Top-down dimensional weight set determines the capture of visual attention: evidence from the PCN component

Visual search for feature singletons is slowed when a task-irrelevant, but more salient distracter singleton is concurrently presented. While there is a consensus that this distracter interference effect can be influenced by internal system settings, it remains controversial at what stage of processing this influence starts to affect visual coding. Advocates of the "stimulus-driven" view maintain that the initial sweep of visual processing is entirely driven by physical stimulus attributes and that top-down settings can bias visual processing only after selection of the most salient item. By contrast, opponents argue that top-down expectancies can alter the initial selection priority, so that focal attention is "not automatically" shifted to the location exhibiting the highest feature contrast. To precisely trace the allocation of focal attention, we analyzed the Posterior-Contralateral-Negativity (PCN) in a task in which the likelihood (expectancy) with which a distracter occurred was systematically varied. Our results show that both high (vs. low) distracter expectancy and experiencing a distracter on the previous trial speed up the timing of the target-elicited PCN. Importantly, there was no distracter-elicited PCN, indicating that participants did not shift attention to the distracter before selecting the target. This pattern unambiguously demonstrates that preattentive vision is top-down modifiable.     

Brain mechanisms mediating auditory attentional capture in humans

The ability to detect and preferentially process salient auditory stimuli, even when irrelevant to a current task, is often critical for adaptive behavior. This stimulus-driven allocation of processing resources is known as "attentional capture." Here we used functional magnetic resonance imaging in humans to investigate brain activity and behavioral effects related to such auditory attentional capture. Participants searched a sequence of tones for a target tone that was shorter or longer than the nontarget tones. An irrelevant singleton feature in the tone sequence resulted in behavioral interference (attentional capture) and activation of parietal and prefrontal cortices only when the singleton was associated with a nontarget tone (nontarget singleton) and not when associated with a target tone (target singleton). In contrast, the presence (vs. absence) of a singleton feature in the sequence was associated with activation of frontal and temporal loci previously associated with auditory change detection. These results suggest that a ventral network involving superior temporal and inferior frontal cortices responds to acoustic variability, regardless of attentional significance, but a dorsal frontoparietal network responds only when a feature singleton captures attention.     

Neural basis for priming of pop-out during visual search revealed with fMRI

Maljkovic and Nakayama first showed that visual search efficiency can be influenced by priming effects. Even "pop-out" targets (defined by unique color) are judged quicker if they appear at the same location and/or in the same color as on the preceding trial, in an unpredictable sequence. Here, we studied the potential neural correlates of such priming in human visual search using functional magnetic resonance imaging (fMRI). We found that repeating either the location or the color of a singleton target led to repetition suppression of blood oxygen level-dependent (BOLD) activity in brain regions traditionally linked with attentional control, including bilateral intraparietal sulci. This indicates that the attention system of the human brain can be "primed," in apparent analogy to repetition-suppression effects on activity in other neural systems. For repetition of target color but not location, we also found repetition suppression in inferior temporal areas that may be associated with color processing, whereas repetition of target location led to greater reduction of activation in contralateral inferior parietal and frontal areas, relative to color repetition. The frontal eye fields were also implicated, notably when both target properties (color and location) were repeated together, which also led to further BOLD decreases in anterior fusiform cortex not seen when either property was repeated alone. These findings reveal the neural correlates for priming of pop-out search, including commonalities, differences, and interactions between location and color repetition. fMRI repetition-suppression effects may arise in components of the attention network because these settle into a stable "attractor state" more readily when the same target property is repeated than when a different attentional state is required.     

Driven to less distraction: rTMS of the right parietal cortex reduces attentional capture in visual search

In visual search, the presence of a highly salient color singletoncan slow or facilitate search for a shape target depending onwhether the singleton is a distractor or coincides with thetarget. This is consistent with an attentional shift (attentionalcapture) to the salient item. This attentional capture can bedriven by bottom–up or top–down processes or both.We investigated the role of the parietal cortex in attentionalcapture by a singleton using repetitive transcranial magneticstimulation. Following disruption to the right posterior parietalcortex by sustained transcranial magnetic stimulation, the reactiontime (RT) cost of the singleton distractor was reduced. At leastpart of this lessening of singleton distraction was due to theelimination of priming (top–down) effects between targetand distractor singletons on consecutive trials. In Experiment2, we presented the different conditions in separate blocksmeaning any effects of the distractor can most likely be attributedto bottom–up processes. Nevertheless, there was stilla decrease in RT interference from the distractor so that areduction in priming cannot provide a full account of the results.The data are consistent with previous work positing that theright parietal cortex directs attention to salient stimuli (e.g.,Constantinidis 2005, Mevorach et al. 2006), while also suggestinga role for the right parietal cortex in the integration of bottom–upsalience information with memories for salient features on priortrials.     

