Electrophysiological evidence for spatiotemporal flexibility in the ventrolateral attention network

Successful completion of many everyday tasks depends on interactions between voluntary attention, which acts to maintain current goals, and reflexive attention, which enables responding to unexpected events by interrupting the current focus of attention. Past studies, which have mostly examined each attentional mechanism in isolation, indicate that volitional and reflexive orienting depend on two functionally specialized cortical networks in the human brain. Here we investigated how the interplay between these two cortical networks affects sensory processing and the resulting overt behavior. By combining measurements of human performance and electrocortical recordings with a novel analytical technique for estimating spatiotemporal activity in the human cortex, we found that the subregions that comprise the reflexive ventrolateral attention network dissociate both spatially and temporally as a function of the nature of the sensory information and current task demands. Moreover, we found that together with the magnitude of the early sensory gain, the spatiotemporal neural dynamics accounted for the high amount of the variance in the behavioral data. Collectively these data support the conclusion that the ventrolateral attention network is recruited flexibly to support complex behaviors.     

Development of brain networks for orienting to novelty

Among the many contributions of I.P. Pavlov to the study of the higher nervous system was his exploration of the physical basis of attention. Pavlov's analysis of the orienting reflex (OR), together with the contributions of Y.N. Sokolov, demonstrated that attention could be studies by the objective methods of neurophysiology. Cognitive neuroscience has continued these effort by using neuroimaging to explore the anatomy and physiology of attention in the working human brain. It is now possible to show that the appearance of a novel visual event invokes multiple attentional networks that work in concert to orient to and process a novel object within a very brief exposure. In this paper, we describe cognitive studies of orienting to novelty in adults, examine the networks of neural areas involved in processing novel objects, and review the development in early life of these attentional networks. Orienting to novelty provides an excellent vehicle for examining how biology and experience shape the mechanisms of self-regulation and cognitive control.     

A biased competition account of attention and memory in Alzheimer's disease

The common view of Alzheimer''s disease (AD) is that of an age-related memory disorder, i.e. declarative memory deficits are the first signs of the disease and associated with progressive brain changes in the medial temporal lobes and the default mode network. However, two findings challenge this view. First, new model-based tools of attention research have revealed that impaired selective attention accompanies memory deficits from early pre-dementia AD stages on. Second, very early distributed lesions of lateral parietal networks may cause these attention deficits by disrupting brain mechanisms underlying attentional biased competition. We suggest that memory and attention impairments might indicate disturbances of a common underlying neurocognitive mechanism. We propose a unifying account of impaired neural interactions within and across brain networks involved in attention and memory inspired by the biased competition principle. We specify this account at two levels of analysis: at the computational level, the selective competition of representations during both perception and memory is biased by AD-induced lesions; at the large-scale brain level, integration within and across intrinsic brain networks, which overlap in parietal and temporal lobes, is disrupted. This account integrates a large amount of previously unrelated findings of changed behaviour and brain networks and favours a brain mechanism-centred view on AD.     

The neurobiology of selective attention.

Research in the field of selective visual attention has recently seen substantial progress in several areas. Neuroimaging and electrical recording results have indicated that selective attention amplifies neural activity in prestriate areas concerned with basic visual processing. Imaging and cellular studies are delineating the networks of anatomical areas that serve as the source of attentional modulation and have suggested that these networks are anatomically distinct from the sites of the resulting amplifications. Cognitive studies of visual search have explored the role of these amplified computations in the integration of visual features into objects. Attentional effects in normal subjects, and their disruption following brain injury, have revealed the mental representations upon which attention operates.     

Sensory gain control (amplification) as a mechanism of selective attention: electrophysiological and neuroimaging evidence.

Both physiological and behavioral studies have suggested that stimulus-driven neural activity in the sensory pathways can be modulated in amplitude during selective attention. Recordings of event-related brain potentials indicate that such sensory gain control or amplification processes play an important role in visual-spatial attention. Combined event-related brain potential and neuroimaging experiments provide strong evidence that attentional gain control operates at an early stage of visual processing in extrastriate cortical areas. These data support early selection theories of attention and provide a basis for distinguishing between separate mechanisms of attentional suppression (of unattended inputs) and attentional facilitation (of attended inputs).     

