Exogenous Attention Research Articles

Effect of attention on ensemble perception: Comparison between exogenous attention, endogenous attention, and depth.

Ensemble perception is an important ability of human beings that allows one to extract summary information for scenes and environments that contain information that far exceeds the processing limit of the visual system. Although attention has been shown to bias ensemble perception, two important questions remain unclear: (1) whether direct manipulations on different types of spatial attention could produce similar effects on ensembles and (2) whether factors potentially influencing the attention distribution, such as depth perception, could evoke an indirect effect of attention on ensemble representation. This study aims to address these questions. In Experiments 1 and 2, two types of precues were used to evoke exogenous and endogenous attention, respectively, and the ensemble color perceptions were examined. We found that both exogenous and endogenous attention biased ensemble representation towards the attended items, and the latter produced a greater effect. In Experiments 3 and 4, we examined whether depth perception could affect color ensembles by indirectly influencing attention allocation in 3D space. The items were separated in two depth planes, and no explicit cues were applied. The results showed that color ensemble was biased to closer items when depth information was task relevant. This suggests that ensemble perception is naturally biased in 3D space, probably through the mechanism of attention. Computational modeling consistently showed that attention exerted a direct shift on the ensemble statistics rather than averaging the feature values over the cued and noncued items, providing evidence against an averaging process of individual perception.

Constraining the Possible Mechanisms Underlying Performance Improvements Following Peripheral Vision Training

AbstractSome training paradigms can improve performance specific to the trained portion of the visual field. Similarly, experience with both pathological and simulated central vision loss can result in compensatory improvement in performance specific to a spared retinal location. The mechanisms underlying these improvements are still debated. Modifications in the capacity to allocate attention to trained locations might explain some of the changes in performance after learning, especially given similarities between behavioral improvements due to training and improvements due to changes in attention. Using a gaze-contingent simulated scotoma paradigm which simulates central vision loss, we tested whether training to use peripheral vision influenced three aspects of visual attention: habitual attention, dynamic exogenous attention, and dynamic endogenous attention. After training, performance improvements were consistent with increased habitual attention to the trained location. Conversely, measures of dynamic shifts of attention (exogenous and endogenous attention) improved similarly in both trained and untrained locations. The lack of retinotopic specificity in dynamic attention improvements suggests that retinotopically specific perceptual improvements following simulated central vision loss are not mediated by dynamic attention changes. On the other hand, habitual attention did change retinotopically, leaving the possibility open that this aspect of attention may drive some retinotopically specific training effects. This work constrains the range of mechanisms that could underlie performance improvements after peripheral vision training, suggesting that habitual attention and dynamic attention are affected independently.

Adaptation and exogenous attention interact in the early visual cortex: A TMS study

Adaptation and exogenous attention interact in the early visual cortex: A TMS study

Dissociable Effects of Endogenous and Exogenous Attention on Crowding: Evidence from Event-Related Potentials.

Crowding is a common visual phenomenon that can significantly impair the recognition of objects in peripheral vision. Two recent behavioral studies have revealed that both exogenous and endogenous attention can alleviate crowding, but exogenous attention seems to be more effective. The present study employed the event-related potential (ERP) technique to explore the electrophysiological characteristics of the influence of these two types of attention on crowding. In the experiment, participants were required to judge whether the letter "T" was upright or inverted, which may be preceded by an exogenous cue or an endogenous cue indicating the location of the target letter. The behavioral results showed that while exogenous cues reduced crowding in all stimulus onset asynchronies (SOAs), endogenous attention took effects only in long SOA. The ERP results indicated that both endogenous and exogenous cues significantly alleviated the inhibition of visual crowding on the N1 component. However, the endogenous cue was effective only under long SOA, while the exogenous cue was effective only under short SOA conditions. In addition, invalid exogenous cues induced a larger P3 wave amplitude than valid ones in the short SOA condition, but endogenous attention did not show such a difference. These results indicate that both endogenous and exogenous attention can alleviate the effects of visual crowding, but they differ in effect size and temporal dynamics.

