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The attentive homunculus: now you see it, now you don't.
The nature of the neural system that directs our attention toward selective items in the extrapersonal world is a longstanding and interesting puzzle. The ability to image the human brain at work non-invasively using positron-emission tomography or functional magnetic resonance has provided the means to investigate this issue. In this article, I review the contributions of brain imaging toward the characterization of attentional control in the human brain. The majority of experiments to date have investigated visual spatial orienting. A consistent pattern of brain areas has been revealed, comprising most notably the posterior parietal cortex around the intraparietal sulcus and frontal regions including the frontal eye fields. The brain areas implicated in the control of visual spatial attention were noted to resemble those involved in the control of eye movements, and direct experimental comparisons supported a tight link between the two systems. The findings suggested a sensible view of the attentional 'homunculus' as a distributed neural system related to the control of eye movements. Eye movements form perhaps the most basic orienting response, and can be shifted rapidly and efficiently based on multiple frames of reference. Some attention experiments using objects and features instead of spatial locations as the target of selection also obtained similar patterns of parietal-frontal activations, rendering further support to this view of the attentional control system. Some recent experiments, however, have cautioned against a premature conclusion regarding the ubiquity of the attentional control system revealed by studies of visual spatial attention. Different parietal and frontal regions become engaged when attention is shifted along non-spatial dimensions, such as when attention is directed toward a particular motor act or toward a specific point in time. In these cases, the neural system resembles those involved in the control of limb movements. The attentional homunculus thus begins to dissolve. The alternative view suggested is that attentional control may be a property of specialized parietal-frontal systems that transform perception into action. Future studies will be needed to validate this view of attention, or to provide a more mature understanding of its true nature.
Orienting attention in time. Modulation of brain potentials.
With the aim of casting light on the neural mechanisms that support our ability to modulate visual attention over time, we recorded event-related potentials (ERPs) while normal human subjects performed a target detection task with temporal contingencies between cue and target stimuli. The task used two central cues, which predicted (80% validity) when a subsequent target would occur (either 600 or 1400 ms after cue onset). Unlike previous tasks of attentional orienting, there was no spatial information provided and all stimuli were presented foveally. Reaction times and ERPs linked to targets presented at the shorter interval showed significant effects linked to attentional orienting. Reaction times were faster when the cues correctly predicted the cue-target interval, suggesting the ability of the brain to use information about time to deploy attentional resources. ERPs differed according to the predicted time interval. In particular, the P300 amplitude and latency were enhanced when the cue predicted the cue-target interval accurately. The ERPs elicited by the cues also differed according to the time interval that they predicted. Differences were observed in potentials linked to motor preparation and expectancies. The results reveal dynamic neural activity involved in orienting attention to time intervals, as well as the consequent modulation of target-related neural activity resulting from differing temporal expectations.
Functional magnetic resonance imaging of sensory and motor cortex: comparison with electrophysiological localization.
Functional magnetic resonance (MR) imaging was performed using a 1.5-tesla MR system to localize sensorimotor cortex. Six neurologically normal subjects were studied by means of axial gradient-echo images with a motor task and one or more sensory tasks: 1) electrical stimulation of the median nerve; 2) continuous brushing over the thenar region; and 3) pulsed flow of compressed air over the palm and digits. An increased MR signal was observed in or near the central sulcus, consistent with the location of primary sensory and motor cortex. Four patients were studied using echo planar imaging sequences and motor and sensory tasks. Three patients had focal refractory seizures secondary to a lesion impinging on sensorimotor cortex. Activation seen on functional MR imaging was coextensive with the location of the sensorimotor area determined by evoked potentials and electrical stimulation. Functional MR imaging provides a useful noninvasive method of localization and functional assessment of sensorimotor cortex.
Qualitative mapping of cerebral blood flow and functional localization with echo-planar MR imaging and signal targeting with alternating radio frequency.
