Pupil size as a robust marker of attentional bias toward nicotine-related stimuli in smokers.

Spatial attention can be magnetically attracted by behaviorally salient stimuli. This phenomenon occasionally conflicts with behavioral goals, leading to maladaptive consequences, as in the case of addiction, in which attentional biases have been described and linked with clinically meaningful variables, such as craving level or dependence intensity. Here, we sought to probe the markers of attentional priority in smokers through eye-tracking measures, by leveraging the established link between eye movements and spatial attention. We were particularly interested in potential markers related to pupil size, because pupil diameter reflects a range of autonomic, affective, and cognitive/attentional reactions to behaviorally significant stimuli and is a robust marker of appetitive and aversive learning. We found that changes in pupil size to nicotine-related visual stimuli could reliably predict, in cross-validated logistic regression, the smoking status of young smokers (showing pupil constriction) better than more traditional proxy measures. The possibility that pupil constriction may reflect a bias toward central vision, for example, attentional capture, is discussed in terms of sensory tuning with respect to nicotine-related stimuli. Pupil size was more sensitive at lower nicotine dependence levels, and at increased abstinence time (though these two variables were collinear). We conclude that pupillometry can provide a robust marker for attentional priority computation and useful indications regarding motivational states and individual attitudes toward conditioned stimuli.

Effects of attentional shifts along the vertical axis on number processing: An eye-tracking study with optokinetic stimulation.

Previous studies suggest that associations between numbers and space are mediated by shifts of visuospatial attention along the horizontal axis. In this study, we investigated the effect of vertical shifts of overt attention, induced by optokinetic stimulation (OKS) and monitored through eye-tracking, in two tasks requiring explicit (number comparison) or implicit (parity judgment) processing of number magnitude. Participants were exposed to black-and-white stripes (OKS) that moved vertically (upward or downward) or remained static (control condition). During the OKS, participants were asked to verbally classify auditory one-digit numbers as larger/smaller than 5 (comparison task; Exp. 1) or as odd/even (parity task; Exp. 2). OKS modulated response times in both experiments. In Exp.1, upward attentional displacement decreased the Magnitude effect (slower responses for large numbers) and increased the Distance effect (slower responses for numbers close to the reference). In Exp.2, we observed a complex interaction between parity, magnitude, and OKS, indicating that downward attentional displacement slowed down responses for large odd numbers. Moreover, eye tracking analyses revealed an influence of number processing on eye movements both in Exp. 1, with eye gaze shifting downwards during the processing of small numbers as compared to large ones; and in Exp. 2, with leftward shifts after large even numbers (6,8) and rightward shifts after large odd numbers (7,9). These results provide evidence of bidirectional links between number and space and extend them to the vertical dimension. Moreover, they document the influence of visuo-spatial attention on processing of numerical magnitude, numerical distance, and parity. Together, our findings are in line with grounded and embodied accounts of numerical cognition.

Close facial emotions enhance physiological responses and facilitate perceptual discrimination.

Accumulating evidence indicates that the peripersonal space (PPS) constitutes a privileged area for efficient processing of proximal stimuli, allowing to flexibly adapt our behavior both to the physical and social environment. Whether and how behavioral and physiological signatures of PPS relate to each other in emotional contexts remains, though, elusive. Here, we addressed this question by having participants to discriminate male from female faces depicting different emotions (happiness, anger or neutral) and presented at different distances (50 cm-300 cm) while we measured the reaction time and accuracy of their responses, as well as pupillary diameter, heart rate and heart rate variability. Results showed facilitation of participants' performances (i.e., faster response time) when faces were presented close compared to far from the participants, even when controlling for retinal size across distances. These behavioral effects were accompanied by significant modulation of participants' physiological indexes when faces were presented in PPS. Interestingly, both PPS representation and physiological signals were affected by features of the seen faces such as the emotional valence, its sex and the participants' sex, revealing the profound impact of social context onto the autonomic state and behavior within PPS. Together, these findings suggest that both external and internal signals contribute in shaping PPS representation.

Atomoxetine modulates the contribution of low-level signals during free viewing of natural images in rhesus monkeys.

Visuo-spatial attentional orienting is fundamental to selectively process behaviorally relevant information, depending on both low-level visual attributes of stimuli in the environment and higher-level factors, such as goals, expectations and prior knowledge. Growing evidence suggests an impact of the locus-cœruleus-norepinephrine (LC-NE) system in attentional orienting that depends on taskcontext. Nonetheless, most of previous studies used visual displays encompassing a target and various distractors, often preceded by cues to orient the attentional focus. This emphasizes the contribution of goal-driven processes, at the expense of other factors related to the stimulus content. Here, we aimed to determine the impact of NE on attentional orienting in more naturalistic conditions, using complex images and without any explicit task manipulation. We tested the effects of atomoxetine (ATX) injections, a NE reuptake inhibitor, on four monkeys during free viewing of images belonging to three categories: landscapes, monkey faces and scrambled images. Analyses of the gaze exploration patterns revealed, first, that the monkeys spent more time on each fixation under ATX compared to the control condition, regard less of the image content. Second, we found that, depending on the image content, ATX modulated the impact of low-level visual salience on attentional orienting. This effect correlated with the effect of ATX on the number and duration of fixations. Taken together, our results demonstrate that ATX adjusts the contribution of salience on attentional orienting depending on the image content, indicative of its role in balancing the role of stimulus-driven and top-down control during free viewing of complex stimuli.