The interaction of brain regions during visual search processing as revealed by transcranial magnetic stimulation

Although it has long been known that right posterior parietal cortex (PPC) has a role in certain visual search tasks, and human motion area V5 is involved in processing tasks requiring attention to motion, little is known about how these areas may interact during the processing of a task requiring the speciality of each. Using transcranial magnetic stimulation (TMS), this study first established the specialization of each area in the form of a double dissociation; TMS to right PPC disrupted processing of a color/form conjunction and TMS to V5 disrupted processing of a motion/form conjunction. The key finding of this study is, however, if TMS is used to disrupt processing of V5 at its critical time of activation during the motion/form conjunction task, concurrent disruption of right PPC now has a significant effect, where TMS at PPC alone does not. Our findings challenge the conventional interpretation of the role of right PPC in conjunction search and spatial attention.     

Cortical mechanisms of feature-based attentional control

A network of fronto-parietal cortical areas is known to be involved in the control of visual attention, but the representational scope and specific function of these areas remains unclear. Recent neuroimaging evidence has revealed the existence of both transient (attention-shift) and sustained (attention-maintenance) mechanisms of space-based and object-based attentional control. Here we investigate the neural mechanisms of feature-based attentional control in human cortex using rapid event-related functional magnetic resonance imaging (fMRI). Subjects viewed an aperture containing moving dots in which dot color and direction of motion changed once per second. At any given moment, observers attended to either motion or color. Two of six motion directions and two of six colors embedded in the stimulus stream cued subjects either to shift attention from the currently attended to the unattended feature or to maintain attention on the currently attended feature. Attentional modulation of the blood oxygenation level dependent (BOLD) fMRI signal was observed in early visual areas that are selective for motion and color. More importantly, both transient and sustained BOLD activity patterns were observed in different fronto-parietal cortical areas during shifts of attention. We suggest these differing temporal profiles reflect complementary roles in the control of attention to perceptual features.     

How many maps are there in visual cortex?

In addition to a topographic map of the retina, mammalian visual cortex contains superimposed, orderly periodic maps of features such as orientation, eye dominance, direction of motion and spatial frequency. There is evidence that these maps are overlaid so as to ensure that all combinations of the different parameters are represented as uniformly as possible across visual space. However, it is unknown to what extent geometrical factors limit the number of periodic maps which might simultaneously be present, given this constraint. This paper attempts to investigate the question by using a dimension reduction model to generate maps of simple, many- dimensional feature spaces onto a model two-dimensional cortex. The feature space included a model retina, plus N binary variables, corresponding to parameters such as ocular dominance or spatial frequency. The results suggest that geometrical factors do not sharply limit the ability of the cortex to represent combinations of parameters in spatially superimposed maps of similar periodicity. Considerations of uniform coverage suggest an upper limit of six or seven maps. A higher limit, of about nine or ten, may be imposed by the numbers of neurons (or minicolumns) available to represent each of 2(N) features within a given small region of cortex.     

Two attentional processes in the parietal lobe

We report fMRI evidence for two attentional processes in parietal cortex. Subjects matched a feature, cued by a word, to a test display of moving colored dots. Either color (red, green) or motion direction (left, right) was cued on mixed scans while only one dimension was cued on blocked scans. An event-related paradigm separated the preparatory activity generated by the cue from the subsequent activity related to the test display. One attentional process specified task information while a second process was motion selective. During the cue period, a pure effect of task specification was observed in left frontal cortex while combined effects of task specification and motion selectivity were observed in left posterior parietal cortex. The frontal task-specification signal may have been the source of the corresponding signal in parietal cortex. Effects of task specification generalized over cue dimension, indicating that the information was coded in a sufficiently abstract form to affect color and motion processing. During the subsequent test period, task-specification and motion-selective signals were again observed in left parietal cortex. Task specification did not significantly affect occipital motion-selective regions, such as MT+, however, indicating that this process did not influence the lower cortical tier of the motion processing stream. These results provide evidence for general and specialized task representations within left parietal cortex during task preparation and execution.     