Exogenous spatial attention selects associated novel bindings in working memory

Real-world environments are complex, demanding a diverse set of cognitive functions such as attention and working memory (WM) to perform adaptive behaviors. However, exogenous attention, characterized as automatic and involuntary, has primarily been studied by focusing on spatial perception. In particular, the ability of pure exogenous retro-cues to select and prioritize not only spatial locations, but also novel stimulus–response (S-R) bindings held in WM remains largely unexplored. Here, in two experimental series, we provide evidence that pure exogenous non-predictive retro-cues can select not only space, but also associated S-R bindings held in WM. Additional evidence from a drift–diffusion model hinted at the possibility that the mechanisms through which exogenous attention selects and prioritizes WM contents depend, at least partially, on the hierarchical relevance of the different dimensions encoded within a specific representation. These results highlight the relationship between pure exogenous attention and complex WM contents and shed light on current theoretical debates about the interaction of attention, memory, and action.

How preferences enslave attention: calling into question the endogenous/exogenous dichotomy from an active inference perspective

AbstractIt is easy to think of attention as a purely sensorimotor, exogenous mechanism divorced from the influence of an agent’s preferences and needs. However, according to the active inference framework, such a strict reduction cannot be straightforwardly invoked, since all cognitive and behavioural processes can at least be described as maximising the evidence for a generative model entailed by the ongoing existence of that agent; that is, the minimisation of variational free energy. As such, active inference models could cast an (embodied) cognitive mechanism like attention, described in this paper as a relevance filter, as constrained (or enslaved) by these prior preferences for which an agent must seek evidence, whether or not such priors are having direct, real-time neurocognitive effects on the sensorimotor loops that couple the attending agent and her surrounding environment. This duality with respect to the role of priors corresponds to a wider, ongoing debate in the active inference community regarding the framework’s explanatory power. More specifically, the debate centres on whether the notion of a generative model and the priors embedded ubiqitously therein act as a purely useful instrumental tool for scientists aiming to model the behaviours of self-organising entities, or, rather, the brain (and body) is genuinely constituted by a predictive hierarchy within which higher-order dynamics constrain and contextualise activity unfolding at lower levels. With a focus on the second (ontologically realist) construal of active inference presented here, this paper argues that in cognitive systems endowed with attentional schema, higher-order preferences do, indeed, impose a demonstrable and powerful modulating effect on the way attention unfolds. Furthermore, these preferences in question transcend the contingent, task-relevant goals that have already been shown to bias attention. Rather, attention is powerfully tuned by the most-deep rooted priors the agent possesses, such that, when sensory evidence against these priors is observed and free energy spikes, the agent attentionally prioritises the homeostatic restoration of these preferred states over their shorter-term desires. This suggests that, at its core, attention is a goal-driven process, which calls into question the putative dichotomy that exists between endogenous (goal-directed) attention and exogenous (stimulus-driven) attention. What emerges in its place is a symbiotic relationship between attention and preferences, whereby the fulfilment of the latter rests on successful application of the former, and the former derives its function from the organismic need to find evidence for the latter.

Fine-tuning exogenous attention at the foveal scale: effects on different spatial frequencies

Fine-tuning exogenous attention at the foveal scale: effects on different spatial frequencies

Threat-induced prosocial behavior: enhanced exogenous attention to protect others from harm

As social animals, humans tend to voluntarily engage in pro-social behavior to prevent harm to others. However, to what extent prosocial behavior can be reflected at the level of less voluntary cognitive processes remains unclear. Here, we examined how threat to others modulates exogenous attention. Fifty-four participants performed an exogenous spatial cueing task where the participant’s performance determined whether electric shocks would be delivered either to themselves or to their anonymous co-participant. Threat of shock to the co-participant elicited orienting and reorienting responses that were faster than in the safe condition and did not differ from performance when participants avoided shocks to themselves. This attentional improvement was not due to speed-accuracy trade off and was associated with arousal, i.e., increased pupil dilation in both threat conditions. Together, these findings suggest that pro-social behavior triggers automatic attentional processes which may be relevant for providing immediate help without relying on reflexive processes.

Endogenous attention enhances contrast appearance regardless of stimulus contrast.