Tracking moving identities: after attending the right location, the identity does not come for free

Although tracking identical moving objects has been studied since the 1980''s, only recently the study into tracking moving objects with distinct identities has started (referred to as Multiple Identity Tracking, MIT). So far, only behavioral studies into MIT have been undertaken. These studies have left a fundamental question regarding MIT unanswered, is MIT a one-stage or a two-stage process? According to the one-stage model, after a location has been attended, the identity is released without effort. However, according to the two-stage model, there are two effortful stages in MIT, attending to a location, and attending to the identity of the object at that location. In the current study we investigated this question by measuring brain activity in response to tracking familiar and unfamiliar targets. Familiarity is known to automate effortful processes, so if attention to identify the object is needed, this should become easier. However, if no such attention is needed, familiarity can only affect other processes (such as memory for the target set). Our results revealed that on unfamiliar trials neural activity was higher in both attentional networks, and visual identification networks. These results suggest that familiarity in MIT automates attentional identification processes, thus suggesting that attentional identification is needed in MIT. This then would imply that MIT is essentially a two-stage process, since after attending the location, the identity does not seem to come for free.     

Control of goal-directed and stimulus-driven attention in the brain

We review evidence for partially segregated networks of brain areas that carry out different attentional functions. One system, which includes parts of the intraparietal cortex and superior frontal cortex, is involved in preparing and applying goal-directed (top-down) selection for stimuli and responses. This system is also modulated by the detection of stimuli. The other system, which includes the temporoparietal cortex and inferior frontal cortex, and is largely lateralized to the right hemisphere, is not involved in top-down selection. Instead, this system is specialized for the detection of behaviourally relevant stimuli, particularly when they are salient or unexpected. This ventral frontoparietal network works as a 'circuit breaker' for the dorsal system, directing attention to salient events. Both attentional systems interact during normal vision, and both are disrupted in unilateral spatial neglect.     

Breakdown of functional connectivity in frontoparietal networks underlies behavioral deficits in spatial neglect

Spatial neglect is a syndrome following stroke manifesting attentional deficits in perceiving and responding to stimuli in the contralesional field. We examined brain network integrity in patients with neglect by measuring coherent fluctuations of fMRI signals (functional connectivity). Connectivity in two largely separate attention networks located in dorsal and ventral frontoparietal areas was assessed at both acute and chronic stages of recovery. Connectivity in the ventral network, part of which directly lesioned, was diffusely disrupted and showed no recovery. In the structurally intact dorsal network, interhemispheric connectivity in posterior parietal cortex was acutely disrupted but fully recovered. This acute disruption, and disrupted connectivity in specific pathways in the ventral network, strongly correlated with impaired attentional processing across subjects. Lastly, disconnection of the white matter tracts connecting frontal and parietal cortices was associated with more severe neglect and more disrupted functional connectivity. These findings support a network view in understanding neglect.     

Neuropeptides,mental performance and aging

The presence of peptidergic neuronal networks in the brain and the modulating action of neuropeptides on brain functions as evidenced by their behavioral influence in particular support the concept that the brain like the peripheral endocrine glands is an endocrine target organ which is as sensitive to treatment with neuropeptides as the peripheral glands are to pituitary hormones. Animal and human data are reviewed showing that neuropeptides related to ACTH/MSH affect motivational and attentional processes and that those related to vasopressin are involved in memory processes. Since these functions decline during aging it is postulated that a decreased bioavailability of neuropeptides in brain of elderly people is associated with specific disturbances in mental performance. Thus, the decreased mental ability of the aged may be restored by treatment with neuropeptides particularly those with little, if any, peripheral, endocrine activity, like the ACTH neuropeptide Org 2766 and the vasopressin neuropeptide DGAVP.     

The behavioral ecology of a cognitive constraint: limited attention

Limited attention may constrain animal behavior in situationsin which the rate of relevant information exceeds the thresholdprocessing capacity of the brain. In the present study, we examinewhy attention is limited by quantifying how attention affectsthe ubiquitous problem of balancing foraging and antipredatoractivity. We analyze how a given attentional capacity affectsfeeding requirements, the optimal attentional focus during predatorscanning, and the probability of detecting predators. Our modelindicates that because of the complex interplay between thecosts and benefits associated with a given attentional capacity,limited attention can be an optimal strategy, which allows effectiveand economical search for cryptic objects.     

