Paying attention to working memory: Similarities in the spatial distribution of attention in mental and physical space.

Selective attention is not limited to information that is physically present in the external world, but can also operate on mental representations in the internal world. However, it is not known whether the mechanisms of attentional selection operate in similar fashions in physical and mental space. We studied the spatial distributions of attention for items in physical and mental space by comparing how successfully distractors were rejected at varying distances from the attended location. The results indicated very similar distribution characteristics of spatial attention in physical and mental space. Specifically, we found that performance monotonically improved with increasing distractor distance relative to the attended location, suggesting that distractor confusability is particularly pronounced for nearby distractors, relative to distractors farther away. The present findings suggest that mental representations preserve their spatial configuration in working memory, and that similar mechanistic principles underlie selective attention in physical and in mental space.

Distinct representations of attentional control during voluntary and stimulus-driven shifts across objects and locations.

Efficient interaction with the sensory environment requires the rapid reallocation of attentional resources between spatial locations, perceptual features, and objects. It is still a matter of debate whether one single domain-general network or multiple independent domain-specific networks mediate control during shifts of attention across features, locations, and objects. Here, we employed functional magnetic resonance imaging to directly compare the neural mechanisms controlling attention during voluntary and stimulus-driven shifts across objects and locations. Subjects either maintained or switched voluntarily and involuntarily their attention to objects located at the same or at a different visual location. Our data demonstrate shift-related activity in multiple frontoparietal, extrastriate visual, and default-mode network regions, several of which were commonly recruited by voluntary and stimulus-driven shifts between objects and locations. However, our results also revealed object- and location-selective activations, which, moreover, differed substantially between voluntary and stimulus-driven attention. These results suggest that voluntary and stimulus-driven shifts between objects and locations recruit partially overlapping, but also separable, cortical regions, implicating the parallel existence of domain-independent and domain-specific reconfiguration signals that initiate attention shifts in dependence of particular demands.

Spatiotemporal dynamics of feature-based attention spread: evidence from combined electroencephalographic and magnetoencephalographic recordings.

Attentional selection on the basis of nonspatial stimulus features induces a sensory gain enhancement by increasing the firing-rate of individual neurons tuned to the attended feature, while responses of neurons tuned to opposite feature-values are suppressed. Here we recorded event-related potentials (ERPs) and magnetic fields (ERMFs) in human observers to investigate the underlying neural correlates of feature-based attention at the population level. During the task subjects attended to a moving transparent surface presented in the left visual field, while task-irrelevant probe stimuli executing brief movements into varying directions were presented in the opposite visual field. ERP and ERMF amplitudes elicited by the unattended task-irrelevant probes were modulated as a function of the similarity between their movement direction and the task-relevant movement direction in the attended visual field. These activity modulations reflecting globally enhanced processing of the attended feature were observed to start not before 200 ms poststimulus and were localized to the motion-sensitive area hMT. The current results indicate that feature-based attention operates in a global manner but needs time to spread and provide strong support for the feature-similarity gain model.

Feature-based attention modulates direction-selective hemodynamic activity within human MT.

Attending to the spatial location or to nonspatial features of a stimulus modulates neural activity in cortical areas that process its perceptual attributes. The feature-based attentional selection of the direction of a moving stimulus is associated with increased firing of individual neurons tuned to the direction of the movement in area V5/MT, while responses of neurons tuned to opposite directions are suppressed. However, it is not known how these multiplicatively scaled responses of individual neurons tuned to different motion-directions are integrated at the population level, in order to facilitate the processing of stimuli that match the perceptual goals. Using functional magnetic resonance imaging (fMRI) the present study revealed that attending to the movement direction of a dot field enhances the response in a number of areas including the human MT region (hMT) as a function of the coherence of the stimulus. Attending the opposite direction, however, lead to a suppressed response in hMT that was inversely correlated with stimulus-coherence. These findings demonstrate that the multiplicative scaling of single-neuron responses by feature-based attention results in an enhanced direction-selective population response within those cortical modules that processes the physical attributes of the attended stimuli. Our results provide strong support for the validity of the "feature similarity gain model" on the integrated population response as quantified by parametric fMRI in humans.