The role of the vestibular system in value attribution to positive and negative reinforcers.

Somatic inputs originating from bioregulatory processes can guide cognition and behavior. One such bodily signal, mostly overlooked so far, is represented by visuo-vestibular coupling and its alteration, which in extreme cases may result in motion sickness. We argued that the inherently perturbed interoceptive state that follows can be a powerful determinant of human motivated behavior, resulting in a blunted response to appetitive stimuli and an exaggerated response to noxious ones. We sought to assess such differential impact of visuo-vestibular mismatches on value through a task involving conflict monitoring. We therefore administered to 42 healthy participants a modified version of the Flankers task, in which distractors (arrows, pointing in either a congruent or incongruent direction) signaled the availability of monetary incentives (gains, losses, or neutral trials). While performing the task, participants received either galvanic vestibular stimulation (GVS), or sham stimulation. We have found impaired behavioral performances when value, which was attached to task-irrelevant information, was at stake. Gains and losses, interestingly, dissociated, and only the latter caused enhanced interference costs in the task, suggesting that negative incentives may be more effective in capturing human attention than positive ones. Finally, we have found some weak evidence for GVS to further increase the processing of losses, as suggested by even larger interference costs in this condition. Results were, however, overall ambiguous, and suggest that much more research is needed to better understand the link between the vestibular system and motivation.

Ipsilesional Impairments of Visual Awareness After Right-Hemispheric Stroke.

Unilateral brain damage following stroke frequently hampers the processing of contralesional space. Whether and how it also affects the processing of stimuli appearing on the same side of the lesion is still poorly understood. Three main alternative hypotheses have been proposed, namely that ipsilesional processing is functionally (i) hyperefficient, (ii) impaired, or (iii) spared. Here, we investigated ipsilesional space awareness through a computerized paradigm that exploits a manipulation of concurrent information processing demands (i.e., multitasking). Twelve chronic right-hemisphere stroke patients with a total lack of awareness for the contralesional side of space were administered a task that required the spatial monitoring of two locations within the ipsilesional hemispace. Targets were presented immediately to the right of a central fixation point (3° eccentricity), or farther to the right toward the screen edge (17° eccentricity), or on both locations. Response to target position occurred either in isolation or while performing a concurrent visual or auditory task. Results showed that most errors occurred when two targets were simultaneously presented and patients were faced with additional task demands (in the visual or auditory modalities). In the context of concurrent visual load, ipsilesional targets presented at the rightmost location were omitted more frequently than those presented closer to fixation. This pattern qualifies ipsilesional processing in right-hemisphere stroke patients as functionally impaired, arguing against the notion of ipsilesional hyperperformance, especially when under visual load.

Spatial grounding of symbolic arithmetic: an investigation with optokinetic stimulation.

Growing evidence suggests that mental calculation might involve movements of attention along a spatial representation of numerical magnitude. Addition and subtraction on nonsymbolic numbers (numerosities) seem to induce a "momentum" effect, and have been linked to distinct patterns of neural activity in cortical regions subserving attention and eye movements. We investigated whether mental arithmetic on symbolic numbers, a cornerstone of abstract mathematical reasoning, can be affected by the manipulation of overt spatial attention induced by optokinetic stimulation (OKS). Participants performed additions or subtractions of auditory two-digit numbers during horizontal (experiment 1) or vertical OKS (experiment 2), and eye movements were concurrently recorded. In both experiments, the results of addition problems were underestimated, whereas results of subtractions were overestimated (a pattern that is opposite to the classic Operational Momentum effect). While this tendency was unaffected by OKS, vertical OKS modulated the occurrence of decade errors during subtractions (i.e., fewer during downward OKS and more frequent during upward OKS). Eye movements, on top of the classic effect induced by OKS, were affected by the type of operation during the calculation phase, with subtraction consistently leading to a downward shift of gaze position and addition leading to an upward shift. These results highlight the pervasive nature of spatial processing in mental arithmetic. Furthermore, the preeminent effect of vertical OKS is in line with the hypothesis that the vertical dimension of space-number associations is grounded in universal (physical) constraints and, thereby, more robust than situated and culture-dependent associations with the horizontal dimension.

Mind the Depth: Visual Perception of Shapes Is Better in Peripersonal Space.

Closer objects are invariably perceived as bigger than farther ones and are therefore easier to detect and discriminate. This is so deeply grounded in our daily experience that no question has been raised as to whether the advantage for near objects depends on other features (e.g., depth itself). In a series of five experiments ( N = 114), we exploited immersive virtual environments and visual illusions (i.e., Ponzo) to probe humans' perceptual abilities in depth and, specifically, in the space closely surrounding our body, termed peripersonal space. We reversed the natural distance scaling of size in favor of the farther object, which thus appeared bigger, to demonstrate a persistent shape-discrimination advantage for close objects. Psychophysical modeling further suggested a sigmoidal trend for this benefit, mirroring that found for multisensory estimates of peripersonal space. We argue that depth is a fundamental, yet overlooked, dimension of human perception and that future studies in vision and perception should be depth aware.