White Matter Hyperintensities among Older Adults Are Associated with Futile Increase in Frontal Activation and Functional Connectivity during Spatial Search

The mechanisms by which aging and other processes can affect the structure and function of brain networks are important to understanding normal age-related cognitive decline. Advancing age is known to be associated with various disease processes, including clinically asymptomatic vascular and inflammation processes that contribute to white matter structural alteration and potential injury. The effects of these processes on the function of distributed cognitive networks, however, are poorly understood. We hypothesized that the extent of magnetic resonance imaging white matter hyperintensities would be associated with visual attentional control in healthy aging, measured using a functional magnetic resonance imaging search task. We assessed cognitively healthy older adults with search tasks indexing processing speed and attentional control. Expanding upon previous research, older adults demonstrate activation across a frontal-parietal attentional control network. Further, greater white matter hyperintensity volume was associated with increased activation of a frontal network node independent of chronological age. Also consistent with previous research, greater white matter hyperintensity volume was associated with anatomically specific reductions in functional magnetic resonance imaging functional connectivity during search among attentional control regions. White matter hyperintensities may lead to subtle attentional network dysfunction, potentially through impaired frontal-parietal and frontal interhemispheric connectivity, suggesting that clinically silent white matter biomarkers of vascular and inflammatory injury can contribute to differences in search performance and brain function in aging, and likely contribute to advanced age-related impairments in cognitive control.     

A computational model for the neurobiological substrates of visual attention

Two interesting and complex tasks are performed by the brain in the process of perception: the integration of characteristics leading to an easier recognition of a pattern as a whole (binding), and the extraction of properties that need to be detailed and analyzed (attention). Attention seems to have a reciprocal relation with binding, inasmuch as the latter promotes the composition of features and their dependencies, while the former selects a single characteristic independently of the remainder. Classically, binding is viewed as a process whereby sets of properties are gathered in representative entities, which are themselves linked to form higher level structures, in a sequence that culminates in the total integration of the pattern features in a localized construct. The convergent axonal projections from one cortical area to another would be the neurobiological mechanism through which binding is achieved. Attention comprises the selective excitation of neuronal networks or pathways that stand for specific pattern properties. The thalamus and its reticular nucleus would then be the anatomical substrate of the attentional focus. In this paper we propose a computational model aiming at bringing together the main (and apparently diverging) ideas about binding and attention. Based on experimental data, a neuronal network representing cortical pyramidal cells is assembled, and its structure and function are related to the binding and attention phenomena. Actually, the convergent projections that enlarge the visual receptive field are associated to binding, while a specific change in the pyramidal cell behavior is responsible for attention. Computer simulations are shown which reproduce the electrophysiology of pyramidal cells and mimic some interesting experimental results in visual attention. We conclude by conjecturing that attention is a driven interruption in the regular process of binding.     

Attention,self-regulation and consciousness.

Consciousness has many aspects. These include awareness of the world, feelings of control over one''s behaviour and mental state (volition), and the notion of continuing self. Focal (executive) attention is used to control details of our awareness and is thus closely related to volition. Experiments suggest an integrated network of neural areas involved in executive attention. This network is associated with our voluntary ability to select among competing items, to correct error and to regulate our emotions. Recent neuroimaging studies suggest that these various functions involve separate areas of the anterior cingulate. We have adopted a strategy of using marker tasks, shown to activate the brain area by imaging studies, as a means of tracing the development of attentional networks. Executive attention appears to develop first to regulate distress during the first year of life. During later childhood the ability to regulate conflict among competing stimuli builds upon the earlier cingulate anatomy to provide a means of cognitive control. During childhood the activation of cingulate structures relates both to the child''s success on laboratory tasks involving conflict and to parental reports of self-regulation and emotional control. These studies indicate a start in understanding the anatomy, circuitry and development of executive attention networks that serve to regulate both cognition and emotion.     

Electrical Neuroimaging Reveals Timing of Attentional Control Activity in Human Brain

Voluntarily shifting attention to a location of the visual field improves the perception of events that occur there. Regions of frontal cortex are thought to provide the top-down control signal that initiates a shift of attention, but because of the temporal limitations of functional brain imaging, the timing and sequence of attentional-control operations remain unknown. We used a new analytical technique (beamformer spatial filtering) to reconstruct the anatomical sources of low-frequency brain waves in humans associated with attentional control across time. Following a signal to shift attention, control activity was seen in parietal cortex 100–200 ms before activity was seen in frontal cortex. Parietal cortex was then reactivated prior to anticipatory biasing of activity in occipital cortex. The magnitudes of early parietal activations were strongly predictive of the degree of attentional improvement in perceptual performance. These results show that parietal cortex, not frontal cortex, provides the initial signals to shift attention and indicate that top-down attentional control is not purely top down.     

