Multisensory enhancement elicited by unconscious visual stimuli.

The merging of information from different senses (i.e., multisensory integration) can facilitate information processing. Processing enhancements have been observed with signals that are irrelevant to the task at hand, and with cues that are non-predictive. Such findings are consistent with the notion that multiple sensory signals are sometimes integrated automatically. Multisensory enhancement has even been reported with stimuli that have been presented subliminally, though only with meaningful multisensory relations that have already been learned. The question of whether there exist cases where multisensory effects occur without either learning or awareness has, though, not been clearly established in the literature to date. Here, we present a case study of a patient with Posterior Cortical Atrophy, who was unable to consciously perceive visual stimuli with our task parameters, yet who nevertheless still exhibited signs of multisensory enhancement even with unlearned relations between audiovisual stimuli. In a simple speeded detection task, both response speed, and the variability of reaction times, decreased in a similar manner to controls for multisensory stimuli. These results are consistent with the view that the conscious perception of stimuli and prior learning are not always a prerequisite for multisensory integration to enhance human performance.

In-group biases and oculomotor responses: beyond simple approach motivation.

An in-group bias describes an individual's bias towards a group that they belong to. Previous studies suggest that in-group bias facilitates approach motor responses, but disrupts avoidance ones. Such motor biases are shown to be more robust when the out-group is threatening. We investigated whether, under controlled visual familiarity and complexity, in-group biases still promote pro-saccade and hinder anti-saccades oculomotor responses. Participants first learned to associate an in-group or out-group label with an arbitrary shape. They were then instructed to listen to the group-relevant auditory cue (name of own and a rival university) followed by one of the shapes. Half of the participants were instructed to look towards the visual target if it matched the preceding group-relevant auditory cue and to look away from it if it did not match. The other half of the participants received reversed instructions. This design allowed us to orthogonally manipulate the effect of in-group bias and cognitive control demand on oculomotor responses. Both pro- and anti-saccades were faster and more accurate following the in-group auditory cue. Independently, pro-saccades were performed better than anti-saccades, and match judgements were faster and more accurate than non-match judgements. Our findings indicate that under higher cognitive control demands individuals' oculomotor responses improved following the motivationally salient cue (in-group). Our findings have important implications for learning and cognitive control in a social context. As we included rival groups, our results might to some extent reflect the effects of out-group threat. Future studies could extend our findings using non-threatening out-groups instead.

Manipulating perceptual parameters in a continuous performance task.

Sustained attention (SA) is among the most studied faculties of human cognition, and thought to be crucial for many aspects of behavior. Measuring SA often relies on performance on a continuous, low-demanding task. Such continuous performance tasks (CPTs) have many variations, and sustained attention is typically estimated based on variability in reaction times. While relying on reaction times may be useful in some cases, it can pose a challenge when working with clinical populations. To increase interpersonal variability in task parameters that do not rely on speed, researchers have increased demands for memory and response inhibition. These approaches, however, may be confounded when used to assess populations that suffer from multiple cognitive deficits. In the current study, we propose a new approach for increasing task variability by increasing the attentional demands. In order to do so, we created a new variation of a CPT - a masked version, where inattention is more likely to cause misidentifying a target. After establishing that masking indeed decreases target detection, we further investigated which task parameter may influence response biases. To do so, we contrasted two versions of the CPT with different target/distractor ratio. We then established how perceptual parameters can be controlled independently in a CPT. Following the experimental manipulations, we tested the MCCPT with aging controls and chronic stroke patients to assure the task can be used with target populations. The results confirm the MCCPT as a task providing high sensitivity without relying on reaction speed, and feasible for patients.

The Neural Basis of Independence Versus Interdependence Orientations: A Voxel-Based Morphometric Analysis of Brain Volume.

Sociocultural research has established independence and interdependence as two fundamental ways of thinking about oneself and the social world. Recent neuroscience studies further demonstrate that these orientations modulate brain activity in various self- and socially related tasks. In the current study, we explored whether the traits of independence and interdependence are reflected in anatomical variations in brain structure. We carried out structural brain imaging on a large sample of healthy participants ( n = 265) who also completed self-report questionnaires of cultural orientations. Voxel-based morphometry analysis demonstrated that a relative focus of independence (vs. interdependence) was associated with increased gray-matter volume in a number of self-related regions, including ventromedial prefrontal cortex, right dorsolateral prefrontal cortex, and right rostrolateral prefrontal cortex. These results provide novel insights into the biological basis of sociocultural orientations.