There has been enduring debate on how attention alters contrast appearance. Recent research indicates that exogenous attention enhances contrast appearance for low-contrast stimuli but attenuates it for high-contrast stimuli. Similarly, one study has demonstrated that endogenous attention heightens perceived contrast for low-contrast stimuli, yet none have explored its impact on high-contrast stimuli. In this study, we investigated how endogenous attention alters contrast appearance, with a specific focus on high-contrast stimuli. In Experiment 1, we utilized the rapid serial visual presentation (RSVP) paradigm to direct endogenous attention, revealing that contrast appearance was enhanced for both low- and high-contrast stimuli. To eliminate potential influences from the confined attention field in the RSVP paradigm, Experiment 2 adopted the letter identification paradigm, deploying attention across a broader visual field. Results consistently indicated that endogenous attention increased perceived contrast for high-contrast stimuli. Experiment 3 employed equiluminant chromatic letters as stimuli in the letter identification task to eliminate potential interference from contrast adaption, which might have occurred in Experiment 2. Remarkably, the boosting effect of endogenous attention persisted. Combining the results from these experiments, we propose that endogenous attention consistently enhances contrast appearance, irrespective of stimulus contrast levels. This stands in contrast to the effects of exogenous attention, suggesting that mechanisms through which endogenous attention alters contrast appearance may differ from those of exogenous attention.

The vestigial pinna-orienting system in humans briefly suppresses superior auricular muscle activity during reflexive orienting toward auditory stimuli.

The vestigial pinna-orienting system in humans is capable of increasing the activity of several auricular muscles in response to lateralized transient auditory stimuli. For example, transient increases in electromyographic activity in the posterior auricular muscle (PAM) to an attention-capturing stimulus have been documented. For the current study, surface electromyograms (EMGs) were recorded from the PAMs and superior auricular muscles (SAMs) of 10 normal-hearing participants. During the experiments, lateralized transient auditory stimuli, such as a crying baby, a shattering vase, or the participant's first names, were presented. These transient stimuli were either presented in silence or when participants actively listened to a podcast. Although ipsilateral PAM activity increased in response to transient stimuli, the SAM displayed the opposite behavior, i.e., a brief, ipsilateral suppression of activity. This suppression of ipsilateral SAM activity was more frequent on the right (75%) than left side (35%), whereas an ipsilateral PAM increase was roughly equal in prevalence on the two sides (left: 90%, right: 95%). During the active listening task, SAM suppression on the right ear was significantly larger in response to ipsilateral stimuli, compared with contralateral ones (P = 0.002), whereas PAM activity increased significantly (P = 0.002). Overall, this study provides evidence of a systematic transient suppression of the SAM during exogenous attention. This could suggest a more complex system than previously assumed, as the presence of synchronized excitatory and inhibitory components in different auricular muscles points toward a coordinated attempt at reflexively orienting the pinna toward a sound.NEW & NOTEWORTHY This study provides evidence that two auricular muscles in humans, the posterior and superior auricular muscles (PAM, SAM), react fundamentally different to lateralized transient auditory stimuli, especially during active listening. Although the PAM reacts with a transient increase in ipsilateral activity, ongoing ipsilateral SAM activity is briefly suppressed at the same time. This indicates the presence of a more complex and nuanced pinna-orienting system, with synchronized excitatory and inhibitory components in humans, than previously suspected.

The disengagement deficit after right-hemisphere damage: Distinct roles of lateral frontal and parietal damage

An influential model of spatial attention postulates three main attention-orienting mechanisms: disengagement, shifting, and engagement. Early research linked disengagement deficits with superior parietal damage, regardless of hemisphere or presence of spatial neglect. Subsequent studies supported the involvement of more ventral parietal regions, especially in the right hemisphere, and linked spatial neglect to deficient disengagement from ipsilateral cues. However, previous lesion studies faced serious limitations, such as small sample sizes and the lack of brain-injured controls without neglect. Additionally, some studies employed symbolic cues or used long cue-target intervals, which may fail to reveal impaired disengagement. We here used a machine-learning approach to conduct lesion-symptom mapping (LSM) on 89 patients with focal cerebral lesions to the left (LH) or right (RH) cerebral hemisphere. A group of 54 healthy participants served as controls. The paradigm used to uncover disengagement deficits employed non-predictive cues presented in the visual periphery and at short cue-target intervals, targeting exogenous attention. The main factors of interest were group (healthy participants, LH, RH), target position (left, right hemifield) and cue validity (valid, invalid). LSM-analyses were performed on two indices: the validity effect, computed as the absolute difference between reaction times (RTs) following invalid compared to valid cues, and the disengagement deficit, determined by the difference between contralesional and ipsilesional validity effects. While LH patients showed general slowing of RTs to contralesional targets, only RH patients exhibited a disengagement deficit from ipsilesional cues. LSM associated the validity effect with a right lateral frontal cluster, which additionally affected subcortical white matter of the right arcuate fasciculus, the corticothalamic pathway, and the superior longitudinal fasciculus. In contrast, the disengagement deficit was related to damage involving the right temporoparietal junction. Thus, our results support the crucial role of right inferior parietal and posterior temporal regions for attentional disengagement, but also emphasize the importance of lateral frontal regions, for the reorienting of attention.