Spatio-temporal analysis of feature-based attention

The cortical mechanisms of feature-selective attention to color and motion cues were studied in humans using combined electrophysiological, magnetoencephalographic, and hemodynamic (functional magnetic resonance imaging) measures of brain activity. Subjects viewed a display of random dots that periodically either changed color or moved coherently. When attention was directed to the color change it elicited enhanced neural activity in visual area V4v, previously shown to be specialized for processing color information. In contrast, when dot movement was attended it produced enhanced activity in the motion-specialized area human MT. Parallel recordings of event-related electrophysiological and magnetoencephalographic responses indicated that the attention-related facilitation of neural activity in these specialized cortical areas occurred rapidly, beginning as early as 90-120 ms after stimulus onset. We conclude that selection of an entire feature dimension (motion or color) boosts neural activity in its specialized cortical module much more rapidly than does selection of one feature value from another (e.g., one color from another), as reported in previous electrophysiological studies. By combining methods with high spatial and temporal resolution it is possible to analyze the precise time course of feature-selective processing in specialized cortical areas.     

Dissociation of stimulus relevance and saliency factors during shifts of visuospatial attention

The control of visuospatial attention entails multiple processes, including both voluntary (endogenous) factors and stimulus-driven (exogenous) factors. Exogenous processes can be triggered by visual targets presented at a previously unattended location, thus capturing attention in a stimulus-driven manner. However, little is known about the relative role of stimulus salience and behavioral relevance for this type of spatial reorienting. Here, we directly assessed how salience and relevance affect activation of the frontoparietal attentional system, using either low-salience but task-relevant target stimuli or salient but task-irrelevant flickering checkerboards. We compared event-related functional magnetic resonance imaging responses for stimuli presented at the unattended versus attended side (invalid minus valid trials), separately for the 2 categories of visual stimuli. We found that task-relevant invalid targets activated the frontoparietal attentional network, demonstrating that this system engages when target stimuli are presented at an unattended location, even when these have a low perceptual salience. Conversely, the presentation of high-salience checkerboards in one hemifield while endogenous attention was engaged elsewhere did not activate the attentional network. These findings indicate that task relevance is critical for stimulus-driven engagement of the attentional network when attentional resources are endogenously allocated somewhere else.     

Monetary incentives enhance processing in brain regions mediating top-down control of attention

To evaluate the effect of an abstract motivational incentive on top-down mechanisms of visual spatial attention, 10 subjects engaged in a target detection task and responded to targets preceded by spatially valid (predictive), invalid (misleading) or neutral central cues under three different incentive conditions: win money (WIN), lose money (LOSE), and neutral (neither gain nor lose). Activation in the posterior cingulate cortex was correlated with visual spatial expectancy, defined as the degree to which the valid cue benefited performance as evidenced by faster reaction times compared to non-directional cues. Winning and losing money enhanced this relationship via overlapping but independent limbic mechanisms. In addition, activity in the inferior parietal lobule was correlated with disengagement (the degree to which invalid cues diminished performance). This relationship was also enhanced by monetary incentives. Finally, incentive enhanced the relationship of activation in the visual cortex to visual spatial expectancy and disengagement for both types of incentive (WIN and LOSE). These results show that abstract incentives enhance neural processing within the attention network in a process- and valence-selective manner. They also show that different cognitive and motivational mechanisms may produce a common effect upon unimodal cortices in order to enhance processing to serve the current behavioral goal.     

Modulation of connectivity in visual pathways by attention: cortical interactions evaluated with structural equation modelling and fMRI

Electrophysiological and neuroimaging studies have shown that attention to visual motion can increase the responsiveness of the motion- selective cortical area V5 and the posterior parietal cortex (PP). Increased or decreased activation in a cortical area is often attributed to attentional modulation of the cortical projections to that area. This leads to the notion that attention is associated with changes in connectivity. We have addressed attentional modulation of effective connectivity using functional magnetic resonance imaging (fMRI). Three subjects were scanned under identical stimulus conditions (visual motion) while varying only the attentional component of the task. Haemodynamic responses defined an occipito-parieto-frontal network, including the, primary visual cortex (V1), V5 and PR A structural equation model of the interactions among these dorsal visual pathway areas revealed increased connectivity between V5 and PP related to attention. On the basis of our analysis and the neuroanatomical pattern of projections from the prefrontal cortex to PP we attributed the source of modulatory influences, on the posterior visual pathway, to the prefrontal cortex (PFC). To test this hypothesis we included the PFC in our model as a 'modulator' of the pathway between V5 and PP, using interaction terms in the structural equation model. This analysis revealed a significant modulatory effect of prefrontal regions on V5 afferents to posterior parietal cortex.      

