of these factors induce spatially selective attentional modulations within regions of human occipital, parietal, and frontal cortex. In addition, the voluntary attentional modulations are temporally sustained, indicating that activity in these regions dynamically tracks the locus of attention. These data show that a convolution of factors, including prior knowledge of location and target-defining features, determines the relative competitive advantage of visual stimuli within multiple stages of the visual system.     

A motion-sensitive area in ferret extrastriate visual cortex: an analysis in pigmented and albino animals

In search of the neuronal substrate for motion analysis in the ferret (Mustela putorius furo), we extracellularly recorded from extrastriate visual cortex in five pigmented and two albino ferrets under general anaesthesia and paralysis. Visual stimulation consisted of large area random dot patterns moving either on a circular path in the frontoparallel plane or expanding and contracting radially. Strongly direction-selective neurons were recorded in a circumscribed area in and just posterior to the suprasylvian sulcus, thus named by us the posterior suprasylvian area (area PSS). Altogether, we recorded 210 (90%) and 95 (72%) PSS neurons in pigmented and albino ferrets, respectively, that were direction selective. In these neurons responses during random dot pattern stimulation in the preferred direction were at least twice as strong than stimulation in the non-preferred direction. Response strength in preferred direction and tuning sharpness of PSS neurons in albinos were significantly reduced when compared to pigmented animals (median values: 34.1 versus 14.8 spikes/s and 142 versus 165 degrees for pigmented and albino ferrets, respectively). Inter-spike-intervals during visual stimulation were significantly shorter in pigmented (median 9 ms) than in albino PSS neurons (median 14 ms). Our data indicate that area PSS may play a crucial role in motion perception in the ferret.     

Impaired face discrimination in acquired prosopagnosia is associated with abnormal response to individual faces in the right middle fusiform gyrus

The middle fusiform gyrus (MFG) and the inferior occipital gyrus (IOG) are activated by both detection and identification of faces. Paradoxically, patients with acquired prosopagnosia following lesions to either of these regions in the right hemisphere cannot identify faces, but can still detect faces. Here we acquired functional magnetic resonance imaging (fMRI) data during face processing in a patient presenting a specific deficit in individual face recognition, following lesions encompassing the right IOG. Using an adaptation paradigm we show that the fMRI signal in the rMFG of the patient, while being larger in response to faces as compared to objects, does not differ between conditions presenting identical and distinct faces, in contrast to the larger response to distinct faces observed in controls. These results suggest that individual discrimination of faces critically depends on the integrity of both the rMFG and the rIOG, which may interact through re-entrant cortical connections in the normal brain.     

Oscillatory activity in human parietal and occipital cortex shows hemispheric lateralization and memory effects in a delayed double-step saccade task

We applied magnetoencephalography (MEG) to record oscillatory brain activity from human subjects engaged in planning a double-step saccade. In the experiments, subjects (n = 8) remembered the locations of 2 sequentially flashed targets (each followed by a 2-s delay), presented in either the left or right visual hemifield, and then made saccades to the 2 locations in sequence. We examined changes in spectral power in relation to target location (left or right) and memory load (one or two targets), excluding error trials based on concurrent eye tracking. During the delay period following the first target, power in the alpha (8-12 Hz) and beta (13-25 Hz) bands was significantly suppressed in the hemisphere contralateral to the target. When the second target was presented, there was a further suppression in the alpha- and beta-band power over both hemispheres. In this period, the same sensors also showed contralateral power enhancements in the gamma band (60-90 Hz), most significantly prior to the initiation of the saccades. Adaptive spatial filtering techniques localized the neural sources of the directionally selective power changes in parieto-occipital areas. These results provide further support for a topographic organization for delayed saccades in human parietal and occipital cortex.     