Neural processing of reward magnitude under varying attentional demands.

Central to the organization of behavior is the ability to represent the magnitude of a prospective reward and the costs related to obtaining it. Therein, reward-related neural activations are discounted in dependence of the effort required to resolve a given task. Varying attentional demands of the task might however affect reward-related neural activations. Here we employed fMRI to investigate the neural representation of expected values during a monetary incentive delay task with varying attentional demands. Following a cue, indicating at the same time the difficulty (hard/easy) and the reward magnitude (high/low) of the upcoming trial, subjects performed an attention task and subsequently received feedback about their monetary reward. Consistent with previous results, activity in anterior-cingulate, insular/orbitofrontal and mesolimbic regions co-varied with the anticipated reward-magnitude, but also with the attentional requirements of the task. These activations occurred contingent on action-execution and resembled the response time pattern of the subjects. In contrast, cue-related activations, signaling the forthcoming task-requirements, were only observed within attentional control structures. These results suggest that anticipated reward-magnitude and task-related attentional demands are concurrently processed in partially overlapping neural networks of anterior-cingulate, insular/orbitofrontal, and mesolimbic regions.

Substantia nigra activity level predicts trial-to-trial adjustments in cognitive control.

Effective adaptation to the demands of a changing environment requires flexible cognitive control. The medial and the lateral frontal cortices are involved in such control processes, putatively in close interplay with the BG. In particular, dopaminergic projections from the midbrain (i.e., from the substantia nigra [SN] and the ventral tegmental area) have been proposed to play a pivotal role in modulating the activity in these areas for cognitive control purposes. In that dopaminergic involvement has been strongly implicated in reinforcement learning, these ideas suggest functional links between reinforcement learning, where the outcome of actions shapes behavior over time, and cognitive control in a more general context, where no direct reward is involved. Here, we provide evidence from functional MRI in humans that activity in the SN predicts systematic subsequent trial-to-trial RT prolongations that are thought to reflect cognitive control in a stop-signal paradigm. In particular, variations in the activity level of the SN in one trial predicted the degree of RT prolongation on the subsequent trial, consistent with a modulating output signal from the SN being involved in enhancing cognitive control. This link between SN activity and subsequent behavioral adjustments lends support to theoretical accounts that propose dopaminergic control signals that shape behavior both in the presence and in the absence of direct reward. This SN-based modulatory mechanism is presumably mediated via a wider network that determines response speed in this task, including frontal and parietal control regions, along with the BG and the associated subthalamic nucleus.

Sound-induced enhancement of low-intensity vision: multisensory influences on human sensory-specific cortices and thalamic bodies relate to perceptual enhancement of visual detection sensitivity.

Combining information across modalities can affect sensory performance. We studied how co-occurring sounds modulate behavioral visual detection sensitivity (d'), and neural responses, for visual stimuli of higher or lower intensity. Co-occurrence of a sound enhanced human detection sensitivity for lower- but not higher-intensity visual targets. Functional magnetic resonance imaging (fMRI) linked this to boosts in activity-levels for sensory-specific visual and auditory cortex, plus multisensory superior temporal sulcus (STS), specifically for a lower-intensity visual event when paired with a sound. Thalamic structures in visual and auditory pathways, the lateral and medial geniculate bodies, respectively (LGB, MGB), showed a similar pattern. Subject-by-subject psychophysical benefits correlated with corresponding fMRI signals in visual, auditory, and multisensory regions. We also analyzed differential "coupling" patterns of LGB and MGB with other regions in the different experimental conditions. Effective-connectivity analyses showed enhanced coupling of sensory-specific thalamic bodies with the affected cortical sites during enhanced detection of lower-intensity visual events paired with sounds. Coupling strength between visual and auditory thalamus with cortical regions, including STS, covaried parametrically with the psychophysical benefit for this specific multisensory context. Our results indicate that multisensory enhancement of detection sensitivity for low-contrast visual stimuli by co-occurring sounds reflects a brain network involving not only established multisensory STS and sensory-specific cortex but also visual and auditory thalamus.