Retinotopy and attention in human occipital, temporal, parietal, and frontal cortex

Novel mapping stimuli composed of biological motion figures were used to study the extent and layout of multiple retinotopic regions in the entire human brain and to examine the independent manipulation of retinotopic responses by visual stimuli and by attention. A number of areas exhibited retinotopic activations, including full or partial visual field representations in occipital cortex, the precuneus, motion-sensitive temporal cortex (extending into the superior temporal sulcus), the intraparietal sulcus, and the vicinity of the frontal eye fields in frontal cortex. Early visual areas showed mainly stimulus-driven retinotopy; parietal and frontal areas were driven primarily by attention; and lateral temporal regions could be driven by both. We found clear spatial specificity of attentional modulation not just in early visual areas but also in classical attentional control areas in parietal and frontal cortex. Indeed, strong spatiotopic activity in these areas could be evoked by directed attention alone. Conversely, motion-sensitive temporal regions, while exhibiting attentional modulation, also responded significantly when attention was directed away from the retinotopic stimuli.     

Mechanisms of top-down facilitation in perception of visual objects studied by FMRI

Prior knowledge regarding the possible identity of an object facilitates its recognition from a degraded visual input, though the underlying mechanisms are unclear. Previous work implicated ventral visual cortex but did not disambiguate whether activity-changes in these regions are causal to or merely reflect an effect of facilitated recognition. We used functional magnetic resonance imaging to study top-down influences on processing of gradually revealed objects, by preceding each object with a name that was congruent or incongruent with the object. Congruently primed objects were recognized earlier than incongruently primed, and this was paralleled by shifts in activation profiles for ventral visual, parietal, and prefrontal cortices. Prior to recognition, defined on a trial-by-trial basis, activity in ventral visual cortex rose gradually but equivalently for congruently and incongruently primed objects. In contrast, prerecognition activity was greater with congruent priming in lateral parietal, retrosplenial, and lateral prefrontal cortices, whereas functional coupling between parietal and ventral visual (and also left lateral prefrontal and parietal) cortices was enhanced in the same context. Thus, when controlling for recognition point and stimulus information, activity in ventral visual cortex mirrors recognition success, independent of condition. Facilitation by top-down cues involves lateral parietal cortex interacting with ventral visual areas, potentially explaining why parietal lesions can lead to deficits in recognizing degraded objects even in the context of top-down knowledge.     

The effects of feature attention on prestimulus cortical activity in the human visual system

Covert attention affects prestimulus activity in the visual cortex. Although most studies investigating neural mechanisms of attention have focused on the effects of spatial attention, attention can also be directed to particular features. To investigate the spatiotemporal nature of feature attention, we measured subjects' brain activity using magnetoencephalography (MEG) and functional magnetic resonance imaging (fMRI) while subjects attended to color or motion of a stimulus based on a visual cue, which was presented 1 s before the stimulus onset. We used the hierarchical Bayesian method that allows us to estimate cortical currents with MEG and fMRI data in the order of millimeters and milliseconds. When subjects attended to color, activity within the color-sensitive area (fusiform gyrus) was selectively enhanced within the prestimulus period. By contrast, when subjects attended to motion, activity within the motion-sensitive area (middle temporal gyrus) was selectively enhanced during this period. This effect was not seen in frontal, parietal, and lower visual areas. Additionally, this effect was transient rather than sustained, suggesting that it differs from temporal aspects of spatial attention. These results suggest that, although both spatial and feature attention modulate prestimulus activity within specific visual areas, neural mechanisms underlying these effects might be different.     

Selective attention increases the dependency of cortical responses on visual motion coherence in man

Attention improves visual discrimination and consequently allows to discern stimuli with low signal-to-noise ratios that otherwise would remain undetected. We used magnetoencephalography (MEG) to test whether neuromagnetic responses recorded from occipito-temporal cortex, reflecting the size of visual motion signals embedded in noise (motion coherence), would mirror the perceptual changes induced by attention. Attention directed to a given hemifield increased and decreased the coherence modulation of the MEG response over contralateral and ipsilateral visual cortex, respectively, indicating a change in the neuronal signal-to-noise ratio at the population level.     