神经导航辅助术中功能区定位切除累及中央前回的脑胶质瘤

目的探讨侵犯运动区的脑胶质瘤手术治疗方法。方法回顾分析12例侵犯中央前回脑胶质瘤病例资料，手术采用神经导航定位病灶和中央前回，术中唤醒麻醉，直接皮质电刺激定位肢体运动或语言运动区，患者清醒状态下切除肿瘤。结果11例术中成功唤醒切除肿瘤，皮质电刺激下7例获得了肢体运动区，2例获得了语言运动区的准确定位。8例（66．7％）达到肿瘤全切除，4例次全切除。8例为低级别胶质瘤，2例为高级别胶质瘤，2例胶质母细胞瘤。术后4例无神经功能缺损，8例出现术后对侧肢体活动障碍或言语障碍，除1例外均在7d至1月内恢复正常。结论导航辅助下的直接皮质电刺激定位功能区和唤醒状态下的肿瘤切除是处理侵犯功能区肿瘤的一种安全、有效的方法，可以获得功能区的准确定位并达到最小程度的功能损伤和最大限度的切除肿瘤。     

Convergence of visual and tactile shape processing in the human lateral occipital complex

We have recently demonstrated using fMRI that a region within the human lateral occipital complex (LOC) is activated by objects when either seen or touched. We term this cortical region LOtv for the lateral occipital tactile-visual region. We report here that LOtv voxels tend to be located in sub-regions of LOC that show preference for graspable visual objects over faces or houses. We further examine the nature of object representation in LOtv by studying its response to stimuli in three modalities: auditory, somatosensory and visual. If objects activate LOtv, irrespective of the modality used, the activation is likely to reflect a highly abstract representation. In contrast, activation specific to vision and touch may reflect common and exclusive attributes shared by these senses. We show here that while object activation is robust in both the visual and the somatosensory modalities, auditory signals do not evoke substantial responses in this region. The lack of auditory activation in LOtv cannot be explained by differences in task performance or by an ineffective auditory stimulation. Unlike vision and touch, auditory information contributes little to the recovery of the precise shape of objects. We therefore suggest that LOtv is involved in recovering the geometrical shape of objects.     

Simultaneous representation of saccade targets and visual onsets in monkey lateral intraparietal area

The monkey's lateral intraparietal area (LIP) has been associated with attention and saccades. LIP neurons have visual on-responses to objects abruptly appearing in their receptive fields (RFs) and sustained activity preceding saccades to the RF. We studied the relationship between the on-responses and delay activity in LIP using a 'stable-array' task. Monkeys viewed eight distinct, continuously illuminated objects, arranged in a circle with at least one object in the RF. A cue flashed instructing the monkey to make a saccade, after a delay, to the stable object physically matching the cue. The location of the cue was fixed in trial blocks, either in or out of the RF. If the cue was outside the RF, neurons developed delay-period activity tuned for the direction of the saccade target at approximately 190 ms after cue onset. If the cue appeared in the RF, neurons initially responded to cue onset and developed tuning for saccade direction only toward the end of the delay period, 390 ms after cue onset. The cue- and saccade-target responses coexisted throughout a significant portion of the delay period. The results show that visual-on responses and delay-period activity in LIP are functionally separable, and that, although highly selective, the salience representation in LIP can contain more than one object at a time.     

Synaptic and spiking dynamics underlying reward reversal in the orbitofrontal cortex

Cognitive and emotional flexibility involve a coordinated interaction between working memory, attention, reward expectations, and the evaluation of rewards and punishers so that behaviour can be changed if necessary. We describe a model at the integrate-and-fire neuronal level of the synaptic and spiking mechanisms which can hold an expectation of a reward rule in working memory, and can reverse the reward rule if expected rewards are not obtained. An example of a reward rule is that stimulus 1 is currently associated with reward, and stimulus 2 with punishment. The attractor-based reward rule working memory incorporates a spike-frequency synaptic adaptation mechanism which supports the neural switching between rules by being shut down by a general inhibitory input produced by punishment, so that when the attractor starts up again is in the opposite state. The mechanism can implement one-trial reward reversal, which is a property of orbitofrontal cortex neurons. We show how this reward rule input can operate in a biased competition way to influence which one of two stimuli is currently associated with reward and which with punishment, and to map the stimuli correctly to the reward or punishment representations, providing a basis for action selection required to obtain the reinforcer.     