PURPOSE: To create qualitative maps of cerebral blood flow (CBF) with the EPISTAR (echo-planar imaging and signal targeting with alternating radio frequency) technique. MATERIALS AND METHODS: The EPISTAR technique was performed in a pig model of hypercapnia and then tested in 26 volunteers by using various paradigms for cortical activation. Echo-planar images were acquired with and without use of a radio-frequency inversion pulse applied to inflowing arterial spins. A qualitative map of CBF was then created by subtracting the image obtained without the radio-frequency pulse from that obtained with the radio-frequency pulse. RESULTS: Progressively more distal portions of the tagged vessels were seen as the inflow time was lengthened until cortical enhancement was seen for inflow times of approximately 1 second or longer. Signal intensity increases from rest to sensorimotor activation ranged from 13% to 193%. CBF changes in the motor strip, primary visual cortex, and the motor area for eye movements were well localized to the cortical gray matter ribbon. CONCLUSION: The EPISTAR technique is a rapid, noninvasive means for creating qualitative maps of CBF.
Modulation of neural activity by motivational and spatial biases.
Motivational biases and spatial attention both modulate neural activity and influence behavioural performance. The time course of motivational bias effects, as well as the relationship between motivation and attention across the time course of information processing, however, are relatively unknown. In the present study, event-related potentials (ERPs) were recorded whilst individuals performed a modified Posner task, in which cue stimuli indicated the reward stakes of a given trial and the probable spatial location of a subsequent target stimulus. Reaction times (RTs) were sensitive to motivation and to attention, with faster responses produced on valid and on rewarded trials. In addition, motivation modulated neural activity from the visual analysis of stimuli, with an earlier N1 peak for rewarded compared with non-rewarded stimuli. Effects of motivation were relatively independent from those of attention until late cognitive processing and response production, where motivation and attention interacted to enhance P300-like potentials and the lateralised readiness potential (LRP). The results suggest that multiple sources of modulatory influences may exist, with motivation and attention exerting independent influences over early stimulus and cognitive processing, followed by a late interaction allowing the construction of a comprehensive stimulus representation that contains information pertaining to both motivational and spatial expectations.
Distinct neural substrates for visual search amongst spatial versus temporal distractors.
Whether the contribution of the superior parietal cortex (BA7) to attention-demanding tasks is strictly spatial in nature remains unresolved. We used functional magnetic resonance imaging to explore the behavioural and neuroanatomical correlates of non-spatial search for a conjunction of features within a stream of temporally-distracting stimuli. In addition, we compared these data to those from a conventional visuo-spatial search task, performed by the same subjects, in order to determine the specificity of right BA7 activation. Mode of stimulus-distribution (spatial versus temporal) and search type (target defined by a single feature or a conjunction of features) were manipulated in a 2 x 2 factorial design. Behaviourally, the temporal conjunction task was shown to index temporal selective attention. Accuracy of detecting a second target varied with the temporal proximity of two successive targets when subjects searched for a conjunction of features, but not a single feature. The temporal conjunction task activated a network of areas including right superior parietal cortex and bilateral regions of intraparietal sulcus, frontal operculum and putamen. The two latter regions were selectively activated by the attentional demands of the temporal conjunction task when compared directly to the attentional demands of the spatial conjunction task, implicating these regions specifically in selective attention among temporally-distracting stimuli. By comparison, only a very medial region of right BA7 was selectively activated by the spatial conjunction task. The more lateral region of BA7 previously reported by other groups was engaged to a similar degree by both spatial and temporal versions of the conjunction search task.
Modulation of human extrastriate visual processing by selective attention to colours and words.
The present study investigated the effect of visual selective attention upon neural processing within functionally specialized regions of the human extrastriate visual cortex. Field potentials were recorded directly from the inferior surface of the temporal lobes in subjects with epilepsy. The experimental task required subjects to focus attention on words from one of two competing texts. Words were presented individually and foveally. Texts were interleaved randomly and were distinguishable on the basis of word colour. Focal field potentials were evoked by words in the posterior part of the fusiform gyrus. Selective attention strongly modulated long-latency potentials evoked by words. The attention effect co-localized with word-related potentials in the posterior fusiform gyrus, and was independent of stimulus colour. The results demonstrated that stimuli receive differential processing within specialized regions of the extrastriate cortex as a function of attention. The late onset of the attention effect and its co-localization with letter string-related potentials but not with colour-related potentials recorded from nearby regions of the fusiform gyrus suggest that the attention effect is due to top-down influences from downstream regions involved in word processing.
Purely endogenous capture of attention by task-defining features proceeds independently from spatial attention.