Factors influencing planning of a familiar grasp to an object: what it is to pick a cup.

We assessed the factors influencing the planning of actions required to manipulate one of two everyday objects with matching dimensions but openings at opposite ends: a cup and a vase. We found that, for cups, measures of movement preparation to reach and grasp the object were influenced by whether the grasp was made to the functional part of the object (wide opening) and whether the action would end in a supinated as opposed to a pronated grasp. These factors interacted such that effects of hand posture were found only when a less familiar grasp was made to the non-functional part of the cup (the base). These effects were not found with the vase, which has a less familiar location for grasping. We interpret the results in terms of a parallel model of action selection, modulated by both the familiarity of the grasp to a part of the object, likely to reflect object 'affordances' and the end state comfort of the action.

Exploring the quiet eye in archery using field- and laboratory-based tasks.

The 'quiet eye' (QE)-a period of extended gaze fixation on a target-has been reported in many tasks that require accurate aiming. Longer quiet eye durations (QEDs) are reported in experts compared to non-experts and on successful versus less successful trials. The QE has been extensively studied in the field; however, the cognitive mechanisms underlying the QE are not yet fully understood. We investigated the QEDs of ten expert and ten novice archers in the field and in the laboratory using a computer-based archery task. The computer task consisted of shooting archery targets using a joystick. Random 'noise' (visual motion perturbation) was introduced at high and low levels to allow for the controlled examination of the effects of task complexity and processing demands. In this computer task, we also tested an additional group of ten non-archers as controls. In both field and computer tasks, eye movements were measured using electro-oculography. The expert archers exhibited longer QED compared to the novice archers in the field task. In the computer task, the archers again exhibited longer QEDs and were more accurate compared to non-archers. Furthermore, expert archers showed earlier QE onsets and longer QEDs during high noise conditions compared to the novices and non-archers. Our findings show skill-based effects on QED in field conditions and in a novel computer-based archery task, in which online (visual) perturbations modulated experts' QEDs. These longer QEDs in experts may be used for more efficient programming in which accurate predictions are facilitated by attention control.

Integration of identity and emotion information in faces: fMRI evidence.

Separate neural systems have been implicated in the recognition of facial identity and emotional expression. A growing number of studies now provide evidence against this modular view by demonstrating that integration of identity and emotion information enhances face processing. Yet, the neural mechanisms that shape this integration remain largely unknown. We hypothesize that the presence of both personal and emotional expression target information triggers changes in functional connectivity between frontal and extrastriate areas in the brain. We report and discuss three important findings. First, the presence of target identity and emotional expression in the same face was associated with super capacity and violations of the independent processing of identity and expression cues. Second, activity in the orbitofrontal cortex (OFC) was associated with the presence of redundant targets and changes in functional connectivity between a particular region of the right OFC (BA11/47) and bilateral visual brain regions (the inferior occipital gyrus (IOG)). Third, these changes in connectivity showed a strong link to behavioural measures of capacity processing. We suggest that the changes in functional connectivity between the right OFC and IOG reduce variability of BOLD responses in the IOG, enhancing integration of identity and emotional expression cues in faces.

Neural Mechanisms of Temporal Resolution of Attention.

The dynamic nature of the world requires that our visual representations are continuously updated. These representations are more precise if there is a narrow time window over which information is averaged. We assess the neural processes of visual updating by testing patients with lesions including inferior parietal cortex, control patients and healthy adults on a continuous visual monitoring task. In Experiment 1, observers kept track of the changing spatial period of a luminance grating and identified the final spatial period after the stimulus disappeared. Healthy older adults and neurological controls were able to perform better than simulated guesses, but only 3 of 11 patients with damage including parietal cortex were able to reach performance that differed from simulated guesses. The effects were unrelated to lesion size. Poor performance on this task is consistent with an inability to selectively attend to the final moment at which the stimulus was seen. To investigate the temporal limits of attention, we varied the rate of stimulus change in Experiment 2. Performance remained poor for some patients even with slow 2.5 Hz change rates. The performance of 4 patients with parietal damage displayed poor temporal precision, namely recovery of performance with slower rates of change.