Natural vision reveals regional specialization to local motion and to contrast-invariant, global flow in the human brain

Visual changes in feature movies, like in real-live, can be partitioned into global flow due to self/camera motion, local/differential flow due to object motion, and residuals, for example, due to illumination changes. We correlated these measures with brain responses of human volunteers viewing movies in an fMRI scanner. Early visual areas responded only to residual changes, thus lacking responses to equally large motion-induced changes, consistent with predictive coding. Motion activated V5+ (MT+), V3A, medial posterior parietal cortex (mPPC) and, weakly, lateral occipital cortex (LOC). V5+ responded to local/differential motion and depended on visual contrast, whereas mPPC responded to global flow spanning the whole visual field and was contrast independent. mPPC thus codes for flow compatible with unbiased heading estimation in natural scenes and for the comparison of visual flow with nonretinal, multimodal motion cues in it or downstream. mPPC was functionally connected to anterior portions of V5+, whereas laterally neighboring putative homologue of lateral intraparietal area (LIP) connected with frontal eye fields. Our results demonstrate a progression of selectivity from local and contrast-dependent motion processing in V5+ toward global and contrast-independent motion processing in mPPC. The function, connectivity, and anatomical neighborhood of mPPC imply several parallels to monkey ventral intraparietal area (VIP).     

Sequence Seeking and Counter Streams: A Computational Model for Bidirectional Information Flow in the Visual Cortex

A computational model is proposed for some general aspects ofinformation flow in the visual cortex. The basic process, called"sequence seeking," is a search for a sequence of mappings,or transformations, linking source and target patterns. Theprocess has two main characteristics: it is bidirectional, bottom-upas well as top-down, and it explores in parallel a large numberof alternative sequences. This operation is performed in a "counterstreams" structure, in which multiple sequences are exploredalong two complementary pathways, an ascending and a descendingone, seeking to meet. A biological embodiment of this modelin cortical circuitry is proposed. The model serves to accountfor known aspects of cortical interconnections and to derivenew predictions.     

Contribution of working memory to transient activation in human inferior prefrontal cortex during performance of the Wisconsin Card Sorting Test.

The Wisconsin Card Sorting Test (WCST) is the standard task paradigm to detect human frontal lobe dysfunction. In this test, subjects sort card stimuli with respect to one of three possible dimensions (color, form and number). These dimensions are changed intermittently, whereupon subjects are required to identify by trial and error a new correct dimension and flexibly shift cognitive set. We decomposed the cognitive requirements at the time of the dimensional changes of the WCST, using functional magnetic resonance imaging (fMRI). By explicitly informing subjects of a new correct dimension, the working memory load for the trial-and-error identification of the new dimension was removed. Event-related fMRI still revealed transient activation time-locked to the dimensional changes in areas in the posterior part of the inferior frontal sulci. However, the activation was significantly smaller than in the original WCST in which subjects had to use working memory to identify the new dimension by trial and error. Furthermore, these areas were found to spatially overlap the areas activated by a working memory task. These results suggest that working memory and set-shifting act cooperatively in the same areas of prefrontal cortex to adapt us to changing environments.     

Lateralized auditory cortical alpha band activity and interregional connectivity pattern reflect anticipation of target sounds

The anticipation of stimuli facilitates the top-down preparation of neuronal tissue involved in the processing of forthcoming targets. Increasing evidence in the visual modality emphasizes the anticipatory adjustment of visual cortex excitability through modulations of oscillatory alpha power. In the auditory system, however, this relationship has not yet been established. Furthermore, the association between anticipatory modulations of auditory alpha power and a potential top-down network within these anticipatory preparation processes remains unexplained. To disclose these processes, we recorded magnetoencephalography while visually cuing participants to attend to either ear and to anticipate forthcoming auditory stimuli. For the cue-stimulus phase, we expected an asymmetric modulation of auditory alpha power when attending to the left or right ear, assuming that frontoparietal regions would phase synchronize with the auditory cortex in an asymmetric pattern. Beamformer source solutions demonstrate an asymmetric modulation of auditory alpha power following visual cues expressed in a strong right auditory alpha power increase when attending to the right ear. Furthermore, the right auditory cortex is functionally connected to the frontal eye fields during the ipsilateral alpha increase. Altogether, these results contribute significantly to the understanding of how auditory anticipation acts on a local as well as on a network level.