Attention can be focused voluntarily and effectively on spatial locations in order to enhance the processing of task-relevant events. However, work on 'attentional capture' has demonstrated that spatial biases can be temporarily reset by transient and salient stimuli, especially if they share defining characteristics with the targets of a task goal. In the current study, we investigated whether the appearance of stimuli containing task-defining features at an unattended location was sufficient to capture attention, even when these were not perceptually salient. We used event-related-potential (ERP) markers to test whether the selection of task-defining features was modulated by top-down spatial attention, and to test whether the appearance of 'unattended targets' transiently disrupted the spatial bias. Surprisingly, the results revealed that ERP markers of selection of task-defining features were equivalent for stimuli appearing at spatially attended and unattended locations. In addition, the presentation of task-defining stimuli at the spatially unattended location induced a short-lived redistribution of the pre-established spatial attention bias toward the 'capture' side. These findings show that task-defining features of a stimulus are automatically processed independently from spatial attention, and suggest the co-existence of multiple sources of top-down biasing signals, which might in part sustain the capture mechanism.
The effects of combined caffeine and glucose drinks on attention in the human brain.
The objective of this research was to measure the effects of energising drinks containing caffeine and glucose, upon mental activity during sustained selective attention. Non-invasive electrophysiological brain recordings were made during a behavioural study of selective attention in which participants received either energising or placebo drinks. We tested specifically whether energising drinks have significant effects upon behavioural measures of performance during a task requiring sustained visual selective attention, as well as on accompanying components of the event-related potential (ERPs) related to information processing in the brain. Forty healthy volunteers were blindly assigned to receive either the energising drink or a similar-tasting placebo drink. The behavioural task involved identifying predefined target stimulus among rapidly presented streams of peripheral visual stimuli, and making speeded motor responses to this stimulus. During task performance, accuracy, reaction times and ongoing brain activity were stored for analysis. The energising drink enhanced behavioural performance both in terms of accuracy and speed of reactions. The energising drink also had significant effects upon the event-related potentials. Effects started from the enhancement of the earliest components (Cl/P1), reflecting early visual cortical processing in the energising-drink group relative to the placebo group over the contralateral scalp. The later N1, N2 and P3 components related to decision-making and responses were also modulated by the energising drink. Energising drinks containing caffeine and glucose can enhance behavioural performance during demanding tasks requiring selective attention. The behavioural benefits are coupled to direct effects upon neural information processing.
Modulation of working-memory maintenance by directed attention.
Many current models of working memory (WM) emphasize a close relationship between WM and attention. Recently it was demonstrated that attention can be dynamically and voluntarily oriented to items held in WM, and it was suggested that directed attention can modulate the maintenance of specific WM representations. Here we used event-related functional magnetic resonance imaging to test the effects of orienting attention to a category of stimuli when participants maintained a variable number of faces and scenes in WM. Retro-cues that indicated the relevant stimulus type for the subsequent WM test modulated maintenance-related activity in extrastriate areas preferentially responsive to face or scene stimuli - fusiform and parahippocampal gyri respectively - in a categorical way. After the retro-cue, the activity level in these areas was larger for the cued category in a load-independent way, suggesting the modulation may also reflect anticipation of the probe stimulus. Activity in associative parietal and prefrontal cortices was also modulated by retro-cues, and additionally co-varied with the number of stimuli of the relevant stimulus category that was being maintained. The findings suggest that these associative areas participate in maintaining the relevant memoranda in a flexible and goal-directed way to guide future behaviour.
Differential modulation of word recognition by semantic and spatial orienting of attention.
In the present study, we investigated the ability to orient attention to abstract associative features of complex stimuli, more specifically, to the semantic categories of visual word stimuli. We compared the behavioral and electrophysiological effects of semantic orienting with those elicited by spatial orienting to word stimuli. Two parallel, cued lexical-decision tasks, with semantic- or spatial-orienting cues, were used. Results showed that both semantic and spatial orienting facilitated behavioral performance. The event-related potential analysis revealed different and non-overlapping patterns of modulation of word processing by semantic and spatial orienting. Modulation by semantic orienting started later, affecting only the potentials linked to conceptual or semantic processing (N300 and N400). The pattern of N300/N400 modulation in the semantic-orienting condition was similar to that observed in semantic-priming tasks, and was compatible with the operation of controlled semantic processes. Spatial orienting significantly enhanced the amplitude of the early visual potential P1 as well as the language-related N400 potential. These findings showed that the similar end-result of behavioral facilitation by semantic and spatial orienting is achieved through largely distinct mechanisms acting upon separate levels of stimulus analysis.