Effect of psychostimulants on distinct attentional parameters in attentional deficit/hyperactivity disorder

Although there is extensive literature about the effects of stimulants on sustained attention tasks in attentional deficit/hyperactivity disorder (ADHD), little is known about the effect of these drugs on other attentional tasks involving different neural systems. In this study we measured the effect of stimulants on ADHD children, both in the electroencephalographic (EEG) activity during sustained attentional tasks and in psychometric performance during selective attentional tasks. These tasks are known to rely on different cortical networks. Our results in children medicated with 10 mg of d-amphetamine administered 60 min before the study indicate (i) a significant increase in amplitude but not latency of the P300 component of the event-related potential (ERP) during the sustained attentional task and (ii) a significant improvement in the reaction times and correct responses in the selective attentional task. In addition to supporting the use of stimulants in children with attentional deficit/hyperactivity disorder, these results show a multifocal activity improvement of cortical structures linked to dopamine, and interestingly, to attention. All these analyses are framed in a wider study of diverse attentional functions in this syndrome.     

Computational model of excitatory/inhibitory ratio imbalance role in attention deficit disorders

Impairments in attentional behaviors, including over-selectivity, under-selectivity, distractibility and difficulty in shift of attention, are widely reported in several developmental disorders, including autism. Uncharacteristic inhibitory to excitatory neuronal number ratio (IER) and abnormal synaptic strength levels in the brain are two broadly accepted neurobiological disorders observed in autistic individuals. These neurobiological findings are contrasting and their relation to the atypical attentional behaviors is not clear yet. In this paper, we take a computational approach to investigate the relation of imbalanced IER and abnormal synaptic strength to some well-documented spectrum of attentional impairments. The computational model is based on a modified version of a biologically plausible neural model of two competing minicolumns in IT cortex augmented with a simple model of top-down attention. Top-down attention is assumed to amplify (attenuates) attended (unattended) stimulus. The inhibitory synaptic strength parameter in the model is set such that typical attentional behavior is emerged. Then, according to related findings, the parameter is changed and the model's attentional behavior is considered. The simulation results show that, without any change in top-down attention, the abnormal inhibitory synaptic strength values - and IER imbalance- result in over-selectivity, under-selectivity, distractibility and difficulty in shift of attention in the model. It suggests that the modeled neurobiological abnormalities can be accounted for the attentional deficits. In addition, the atypical attentional behaviors do not necessarily point to impairments in top-down attention. Our simulations suggest that limited changes in the inhibitory synaptic strength and variations in top-down attention signal affect the model's attentional behaviors in the same way. So, limited deficits in the inhibitory strength may be alleviated by appropriate change in top-down attention biasing. Nevertheless, our model proposes that this compensation is not possible for very high and very low values of the inhibitory strength.     

Temporal Dynamics of Visual Attention Measured with Event-Related Potentials

How attentional modulation on brain activities determines behavioral performance has been one of the most important issues in cognitive neuroscience. This issue has been addressed by comparing the temporal relationship between attentional modulations on neural activities and behavior. Our previous study measured the time course of attention with amplitude and phase coherence of steady-state visual evoked potential (SSVEP) and found that the modulation latency of phase coherence rather than that of amplitude was consistent with the latency of behavioral performance. In this study, as a complementary report, we compared the time course of visual attention shift measured by event-related potentials (ERPs) with that by target detection task. We developed a novel technique to compare ERPs with behavioral results and analyzed the EEG data in our previous study. Two sets of flickering stimulus at different frequencies were presented in the left and right visual hemifields, and a target or distracter pattern was presented randomly at various moments after an attention-cue presentation. The observers were asked to detect targets on the attended stimulus after the cue. We found that two ERP components, P300 and N2pc, were elicited by the target presented at the attended location. Time-course analyses revealed that attentional modulation of the P300 and N2pc amplitudes increased gradually until reaching a maximum and lasted at least 1.5 s after the cue onset, which is similar to the temporal dynamics of behavioral performance. However, attentional modulation of these ERP components started later than that of behavioral performance. Rather, the time course of attentional modulation of behavioral performance was more closely associated with that of the concurrently recorded SSVEPs analyzed. These results suggest that neural activities reflected not by either the P300 or N2pc, but by the SSVEPs, are the source of attentional modulation of behavioral performance.     

