Visual attention: spotlight on the primary visual cortex.

Visual search tasks appear to involve spatially selective attention to the target, but evidence for attentional modulation in the visual area with the most precise retinotopic organization V1 has been elusive. Recent imaging studies show that spatial attention can indeed enhance visual responses in human V1.     

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.     

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.     

Dynamic shape integration in extrastriate cortex

BACKGROUND: In anorthoscopic viewing conditions, observers can perceive a moving object through a narrow slit even when only portions of its contour are visible at any time. We used fMRI to examine the contribution of early and later visual cortical areas to dynamic shape integration. Observers' success at integrating the shape of the slit-viewed object was manipulated by varying the degree to which the stimulus was dynamically distorted. Line drawings of common objects were either moderately distorted, strongly distorted, or shown undistorted. Phenomenologically, increasing the stimulus distortion made both object shape and motion more difficult to perceive.RESULTS: We found that bilateral cortical activity in portions of the ventral occipital cortex, corresponding to known object areas within the lateral occipital complex (LOC), was inversely correlated with the degree of stimulus distortion. We found that activity in left MT+, the human cortical area specialized for motion, showed a similar pattern as the ventral occipital region. The LOC also showed greater activity to a fully visible moving object than to the undistorted slit-viewed object. Area MT+, however, showed more equivalent activity to both the slit-viewed and fully visible moving objects.CONCLUSIONS: In early retinotopic cortex, the distorted and undistorted stimuli elicited the same amount of activity. Higher visual areas, however, were correlated with the percept of the coherent object, and this correlation suggests that the shape integration is mediated by later visual cortical areas. Motion information from the dorsal stream may project to the LOC to produce the shape percept.     

Interhemisphere differences of extrastriate cortical activation during attention and selection of lateralized visual stimuli in humans

The cortical activation was estimated by event-related potentials (ERPs) methods during selection tasks of lateralized visual stimuli in right and left hemi-fields requiring different forms of attention: 1. Attention of a stimuli form, 2. Attention of a stimuli position, 3. Combined attention of form and position. 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 complex [N1--P3]. The differences between ERPs at contra- and ipsilateral stimuli in the right and left hemispheres were considered as indices of asymmetry. The asymmetry was revealed in right hemisphere in all kinds of attention forms. The level (amplitude) of right-side asymmetry was depended on the level of attention: The significant relation between the right-side asymmetry and subjects' reaction time was also revealed. It is proposed that such an asymmetry is the evidence of better spatial differentiation of visual stimuli in right hemisphere in humans.     

Distribution of attention modulates salience signals in early visual cortex

Previous research has shown that the extent to which people spread attention across the visual field plays a crucial role in visual selection and the occurrence of bottom-up driven attentional capture. Consistent with previous findings, we show that when attention was diffusely distributed across the visual field while searching for a shape singleton, an irrelevant salient color singleton captured attention. However, while using the very same displays and task, no capture was observed when observers initially focused their attention at the center of the display. Using event-related fMRI, we examined the modulation of retinotopic activity related to attentional capture in early visual areas. Because the sensory display characteristics were identical in both conditions, we were able to isolate the brain activity associated with exogenous attentional capture. The results show that spreading of attention leads to increased bottom-up exogenous capture and increased activity in visual area V3 but not in V2 and V1.     

Specialized color modules in macaque extrastriate cortex

Imaging studies are consistent with the existence of brain regions specialized for color, but electrophysiological studies have produced conflicting results. Here we address the neural basis for color, using targeted single-unit recording in alert macaque monkeys, guided by functional magnetic resonance imaging (fMRI) of the same subjects. Distributed within posterior inferior temporal cortex, a large region encompassing V4, PITd, and posterior TEO that some have proposed functions as a single visual complex, we found color-biased fMRI hotspots that we call "globs," each several millimeters wide. Almost all cells located in globs showed strong luminance-invariant color tuning and some shape selectivity. Cells in different globs represented distinct visual field locations, consistent with the coarse retinotopy of this brain region. Cells in "interglob" regions were not color tuned, but were more strongly shape selective. Neither population was direction selective. These results suggest that color perception is mediated by specialized neurons that are clustered within the extrastriate brain.     

Effects of attention on the reliability of individual neurons in monkey visual cortex.

To determine the physiological mechanisms underlying the enhancement of performance by attention, we examined how attention affects the ability of isolated neurons to discriminate orientation by investigating the reliability of responses with and without attention. Recording from 262 neurons in cortical area V4 while two rhesus macaques did a delayed match-to-sample task with oriented stimuli, we found that attention did not produce detectable changes in the variability of neuronal responses but did improve the orientation discriminability of the neurons. We also found that attention did not change the relationship between burst rate and response rate. Our results are consistent with the idea that attention selects groups of neurons for a multiplicative enhancement in response strength.     