Sub-second "temporal attention" modulates alpha rhythms. A high-resolution EEG study.
In the present high-resolution electroencephalographic (EEG) study, event-related desynchronization/synchronization (ERD/ERS) of alpha rhythms was computed during an S1-S2 paradigm, in which a visual cue (S1) predicted a SHORT (600 ms) or LONG (1400 ms) foreperiod, preceding a visual go stimulus (S2) triggering right or left finger movement. Could orienting attention to a selective point in time influence the alpha rhythms as a function of the SHORT vs. LONG foreperiod? Stronger selective attentional modulations were predicted for the SHORT than LONG condition. EEG data from 54 channels were "depurated" from phase-locked visual evoked potentials and spatially enhanced by surface Laplacian estimation (i.e., final data analysis was conducted on 16 subjects having a sufficient number of artifact-free EEG single trials). Low-band alpha rhythms (about 6-10 Hz) were supposed to be related to anticipatory attentional processes, whereas high-band alpha rhythms (10-12 Hz) would indicate task-specific visuo-motor processes. Compared to the LONG condition (foreperiod), the SHORT condition induced a quicker and stronger ERS at low-band alpha rhythm (about 6-8 Hz) over midline and bilateral prefrontal, sensorimotor, and posterior parietal areas. In contrast, the concomitant high-band alpha (about 10-12 Hz) ERD/ERS showed no significant difference between the two conditions. In conclusion, temporal attention for a sub-second delay (800 ms) did modulate low-band alpha rhythm over large regions of both cortical hemispheres.
Modulation of semantic processing by spatial selective attention.
The effects of spatial selective attention upon ERPs associated with the processing of word stimuli were investigated. While subjects maintained central eye fixation, ERPs were recorded to words presented to the left and right visual fields. In each of 6 runs, subjects focussed attention to alternate fields to perform a category-detection task. Pairs of semantically related and repeated words were embedded in the word lists presented to the attended and unattended visual fields. Consistent with prior studies, the P1-N1 visual ERP was larger when elicited by words in attended spatial locations. A large negative slow wave identified as N400 was elicited by attended, but not unattended, words. For attended words, N400 was smaller for semantically primed or repeated words. We concluded that spatial selective attention can modulate the degree to which words are processed, and that the cognitive processes associated with N400 are not automatic.
Dissociating explicit timing from temporal expectation with fMRI.
Explicit timing is engaged whenever subjects make a deliberate estimate of discrete duration in order to compare it with a previously memorised standard. Conversely, implicit timing is engaged, even without a specific instruction to time, whenever sensorimotor information is temporally structured and can be used to predict the duration of future events. Both emergent timing (motor) and temporal expectation (perceptual) are forms of implicit timing. Recent fMRI studies demonstrate discrete neural substrates for explicit and implicit timing. Specifically, basal ganglia are activated almost invariably by explicit timing, with co-activation of prefrontal, premotor and cerebellar areas being more context-dependent. Conversely, implicit perceptual timing (or "temporal expectation") recruits cortical action circuits, comprising inferior parietal and premotor areas, highlighting its role in the optimisation of prospective behaviour.
Selective attention to specific features within objects: behavioral and electrophysiological evidence.