Silence is golden: transient neural deactivation in the prefrontal cortex during attentive reading

It is becoming increasingly clear that attention-demanding tasks engage not only activation of specific cortical regions but also deactivation of other regions that could interfere with the task at hand. At the same time, electrophysiological studies in animals and humans have found that the participation of cortical regions to cognitive processes translates into local synchronization of rhythmic neural activity at frequencies above 40 Hz (so-called gamma-band synchronization). Such synchronization is seen as a potential facilitator of neural communication and synaptic plasticity. We found evidence that cognitive processes can also involve the disruption of gamma-band activity in high-order brain regions. Intracerebral electroencephalograms were recorded in 3 epileptic patients during 2 reading tasks. Visual presentation of words induced a strong deactivation in a broad (20-150 Hz) frequency range in the left ventral lateral prefrontal cortex, in parallel with gamma-band activations within the reading network, including Broca's area. The observed energy decrease in neural signals was reproducible across patients. It peaked around 500 ms after stimulus onset and appeared subject to attention-modulated amplification. Our results suggest that cognition might be mediated by a coordinated interaction between regional gamma-band synchronizations and desynchronizations, possibly reflecting enhanced versus reduced local neural communication.     

Serial and parallel processing in the human auditory cortex: a magnetoencephalographic study

Although anatomical, histochemical and electrophysiological findings in both animals and humans have suggested a parallel and serial mode of auditory processing, precise activation timings of each cortical area are not well known, especially in humans. We investigated the timing of arrival of signals to multiple cortical areas using magnetoencephalography in humans. Following click stimuli applied to the left ear, activations were found in six cortical areas in the right hemisphere: the posteromedial part of Heschl's gyrus (HG) corresponding to the primary auditory cortex (PAC), the anterolateral part of the HG region on or posterior to the transverse sulcus, the posterior parietal cortex (PPC), posterior and anterior parts of the superior temporal gyrus (STG), and the planum temporale (PT). The mean onset latencies of each cortical activity were 17.1, 21.2, 25.3, 26.2, 30.9 and 47.6 ms respectively. These results suggested a serial model of auditory processing along the medio-lateral axis of the supratemporal plane and, in addition, implied the existence of several parallel streams running postero-superiorly (from the PAC to the belt region and then to the posterior STG, PPC or PT) and anteriorly (PAC-belt-anterior STG).     

Functional specializations in lateral prefrontal cortex associated with the integration and segregation of information in working memory

Control processes are thought to play an important role in working memory (WM), by enabling the coordination, transformation, and integration of stored information. Yet little is known about the neural mechanisms that subserve such control processes. This study examined whether integration operations within WM involve the activation of distinct neural mechanisms within lateral prefrontal cortex (PFC). Event-related functional magnetic resonance imaging was used to monitor brain activity while participants performed a mental arithmetic task. In the integration (IN) condition, a WM preload item had to be mentally inserted into the last step of the math problem. This contrasted with the segregation (SG) condition, which also required maintenance of the WM preload while performing mental arithmetic but had no integration requirement. Two additional control conditions involved either ignoring the preload (math only condition) or ignoring the math problem (recall only condition). Left anterior PFC (Brodmann's Area [BA] 46/10) was selectively engaged by integration demands, with activation increasing prior to, as well as during the integration period. A homologous right anterior PFC region showed selectively increased activity in the SG condition during the period in which the math problem and preload digit were reported. Left middorsolateral PFC regions (BA 9/46) showed increased, but equivalent, activity in both the SG and IN conditions relative to both control conditions. These results provide support for the selective role of lateral PFC in cognitive control over WM and suggest more specific hypotheses regarding dissociable PFC mechanisms involved during the integration and segregation of stored WM items.