Phase locking of single neuron activity to theta oscillations during working memory in monkey extrastriate visual cortex

Working memory has been linked to elevated single neuron discharge in monkeys and to oscillatory changes in the human EEG, but the relation between these effects has remained largely unexplored. We addressed this question by measuring local field potentials and single unit activity simultaneously from multiple electrodes placed in extrastriate visual cortex while monkeys were performing a working memory task. We describe a significant enhancement in theta band energy during the delay period. Theta oscillations had a systematic effect on single neuron activity, with neurons emitting more action potentials near their preferred angle of each theta cycle. Sample-selective delay activity was enhanced if only action potentials emitted near the preferred theta angle were considered. Our results suggest that extrastriate visual cortex is involved in short-term maintenance of information and that theta oscillations provide a mechanism for structuring the recurrent interaction between neurons in different brain regions that underlie working memory.     

Activation of the human cortex during visual attention and selection

Cortical activation in visual discrimination tasks was estimated by measurement of the CNV (contingent negative variation) and N1-P3 components of visual ERPs in frontal, parietal, occipital and temporal leads recorded in 18 young healthy adults. In all investigated tasks, the maximal values of CNV and ERPa were observed in parietal regions. The estimation of cortical readiness state (CNV) is quite a useful procedure in the attention tasks because amplitude and stability of ERPs depend on preceding cortical excitability. The prevalence of parietal activation in visual attention tasks may be considered as the dominance of occipito-parietal way (stream) in human visual attention system.     

Selective attention and memory: evoked potentials in visual and auditory verbal competition

The brain mechanisms of the selective verbal attention were studied using evoked potential (EP) technique. It was shown that the late "cognitive" EP components (400-800 ms) related to memory function were more positive to the words presented via a relevant sensory channel and more negative to irrelevant words. The hypothesis is advanced that words delivered via two competing sensory channels, such as visual and auditory, are perceived, i.e., the subject sees and hears them. However, irrelevant signals are not stored in episodic memory due to the active inhibition of the information transmission to the hippocampal structures.     

Restoration of ocular dominance plasticity mediated by adenosine 3',5'-monophosphate in adult visual cortex.

Noradrenaline (NA)-stimulated beta-adrenoreceptors activate adenylate cyclase via excitatory G-proteins (Gs). Activated adenylate cyclase in turn promotes the production of cAMP. Critical roles of cAMP-dependent protein kinase A (PKA) in divergent cellular functions have been shown, including memory, learning and neural plasticity. Ocular dominance plasticity (ODP) is strongly expressed in early postnatal life and usually absent in the mature visual cortex. Here, we asked whether the activation of cAMP-dependent PKA could restore ODP to the aplastic visual cortex of adult cats. Concurrent with brief monocular deprivation, each of the following cAMP-related drugs was directly and continuously infused in the adult visual cortex: cholera toxin (a Gs-protein stimulant), forskolin (a Gs-protein-independent activator of adenylate cyclase) and dibutyryl cAMP (a cAMP analogue). We found that the ocular dominance distribution became W-shaped, the proportion of binocular cells being significantly lower than that in respective controls. We concluded that the activation of cAMP cascades rapidly restores ODP to the adult visual cortex, though moderately. The finding further extends the original hypothesis that the NA-beta-adrenoreceptors system is a neurochemical mechanism of cortical plasticity.     

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.     

Feature-based attention increases the selectivity of population responses in primate visual cortex

BACKGROUND: Attending to the spatial location or to nonspatial features of visual stimuli can modulate neuronal responses in primate visual cortex. The modulation by spatial attention changes the gain of sensory neurons and strengthens the representation of attended locations without changing neuronal selectivities such as directionality, i.e., the ratio of responses to preferred and anti-preferred directions of motion. Whether feature-based attention acts in a similar manner is unknown. RESULTS: To clarify this issue, we recorded the responses of 135 direction-selective neurons in the middle temporal area (MT) of two macaques to an unattended moving random dot pattern (the distractor) positioned inside a neuron's receptive field while the animals attended to a second moving pattern positioned in the opposite hemifield. Responses to different directions of the distractor were modulated by the same factor (approximately 12%) as long as the attended direction remained unchanged. On the other hand, systematically changing the attended direction from a neuron's preferred to its anti-preferred direction caused a systematic change of the attentional modulation from an enhancement to a suppression, increasing directionality by about 20%. CONCLUSIONS: The results show that (1) feature-based attention exerts a multiplicative modulation upon neuronal responses and that the strength of this modulation depends on the similarity between the attended feature and the cell's preferred feature, in line with the feature-similarity gain model, and (2) at the level of the neuronal population, feature-based attention increases the selectivity for attended features by increasing the responses of neurons preferring this feature value while decreasing responses of neurons tuned to the opposite feature value.     