Structural Organization of the Corpus Callosum Predicts Attentional Shifts after Continuous Theta Burst Stimulation.

Repetitive transcranial magnetic stimulation (rTMS) applied over the right posterior parietal cortex (PPC) in healthy participants has been shown to trigger a significant rightward shift in the spatial allocation of visual attention, temporarily mimicking spatial deficits observed in neglect. In contrast, rTMS applied over the left PPC triggers a weaker or null attentional shift. However, large interindividual differences in responses to rTMS have been reported. Studies measuring changes in brain activation suggest that the effects of rTMS may depend on both interhemispheric and intrahemispheric interactions between cortical loci controlling visual attention. Here, we investigated whether variability in the structural organization of human white matter pathways subserving visual attention, as assessed by diffusion magnetic resonance imaging and tractography, could explain interindividual differences in the effects of rTMS. Most participants showed a rightward shift in the allocation of spatial attention after rTMS over the right intraparietal sulcus (IPS), but the size of this effect varied largely across participants. Conversely, rTMS over the left IPS resulted in strikingly opposed individual responses, with some participants responding with rightward and some with leftward attentional shifts. We demonstrate that microstructural and macrostructural variability within the corpus callosum, consistent with differential effects on cross-hemispheric interactions, predicts both the extent and the direction of the response to rTMS. Together, our findings suggest that the corpus callosum may have a dual inhibitory and excitatory function in maintaining the interhemispheric dynamics that underlie the allocation of spatial attention. SIGNIFICANCE STATEMENT: The posterior parietal cortex (PPC) controls allocation of attention across left versus right visual fields. Damage to this area results in neglect, characterized by a lack of spatial awareness of the side of space contralateral to the brain injury. Transcranial magnetic stimulation over the PPC is used to study cognitive mechanisms of spatial attention and to examine the potential of this technique to treat neglect. However, large individual differences in behavioral responses to stimulation have been reported. We demonstrate that the variability in the structural organization of the corpus callosum accounts for these differences. Our findings suggest novel dual mechanism of the corpus callosum function in spatial attention and have broader implications for the use of stimulation in neglect rehabilitation.

Structural Variability within Frontoparietal Networks and Individual Differences in Attentional Functions: An Approach Using the Theory of Visual Attention.

Visuospatial attention allows us to select and act upon a subset of behaviorally relevant visual stimuli while ignoring distraction. Bundesen's theory of visual attention (TVA) (Bundesen, 1990) offers a quantitative analysis of the different facets of attention within a unitary model and provides a powerful analytic framework for understanding individual differences in attentional functions. Visuospatial attention is contingent upon large networks, distributed across both hemispheres, consisting of several cortical areas interconnected by long-association frontoparietal pathways, including three branches of the superior longitudinal fasciculus (SLF I-III) and the inferior fronto-occipital fasciculus (IFOF). Here we examine whether structural variability within human frontoparietal networks mediates differences in attention abilities as assessed by the TVA. Structural measures were based on spherical deconvolution and tractography-derived indices of tract volume and hindrance-modulated orientational anisotropy (HMOA). Individual differences in visual short-term memory (VSTM) were linked to variability in the microstructure (HMOA) of SLF II, SLF III, and IFOF within the right hemisphere. Moreover, VSTM and speed of information processing were linked to hemispheric lateralization within the IFOF. Differences in spatial bias were mediated by both variability in microstructure and volume of the right SLF II. Our data indicate that the microstructural and macrostrucutral organization of white matter pathways differentially contributes to both the anatomical lateralization of frontoparietal attentional networks and to individual differences in attentional functions. We conclude that individual differences in VSTM capacity, processing speed, and spatial bias, as assessed by TVA, link to variability in structural organization within frontoparietal pathways.

Temporal Binding and Segmentation in Visual Search: A Computational Neuroscience Analysis.