Learning to discriminate the eye-of-origin during continuous flash suppression.

Helmholtz asked whether one could discriminate which eye is the origin of one's perception merely based on the retinal signals. Studies to date showed that participants' ability to tell the eye-of-origin most likely depends on contextual cues. Nevertheless, it has been shown that exogenous attention can enhance performance for monocularly presented stimuli. We questioned whether adults can be trained to discriminate the eye-of-origin of their perceptions and if this ability depends on the strength of the monocular channels. We used attentional feed-forward training to improve the subject's eye-of-origin discrimination performance with voluntary attention. During training, participants received a binocular cue to inform them of the eye-of-origin of an upcoming target. Using continuous flash suppression, we also measured the signal strength of the monocular targets to see any possible modulations related to the cues. We collected confidence ratings from the participants about their eye-of-origin judgements to study in further detail whether metacognition has access to this information. Our results show that, even though voluntary attention did not alter the strength of the monocular channels, eye-of-origin discrimination performance improved following the training. A similar pattern was observed for confidence. The results from the feedforward attentional training and the increase in subjective confidence point towards a high-level decisional mechanism being responsible for the eye-of-origin judgements. We propose that this high-level process is informed by subtle sensory cues such as the differences in luminance or contrast in the two monocular channels.

Attentional development is altered in toddlers with congenital heart disease.

Congenital Heart Disease (CHD) is the most common congenital abnormality. Survival rates are over 90%, however infants with CHD remain at high risk of attention and executive function impairments. These abilities are difficult to assess in toddlers because clinical assessments rely on language abilities which are commonly delayed in CHD. Our aim was to characterise visual attention in toddlers with CHD compared to controls and identify associations with parent-rated effortful control. Thirty toddlers with CHD (19 male, median (IQR) age at assessment 22.2 (22-23.1) months) and 66 controls from the developing human connectome project (36 male, age at assessment 22 (21.5-23.8) months) using eye-tracking tasks designed to assess multiple components of visual attention. Analyses of co-variance and regressions were used to identify differences between groups and relationships between gaze behaviours and parent-rated effortful control. Toddlers with CHD were less accurate when switching behaviours (set-shifting) [median (IQR) 79%, (28-100)] compared to controls [100% (86-100), pFDR=0.032], with worse accuracy associated with lower parent-rated effortful control in CHD but not controls (interaction pFDR=0.028). Reaction times were slower during selective [CHD 1243ms (986-1786), controls 1065ms (0851-1397), pFDR<0.001] and exogenous attention tasks [CHD 312ms (279-358), control 289 (249-331), (pFDR=0.032) and endogenous attention was less mature (prolonged looks at facial stimuli CHD 670ms (518-885), control 500ms (250-625), (pFDR=0.006). These results were unrelated to differences in cognition or socioeconomic status. In contrast, the allocation of attentional resources was preserved in CHD. We identified a profile of altered attention and early executive functioning development in CHD. Eye-tracking may provide clinically feasible, early objective measures of attention and executive function development in CHD.

Intracortical recordings reveal vision-to-action cortical gradients driving human exogenous attention

Exogenous attention, the process that makes external salient stimuli pop-out of a visual scene, is essential for survival. How attention-capturing events modulate human brain processing remains unclear. Here we show how the psychological construct of exogenous attention gradually emerges over large-scale gradients in the human cortex, by analyzing activity from 1,403 intracortical contacts implanted in 28 individuals, while they performed an exogenous attention task. The timing, location and task-relevance of attentional events defined a spatiotemporal gradient of three neural clusters, which mapped onto cortical gradients and presented a hierarchy of timescales. Visual attributes modulated neural activity at one end of the gradient, while at the other end it reflected the upcoming response timing, with attentional effects occurring at the intersection of visual and response signals. These findings challenge multi-step models of attention, and suggest that frontoparietal networks, which process sequential stimuli as separate events sharing the same location, drive exogenous attention phenomena such as inhibition of return.