Neural correlates of exemplar novelty processing under different spatial attention conditions.

The detection of novel events and their identification is a basic prerequisite in a rapidly changing environment. Recently, the processing of novelty has been shown to rely on the hippocampus and to be associated with activity in reward-related areas. The present study investigated the influence of spatial attention on neural processing of novel relative to frequently presented standard and target stimuli. Never-before-seen Mandelbrot-fractals absent of semantic content were employed as stimulus material. Consistent with current theories, novelty activated a widespread network of brain areas including the hippocampus. No activity, however, could be observed in reward-related areas with the novel stimuli absent of a semantic meaning employed here. In the perceptual part of the novelty-processing network a region in the lingual gyrus was found to specifically process novel events when they occurred outside the focus of spatial attention. These findings indicate that the initial detection of unexpected novel events generally occurs in specialized perceptual areas within the ventral visual stream, whereas activation of reward-related areas appears to be restricted to events that do possess a semantic content indicative of the biological relevance of the stimulus.

On perceived synchrony-neural dynamics of audiovisual illusions and suppressions.

Whenever temporally incongruent audiovisual sequences are presented, the perceived flash rate follows the physical flutter rate. Increasing the auditory flutter rate increases the perceived flicker rate (visual illusions). Likewise, decreasing the flutter rate decreases the perceived flicker rate (visual suppressions). Here, we investigated the electrophysiological correlates of this perceptual phenomenon. Two sequences of visual flashes and auditory beeps were presented either synchronously (both visual flashes (F) and auditory beeps (B) at 3 or 5 Hz, respectively) or asynchronously at different rates (3F5B or 5F3B). Event-related potentials were acquired, while subjects reported the perceived number of flashes (response options: 3, 4, and 5). During asynchronous trials, subjects' flash counts were significantly higher when the flutter rate exceeded the flicker rate (i.e. visual illusions occurred); and lower flutter rate was below the flicker rate (i.e. visual suppressions occurred). Differential brain responses for reported illusions and suppressions (incorrect flash counts) vs. no-illusions/suppressions (correct flash counts) were found over parieto-occipital sites, followed by slow modulations over frontal and occipital areas. Importantly, the modulation over occipital electrodes starting around 500 ms had an inverse polarity for illusions vs. suppressions. These results provide evidence that both sound-induced visual illusions and suppressions are mediated by an interplay of distributed brain regions, in the attempt to fuse asynchronous audiovisual stimuli into a synchronous percept.

Neural mechanisms of spatial- and feature-based attention: a quantitative analysis.

Attentional selection can be based on spatial locations, non-spatial stimulus features, or entire objects as integrated feature ensembles. Several studies reported attentional modulations in those regions that process the constituent features of the presented stimuli. Here we employed functional magnetic resonance imaging (fMRI) to directly compare the magnitude of space- and/or feature-based attentional modulations while subjects directed their attention to a particular color (red or green) of a transparent surface and at the same time to a spatial location (left or right visual field). The experimental design made it possible to disentangle and quantify the hemodynamic activity elicited by identical physical stimuli when attention was directed to spatial locations and/or stimulus features. The highest modulations were observed when the attentional selection was based on spatial location. Attended features also elicited a response increase relative to unattended features when their spatial location was attended. Importantly, at unattended locations, a response increase upon feature-based selection was observed in motion-sensitive but not in color-related areas. This suggests that compared to color, motion stimuli are more effective in capturing attention at unattended locations leading to a competitive advantage. These results support the idea of a high biological relevance of the feature motion in the visual world.