Human visual search operates not only over space but also over time, as old items remain in the visual field and new items appear. Preview search (where one set of distractors appears before the onset of a second set) has been used as a paradigm to study search over time and space [Watson, D. G., & Humphreys, G. W. Visual marking: Prioritizing selection for new objects by top-down attentional inhibition of old objects. Psychological Review, 104, 90-122, 1997], with participants showing efficient search when old distractors can be ignored and new targets prioritized. The benefits of preview search are lost, however, if a temporal gap is introduced between a first presentation of the old items and the re-presentation of all the items in the search display [Kunar, M. A., Humphreys, G. W., & Smith, K. J. History matters: The preview benefit in search is not onset capture. Psychological Science, 14, 181-185, 2003a], consistent with the old items being bound by temporal onset to the new stimuli. This effect of temporal binding can be eliminated if the old items reappear briefly before the new items, indicating also a role for the memory of the old items. Here we simulate these effects of temporal coding in search using the spiking search over time and space model [Mavritsaki, E., Heinke, D., Allen, H., Deco, G., & Humphreys, G. W. Bridging the gap between physiology and behavior: Evidence from the sSoTS model of human visual attention. Psychological Review, 118, 3-41, 2011]. We show that a form of temporal binding by new onsets has to be introduced to the model to simulate the effects of a temporal gap, but that effects of the memory of the old item can stem from continued neural suppression across a temporal gap. We also show that the model can capture the effects of brain lesion on preview search under the different temporal conditions. The study provides a proof-of-principle analysis that neural suppression and temporal binding can be sufficient to account for human search over time and space.

Unconscious Familiarity-based Color-Form Binding: Evidence from Visual Extinction.

There is good evidence that early visual processing involves the coding of different features in independent brain regions. A major question, then, is how we see the world in an integrated manner, in which the different features are "bound" together. A standard account of this has been that feature binding depends on attention to the stimulus, which enables only the relevant features to be linked together [Treisman, A., & Gelade, G. A feature-integration theory of attention. Cognitive Psychology, 12, 97-136, 1980]. Here we test this influential idea by examining whether, in patients showing visual extinction, the processing of otherwise unconscious (extinguished) stimuli is modulated by presenting objects in their correct (familiar) color. Correctly colored objects showed reduced extinction when they had a learned color, and this color matched across the ipsi- and contralesional items (red strawberry + red tomato). In contrast, there was no reduction in extinction under the same conditions when the stimuli were colored incorrectly (blue strawberry + blue tomato; Experiment 1). The result was not due to the speeded identification of a correctly colored ipsilesional item, as there was no benefit from having correctly colored objects in different colors (red strawberry + yellow lemon; Experiment 2). There was also no benefit to extinction from presenting the correct colors in the background of each item (Experiment 3). The data suggest that learned color-form binding can reduce extinction even when color is irrelevant for the task. The result is consistent with preattentive binding of color and shape for familiar stimuli.

Brain networks of temporal preparation: A multiple regression analysis of neuropsychological data.

There are only a few studies on the brain networks involved in the ability to prepare in time, and most of them followed a correlational rather than a neuropsychological approach. The present neuropsychological study performed multiple regression analysis to address the relationship between both grey and white matter (measured by magnetic resonance imaging in patients with brain lesion) and different effects in temporal preparation (Temporal orienting, Foreperiod and Sequential effects). Two versions of a temporal preparation task were administered to a group of 23 patients with acquired brain injury. In one task, the cue presented (a red versus green square) to inform participants about the time of appearance (early versus late) of a target stimulus was blocked, while in the other task the cue was manipulated on a trial-by-trial basis. The duration of the cue-target time intervals (400 versus 1400ms) was always manipulated within blocks in both tasks. Regression analysis were conducted between either the grey matter lesion size or the white matter tracts disconnection and the three temporal preparation effects separately. The main finding was that each temporal preparation effect was predicted by a different network of structures, depending on cue expectancy. Specifically, the Temporal orienting effect was related to both prefrontal and temporal brain areas. The Foreperiod effect was related to right and left prefrontal structures. Sequential effects were predicted by both parietal cortex and left subcortical structures. These findings show a clear dissociation of brain circuits involved in the different ways to prepare in time, showing for the first time the involvement of temporal areas in the Temporal orienting effect, as well as the parietal cortex in the Sequential effects.