Probing the role of the vestibular system in motivation and reward-based attention.

The vestibular system has widespread connections in the central nervous system. Several activation loci following vestibular stimulations have been notably reported in deep, limbic areas that are otherwise difficult to reach and modulate in healthy subjects. Following preliminary evidence, suggesting that such stimulations might affect mood and affective processing, we wondered whether the vestibular system is also involved in motivation. Evolutionary accounts suggest that visuo-vestibular mismatches might have a role in preventing the search for and exploitation of goods that previously resulted in aversive reactions, as they would be a fine warning signal which follows the contact with or ingestion of noxious neurotoxins. The first question was thus whether vestibular stimulation alters sensitivity to reward. Secondly, we sought to assess whether attention is allocated in space differently when cued by highly motivational stimuli, and if this interplay is further modulated by the vestibular system. In order to evaluate both motivational and attentional assets, we administered a Posner-like cueing task to 30 healthy subjects concurrently receiving sham or galvanic vestibular stimulation (GVS; Left-Anodal and Right-Anodal configurations). The participants had to discriminate targets appearing in either exogenously cued or uncued locations (50% validity); cues predicted the amount of points (0, 2, or 10) and thus money that they could earn for a correct response. The results highlight a robust inhibition of return (IOR) (faster responses for invalidly-cued targets) which was not modulated by different levels of reward or GVS. Across all stimulation sessions, rewards exerted a powerful beneficial effect over performance: reaction times were faster when rewards were at stake. However, this effect was largest in sham, but greatly reduced in GVS conditions, most notably with the Right-Anodal configuration. This is the first evidence for a decreased sensitivity to rewards causally induced by a perturbation of the vestibular system. While future studies will shed light on its neural underpinnings and clinical implications, here we argue that GVS could be a safe and promising way to enrich our understanding of reward processes and eventually tackle the management of patients with aberrant sensitivity to rewards.

Multi-tasking uncovers right spatial neglect and extinction in chronic left-hemisphere stroke patients.

Unilateral Spatial Neglect, the most dramatic manifestation of contralesional space unawareness, is a highly heterogeneous syndrome. The presence of neglect is related to core spatially lateralized deficits, but its severity is also modulated by several domain-general factors (such as alertness or sustained attention) and by task demands. We previously showed that a computer-based dual-task paradigm exploiting both lateralized and non-lateralized factors (i.e., attentional load/multitasking) better captures this complex scenario and exacerbates deficits for the contralesional space after right hemisphere damage. Here we asked whether multitasking would reveal contralesional spatial disorders in chronic left-hemisphere damaged (LHD) stroke patients, a population in which impaired spatial processing is thought to be uncommon. Ten consecutive LHD patients with no signs of right-sided neglect at standard neuropsychological testing performed a computerized spatial monitoring task with and without concurrent secondary tasks (i.e., multitasking). Severe contralesional (right) space unawareness emerged in most patients under attentional load in both the visual and auditory modalities. Multitasking affected the detection of contralesional stimuli both when presented concurrently with an ipsilesional one (i.e., extinction for bilateral targets) and when presented in isolation (i.e., left neglect for right-sided targets). No spatial bias emerged in a control group of healthy elderly participants, who performed at ceiling, as well as in a second control group composed of patients with Mild Cognitive Impairment. We conclude that the pathological spatial asymmetry in LHD patients cannot be attributed to a global reduction of cognitive resources but it is the consequence of unilateral brain damage. Clinical and theoretical implications of the load-dependent lack of awareness for contralesional hemispace following LHD are discussed.

Different effects of numerical magnitude on visual and proprioceptive reference frames.

This study assessed whether numerical magnitude affects the setting of basic spatial coordinates and reference frames, namely the subjective straight ahead. Three tasks were given to 24 right-handed healthy participants: a proprioceptive and a visuo-proprioceptive task, requiring pointing to the subjective straight ahead, and a visual task, requiring a perceptual judgment about the straight ahead position of a light moving left-to-right, or right-to-left. A control task, requiring the bisection of rods of different lengths, was also given. The four tasks were performed under conditions of passive auditory numerical (i.e., listening to small, "2," and large, "8," numbers), and neutral auditory-verbal ("blah") stimulation. Numerical magnitude modulated the participants' deviations in the visual straight ahead task, when the movement of the light was from left-to-right, with the small number bringing about a leftward deviation, the large number a rightward deviation. A similar directional modulation was found in the rod bisection task, in line with previous evidence. No effects of numerical magnitude were found on the proprioceptive and visuo-proprioceptive straight ahead tasks. These results suggest that the spatial effects induced by the activation of the mental number line extend to an egocentric frame of reference but only when a portion of horizontal space has to be "actively" explored.