Evidence regarding the ability of attention to bias neural processing at the level of single features has been gathering steadily, but most of the experiments to date used arrays with multiple objects and locations, making it difficult to rule out indirect influences from object or spatial attention. To investigate feature-specific selective attention, we have assessed the ability to select and ignore individual features within the same object. We used a negative-priming paradigm in which the color or the direction of internal motion of the object could determine the relevant response. Bidimensional (colored and moving) and unidimensional (colored and stationary, or gray and moving) stimuli appeared in unpredictable order. In successive blocks, participants were instructed that one feature dimension was dominant. During that block, participants responded according to the dominant dimension for bidimensional stimuli. For unidimensional stimuli, participants responded to the only dimension of the stimulus that afforded a response, regardless of the instruction for the block. The ability to inhibit irrelevant task information at the level of specific features (negative priming for features) was indexed by a decrease in performance to detect one particular feature value (e.g., red) if the same feature value (red) but not another color value (green) had been ignored in the previous bidimensional stimulus. Behavioral results confirmed the existence of inhibitory, negative-priming mechanisms at the single-feature level for both color and motion dimensions of stimuli. Event-related potentials recorded during task performance revealed the dynamics of neural modulation by feature attention. Comparisons were made using the identical physical stimuli under different conditions of attention to isolate purely attentional effects. Processing of identical bidimensional stimuli was compared as a function of the dimension of attention (color, motion). Processing of identical unidimensional stimuli that followed bidimensional stimuli was also compared to identify possible effects of feature-specific negative priming. The electrophysiological effects revealed that inhibition of irrelevant features leads to modulation of brain activity during early stages of perceptual analysis.
Orienting attention to locations in perceptual versus mental representations.
Extensive clinical and imaging research has characterized the neural networks mediating the adaptive distribution of spatial attention. In everyday behavior, the distribution of attention is guided not only by extrapersonal targets but also by mental representations of their spatial layout. We used event-related functional magnetic resonance imaging to identify the neural system involved in directing attention to locations in arrays held as mental representations, and to compare it with the system for directing spatial attention to locations in the external world. We found that these two crucial aspects of spatial cognition are subserved by extensively overlapping networks. However, we also found that a region of right parietal cortex selectively participated in orienting attention to the extrapersonal space, whereas several frontal lobe regions selectively participated in orienting attention within on-line mental representations.
The neural system of language: structure and development.
Recent neuroimaging and neuropsychological research in adults and infants suggests that the neural system for language is widely distributed and shares organizational principles with other cognitive systems in the brain. Connectionist modelling has clarified that networks operating with associative mechanisms can display properties typically associated with genetically predetermined and dedicated symbolic functions.
The posterior cingulate and medial prefrontal cortex mediate the anticipatory allocation of spatial attention.
The purpose of this study was to identify brain regions underlying internally generated anticipatory biases toward locations where significant events are expected to occur. Subjects fixated centrally and responded to peripheral targets preceded by a spatially valid (predictive), invalid (misleading), or neutral central cue while undergoing fMRI scanning. In some validly cued trials, reaction time was significantly shorter than in trials with neutral cues, indicating that the cue had successfully induced a spatial redistribution of motivational valence, manifested as expectancy. The largest cue benefits led to selectively greater activations within the posterior cingulate and medial prefrontal cortex. These two areas thus appear to establish a neural interface between attention and motivation. An inverse relationship to cue benefit was seen in the parietal cortex, suggesting that spatial expectancy may entail the inhibition of attention-related areas to reduce distractibility by events at irrelevant locations.
Covert visual spatial orienting and saccades: overlapping neural systems.
We used functional magnetic resonance imaging (fMRI) to investigate the functional anatomical relationship between covert orienting of visual spatial attention and execution of saccadic eye movements. Brain areas engaged by shifting spatial attention covertly and by moving the eyes repetitively toward visual targets were compared and contrasted directly within the same subjects. The two tasks activated highly overlapping neural systems and showed that common parietal and frontal regions are more activated during the covert task than the overt oculomotor condition. The possible nature of the relationship between these two operations is discussed.
Functional localization of the system for visuospatial attention using positron emission tomography.
PET was used to image the neural system underlying visuospatial attention. Analysis of data at both the group and individual-subject level provided anatomical resolution superior to that described to date. Six right-handed male subjects were selected from a pilot behavioural study in which behavioural responses and eye movements were recorded. The attention tasks involved covert shifts of attention, where peripheral cues indicated the location of subsequent target stimuli to be discriminated. One attention condition emphasized reflexive aspects of spatial orientation, while the other required controlled shifts of attention. PET activations agreed closely with the cortical regions recently proposed to form the core of a neural network for spatial attention. The two attention tasks evoked largely overlapping patterns of neural activation, supporting the existence of a general neural system for visuospatial attention with regional functional specialization. Specifically, neocortical activations were observed in the right anterior cingulate gyrus (Brodmann area 24), in the intraparietal sulcus of right posterior parietal cortex, and in the mesial and lateral premotor cortices (Brodmann area 6).