视觉注意与神经振荡

人脑每时每刻都要接收大量视觉信息，由于人脑加工信息的能力有限，所以在较大视野内将注意分配给相关信息，同时抑制引起注意分散的不相关信息，对执行目标导向的行为至关重要。这种对视觉信息的选择性和主动性加工以适应当前目标的过程被称作视觉注意（visual attention），且视觉注意可分为自上而下的注意与自下而上的注意两种不同功能。由于来自大脑电信号的神经振荡活动在认知加工中发挥重要作用，已有研究综述了视觉注意与神经振荡（neural oscillation）的密切关系，但并未涉及不同的注意功能与神经振荡的关系。本文系统性调查了不同注意功能与神经振荡的关系，发现额-顶区域的theta频带振荡活动反映了自上而下的认知控制，而后部脑区的theta振荡与自下而上的注意相关。顶-枕区域alpha振荡的偏侧化有助于注意分配，而alpha频带的大规模同步促成了注意对视皮层自上而下的影响。Beta振荡介导了自上而下的信息与自下而上的信息之间的互动，作为信息载体促进了视觉信息处理。Gamma振荡则可能与自上而下和自下而上的注意间整合相关。本文就视觉注意功能与神经振荡关系的研究现状展开综述，旨在揭示不同的神经振荡活动在特定的视觉注意功能中的作用。     

Emotional and cognitive changes during adolescence

Adolescence is a critical period for maturation of neurobiological processes that underlie higher cognitive functions and social and emotional behavior. Recent studies have applied new advances in magnetic resonance imaging to increase understanding of the neurobiological changes that occur during the transition from childhood to early adulthood. Structural imaging data indicate progressive and regressive changes in the relative volumes of specific brain regions, although total brain volume is not significantly altered. The prefrontal cortex matures later than other regions and its development is paralleled by increased abilities in abstract reasoning, attentional shifting, response inhibition and processing speed. Changes in emotional capacity, including improvements in affective modulation and discrimination of emotional cues, are also seen during adolescence. Functional imaging studies using cognitive and affective challenges have shown that frontal cortical networks undergo developmental changes in processing. In summary, brain regions that underlie attention, reward evaluation, affective discrimination, response inhibition and goal-directed behavior undergo structural and functional re-organization throughout late childhood and early adulthood. Evidence from recent imaging studies supports a model by which the frontal cortex adopts an increasingly regulatory role. These neurobiological changes are believed to contribute, in part, to the range in cognitive and affective behavior seen during adolescence.     

The Pattern and Loci of Training-Induced Brain Changes in Healthy Older Adults Are Predicted by the Nature of the Intervention

There is enormous interest in designing training methods for reducing cognitive decline in healthy older adults. Because it is impaired with aging, multitasking has often been targeted and has been shown to be malleable with appropriate training. Investigating the effects of cognitive training on functional brain activation might provide critical indication regarding the mechanisms that underlie those positive effects, as well as provide models for selecting appropriate training methods. The few studies that have looked at brain correlates of cognitive training indicate a variable pattern and location of brain changes - a result that might relate to differences in training formats. The goal of this study was to measure the neural substrates as a function of whether divided attentional training programs induced the use of alternative processes or whether it relied on repeated practice. Forty-eight older adults were randomly allocated to one of three training programs. In the SINGLE REPEATED training, participants practiced an alphanumeric equation and a visual detection task, each under focused attention. In the DIVIDED FIXED training, participants practiced combining verification and detection by divided attention, with equal attention allocated to both tasks. In the DIVIDED VARIABLE training, participants completed the task by divided attention, but were taught to vary the attentional priority allocated to each task. Brain activation was measured with fMRI pre- and post-training while completing each task individually and the two tasks combined. The three training programs resulted in markedly different brain changes. Practice on individual tasks in the SINGLE REPEATED training resulted in reduced brain activation whereas DIVIDED VARIABLE training resulted in a larger recruitment of the right superior and middle frontal gyrus, a region that has been involved in multitasking. The type of training is a critical factor in determining the pattern of brain activation.