Biased towards food: Electrophysiological evidence for biased attention to food stimuli.

We investigated the neural mechanisms involved in bias for food stimuli in our visual environment using event related lateralized (ERL) responses. The participants were presented with a cue (food or non-food item) to either identify or hold in working memory. Subsequently, they had to search for a target in a 2-item display where target and distractor stimuli were each flanked by a picture of a food or a non-food item. The behavioural data showed that performance was strongly affected by food cues, especially when food was held in WM compared to when the cues were merely identified. The temporal dynamics of electrophysiological measures of attention (the N1pc and N2pc) showed that the orienting of attention towards food stimuli was associated with two different mechanisms; an early stage of attentional suppression followed by a later stage of attentional orienting towards food stimuli. In contrast, non-food cues were associated only with the guidance of attention to or away from cued stimuli on valid and invalid trials. The results demonstrate that food items, perhaps due to their motivational significance modulate the early orienting of attention, including an initial suppressive response to food items.

Lesion-symptom mapping of self-prioritization in explicit face categorization: distinguishing hypo- and hyper-self-biases.

People make faster familiarity decisions for their own face compared with a familiar other. Lesion studies diverge on whether this self-face prioritization (SFP) effect is associated with functional processes isolated in the left or right hemispheres. To assess both decreases (hypo-) and increases (hyper-) in SFP after brain lesion, we asked patients with chronic deficits to perform familiarity judgments to images of their own face, a familiar other, or unfamiliar faces. Of 30 patients, 7 showed hypo- and 6 showed hyper-self-bias effects, comparing responses with their own faces versus responses with a familiar other. Hyper-self-bias correlated with reduced executive control function and, at a neural level, this was associated with lesions to the left prefrontal and superior temporal cortices. In contrast, reduced self-prioritization was associated with damage to the right inferior temporal structures including the hippocampus and extending to the fusiform gyrus. In addition, lesions affecting fibers crossing the right temporal cortex, potentially disconnecting occipital-temporal from frontal regions, diminished the self-bias effect. The data highlight that self-prioritized face processing is linked to regions in the right hemisphere associated with face recognition memory and it also calls on executive processes in the left hemisphere that normally modulate self-prioritized attention.

The salient self: the left intraparietal sulcus responds to social as well as perceptual-salience after self-association.

Perceptual learning is associated with experience-based changes in stimulus salience. Here, we use a novel procedure to show that learning a new association between a self-label and a neutral stimulus produces fast alterations in social salience measured by interference when targets associated with other people have to be selected in the presence of self-associated distractors. Participants associated neutral shapes with either themselves or a friend, over a short run of training trials. Subsequently, the shapes had to be identified in hierarchical (global-local) forms. The data show that giving a shape greater personal significance by associating it with the self had effects on visual selection equivalent to altering perceptual salience. Similar to previously observed effects linked to when perceptually salient distractors are ignored, effects of a self-associated distractor also increased activation in the left intraparietal cortex sulcus. The results show that self-associations to sensory stimuli rapidly modulate neural responses in a manner similar to changes in perceptual saliency. The self-association procedure provides a new way to understand how personal significance affects behavior.

Risk-sensitive decision-making in patients with posterior parietal and ventromedial prefrontal cortex injury.

Successful choice under risk requires the integration of information about outcome probabilities and values and implicates a brain network including the ventromedial prefrontal cortex (vmPFC) and posterior parietal cortex (pPAR). Damage to the vmPFC is linked to poor decision-making and increased risk-taking. Electrophysiological and neuroimaging data implicate the pPAR in the processing of reward probability during choice, but the causal contribution of this area has not been established. We compared patients with lesions to the pPAR (n = 13), vmPFC (n = 13), and healthy volunteers (n = 22) on the Roulette Betting Task, a measure of risk-sensitive decision-making. Both lesion groups were impaired in adjusting their bets to the probability of winning. This impairment was correlated with the extent of pPAR, but not vmPFC, damage. In addition, the vmPFC group chose higher bets than healthy controls overall, an effect that correlated with lesion volume in the medial orbitofrontal cortex. Both lesion groups earned fewer points than healthy controls. The groups did not differ on 2 tasks assessing probabilistic reasoning outside of a risk-reward context. Our results demonstrate the causal involvement of both the pPAR and vmPFC in risk-sensitive choice and indicate distinguishable roles of these areas in probability processing and risk appetite.