Selective visual processing across competition episodes: a theory of task-driven visual attention and working memory

The goal of this review is to introduce a theory of task-driven visual attention and working memory (TRAM). Based on a specific biased competition model, the ‘theory of visual attention’ (TVA) and its neural interpretation (NTVA), TRAM introduces the following assumption. First, selective visual processing over time is structured in competition episodes. Within an episode, that is, during its first two phases, a limited number of proto-objects are competitively encoded—modulated by the current task—in activation-based visual working memory (VWM). In processing phase 3, relevant VWM objects are transferred via a short-term consolidation into passive VWM. Second, each time attentional priorities change (e.g. after an eye movement), a new competition episode is initiated. Third, if a phase 3 VWM process (e.g. short-term consolidation) is not finished, whereas a new episode is called, a protective maintenance process allows its completion. After a VWM object change, its protective maintenance process is followed by an encapsulation of the VWM object causing attentional resource costs in trailing competition episodes. Viewed from this perspective, a new explanation of key findings of the attentional blink will be offered. Finally, a new suggestion will be made as to how VWM items might interact with visual search processes.     

Spatial attention changes excitability of human visual cortex to direct stimulation

Conscious perception depends not only on sensory input, but also on attention [1, 2]. Recent studies in monkeys [3-6] and humans [7-12] suggest that influences of spatial attention on visual awareness may reflect top-down influences on excitability of visual cortex. Here we tested this specifically, by providing direct input into human visual cortex via cortical transcranial magnetic stimulation (TMS) to produce illusory visual percepts, called phosphenes. We found that a lower TMS intensity was needed to elicit a conscious phosphene when its apparent spatial location was attended, rather than unattended. Our results indicate that spatial attention can enhance visual-cortex excitability, and visual awareness, even when sensory signals from the eye via the thalamic pathway are bypassed.     

Spatiotopic coding of BOLD signal in human visual cortex depends on spatial attention

The neural substrate of the phenomenological experience of a stable visual world remains obscure. One possible mechanism would be to construct spatiotopic neural maps where the response is selective to the position of the stimulus in external space, rather than to retinal eccentricities, but evidence for these maps has been inconsistent. Here we show, with fMRI, that when human subjects perform concomitantly a demanding attentive task on stimuli displayed at the fovea, BOLD responses evoked by moving stimuli irrelevant to the task were mostly tuned in retinotopic coordinates. However, under more unconstrained conditions, where subjects could attend easily to the motion stimuli, BOLD responses were tuned not in retinal but in external coordinates (spatiotopic selectivity) in many visual areas, including MT, MST, LO and V6, agreeing with our previous fMRI study. These results indicate that spatial attention may play an important role in mediating spatiotopic selectivity.     

Color coding in the cortex: a modified approach to bottom-up visual attention

Itti and Koch’s (Vision Research 40:1489–1506, 2000) saliency-based visual attention model is a broadly accepted model that describes how attention processes are deployed in the visual cortex in a pure bottom-up strategy. This work complements their model by modifying the color feature calculation. Evidence suggests that S-cone responses are elicited in the same spatial distribution and have the same sign as responses to M-cone stimuli; these cells are tentatively referred to as red-cyan. For other cells, the S-cone input seems to be aligned with the L-cone input; these cells might be green-magenta cells. To model red-cyan and green-magenta double-opponent cells, we implement a center-surround difference approach of the aforementioned model. The resulting color maps elicited enhanced responses to color salient stimuli when compared to the classic ones at high statistical significance levels. We also show that the modified model improves the prediction of locations attended by human viewers.     

Computations in the early visual cortex.

This paper reviews some of the recent neurophysiological studies that explore the variety of visual computations in the early visual cortex in relation to geometric inference, i.e. the inference of contours, surfaces and shapes. It attempts to draw connections between ideas from computational vision and findings from awake primate electrophysiology. In the classical feed-forward, modular view of visual processing, the early visual areas (LGN, V1 and V2) are modules that serve to extract local features, while higher extrastriate areas are responsible for shape inference and invariant object recognition. However, recent findings in primate early visual systems reveal that the computations in the early visual cortex are rather complex and dynamic, as well as interactive and plastic, subject to influence from global context, higher order perceptual inference, task requirement and behavioral experience. The evidence argues that the early visual cortex does not merely participate in the first stage of visual processing, but is involved in many levels of visual computation.     

Visual search and event-related potentials in the extrastriate cortical areas in humans

Parameters of visual search (reaction time, mistakes) were investigated in 11 young healthy volunteers, under changing parameters of target symbol (form, color and location) in surroundings of white distracters (flankers). In 6 cortical leads: P3, P4, T3, T4, T5, T6, monopolar event-related potentials (ERPs) were registered and analyzed for the late endogenous components: N2, P3, because these very components have changed under verifying of search (so called late selection). In the hard search (similar target and distractors) the increase of search time was accompanied by the delay of P3 component and diminishing of its amplitude. The definite (the knowledge of) target position resulted by the decrease search time and decrease of P3 wave latency as compared with situation of uncertainty in respect to the position of target. Alteration of the target stimulus colour were led to abolishment (cancel) of inhibition effect of distracters: parameters of IRP have not differed from parameters of ERPs to solid target stimulus without the flankers. A high correlation of search time and parameters of P3 wave was revealed. Changes of IRPs in different kinds of search were similar in parietal and temporal leads. This suggests that the parietal and temporal cortical areas function as united system, under search of target in environment.