Coupling social attention to the self forms a network for personal significance.

Prior social psychological studies show that newly assigned personal significance can modulate high-level cognitive processes, e.g., memory and social evaluation, with self- and other-related information processed in dissociated prefrontal structure: ventral vs. dorsal, respectively. Here, we demonstrate the impact of personal significance on perception and show the neural network that supports this effect. We used an associative learning procedure in which we "tag" a neutral shape with a self-relevant label. Participants were instructed to associate three neutral shapes with labels for themselves, their best friend, or an unfamiliar other. Functional magnetic resonance imaging data were acquired while participants judged whether the shape-label pairs were maintained or swapped. Behaviorally, participants rapidly tagged a neutral stimulus with self-relevance, showing a robust advantage for self-tagged stimuli. Self-tagging responses were associated with enhanced activity in brain regions linked to self-representation [the ventral medial prefrontal cortex (vmPFC)] and to sensory-driven regions associated with social attention [the left posterior superior temporal sulcus (LpSTS)]. In contrast, associations formed with other people recruited a dorsal frontoparietal control network, with the two networks being inversely correlated. Responses in the vmPFC and LpSTS predicted behavioral self-bias effects. Effective connectivity analyses showed that the vmPFC and the LpSTS were functionally coupled, with the strength of coupling associated with behavioral self-biases. The data show that assignment of personal social significance affects perceptual matching by coupling internal self-representations to brain regions modulating attentional responses to external stimuli.

A Neural Decomposition of Visual Search Using Voxel-based Morphometry.

The ability to search efficiently for visual targets among distractors can break down after a variety of brain lesions, but the specific processes affected by the lesions are unclear. We examined search over space (conjunction search) and over time plus space (preview search) in a consecutive series of patients with acquired brain lesions. We also assessed performance on standard neuropsychological measures of visuospatial short-term memory (Corsi Block), sustained attention and memory updating (the contrast between forward and backward digit span), and visual neglect. Voxel-based morphometry analyses revealed regions in the occipital (middle occipital gyrus), posterior parietal (angular gyrus), and temporal cortices (superior and middle temporal gyri extending to the insula), along with underlying white matter pathways, associated with poor search. Going beyond standard voxel-based morphometry analyses, we then report correlation measures of structural damage in these regions and the independent neuropsychological measures of other cognitive functions. We find distinct patterns of correlation in areas linked to poor search, suggesting that the areas play functionally different roles in search. We conclude that neuropsychological disorders of search can be linked to necessary and distinct cognitive functions, according to the site of lesion.

Visual search in depth: The neural correlates of segmenting a display into relevant and irrelevant three-dimensional regions.

Visual perception is facilitated by the ability to selectively attend to relevant parts of the world and to ignore irrelevant regions or features. In visual search tasks, viewers are able to segment displays into relevant and irrelevant items based on a number of factors including the colour, motion, and temporal onset of the target and distractors. Understanding the process by which viewers prioritise relevant parts of a display can provide insights into the effect of top-down control on visual perception. Here, we investigate the behavioural and neural correlates of segmenting a display according to the expected three-dimensional (3D) location of a target. We ask whether this segmentation is based on low-level visual features (e.g. common depth or common surface) or on higher-order representations of 3D regions. Similar response-time benefits and neural activity were obtained when items fell on common surfaces or within depth-defined volumes, and when displays were vertical (such that items shared a common depth/disparity) or were tilted in depth. These similarities indicate that segmenting items according to their 3D location is based on attending to a 3D region, rather than a specific depth or surface. Segmenting the items in depth was mainly associated with increased activation in depth-sensitive parietal regions rather than in depth-sensitive visual regions. We conclude that segmenting items in depth is primarily achieved via higher-order, cue invariant representations rather than through filtering in lower-level perceptual regions.