A bio-inspired exogenous attention-based architecture for social robots

Social robots are increasingly becoming part of our society. In this sense, such robots’ interaction capabilities depend largely on their ability to detect and respond to stimuli in their environment as humans would. This paper describes a bio-inspired exogenous attention-based perception architecture for social robots. Exogenous attention regulates those involuntary attentional processes driven by the salience of perceived stimuli. The architecture incorporates bio-inspired concepts, such as inhibition to return, focus of attention selection and detecting salient stimuli, that have been validated on a real social robot. We have integrated several multimodal mechanisms for recognising salient visual, auditory, and tactile stimuli. Then, the attention architecture’s performance was evaluated by comparing it to a ground truth gathered from interactions of 50 users, obtaining comparable responses in real-time.

Mice with an autism-associated R451C mutation in neuroligin-3 show intact attention orienting but atypical responses to methylphenidate and atomoxetine in the mouse-Posner task.

Atypical attention orienting has been associated with some autistic symptoms, but the neural mechanisms remain unclear. The human Posner task, a classic attention orienting paradigm, was recently adapted for use with mice, supporting the investigation of the neurobiological underpinnings of atypical attention orienting in preclinical mouse models. The current study tested mice expressing the autism-associated R451C gene mutation in neuroligin-3 (NL3) on the mouse-Posner (mPosner) task. NL3R451C and wild-type (WT) mice were trained to respond to a validly or invalidly cued target on a touchscreen. The cue was a peripheral non-predictive flash in the exogenous task and a central spatially predictive image in the endogenous task. The effects of dopaminergic- and noradrenergic-modulating drugs, methylphenidate and atomoxetine, on task performance were assessed. In both tasks, mice were quicker and more accurate in the validly versus invalidly cued trials, consistent with results in the human Posner task. NL3R451C and WT mice showed similar response times and accuracy but responded differently when treated with methylphenidate and atomoxetine. Methylphenidate impaired exogenous attention disengagement in NL3R451C mice but did not significantly affect WT mice. Atomoxetine impaired endogenous orienting in WT mice but did not significantly affect NL3R451C mice. NL3R451C mice demonstrated intact attention orienting but altered responses to the pharmacological manipulation of the dopaminergic and noradrenergic networks. These findings expand our understanding of the NL3R451C mutation by suggesting that this mutation may lead to selective alterations in attentional processes.

A lateralized alpha-band marker of the interference of exogenous attention over endogenous attention.

Scientists differentiate top-down from bottom-up visuospatial attention processes. While bottom-up attention is rapidly engaged by emerging demands from the environment, top-down attention in contrast reflects slow voluntary shifts of spatial attention. Several lines of research substantiate the idea that top-down and bottom-up attentional processes involve distinct functional systems. An increasing number of studies, however, argue that both attention systems share brain processing resources. The current study examines how salient visual events that engage bottom-up processes interfere with top-down attentional processes. Using neurophysiological recordings and multivariate pattern classification techniques, the authors show that these patterns of interference occur within the alpha-band of neural activity (8-12Hz), which implies that bottom-up and top-down attention processes share this narrow-band frequency brain resource. The results further demonstrate that patterns of alpha-band activity explains, in part, the interference between top-down and bottom-up attention at the behavioral level.

Invited Session I: 30 years of normalization in the visual system: Testing and expanding the Reynolds & Heeger Normalization Model of Attention.

I present evidence supporting key predictions of the Reynolds-Heeger Normalization Model of Attention: 1) Both exogenous and endogenous covert spatial attention, can affect performance by contrast or response gain changes, depending on the stimulus size and the relative size of the attention field. 2) Feature-based attention improves performance, consistent with a change in response gain, when the featural extent of the attention field is small (low uncertainty) or large (high uncertainty) relative to the featural extent of the stimulus. 3) An expanded model of spatial resolution and attention that distinguishes between exogenous and endogenous attention, focusing on texture-based segmentation as a model system, which has revealed a dissociation between these types of attention across eccentricity. Our model emulates sensory encoding to segment figures from their background and predict performance. To explain attentional effects, exogenous and endogenous attention require separate operating regimes across spatial frequency. The model reproduces behavioral performance across several experiments and resolves three unexplained phenomena: 1) the parafoveal advantage in segmentation, 2) the uniform improvements across eccentricity by endogenous attention, and 3) the peripheral improvements and central impairments by exogenous attention. This model provides a generalizable framework for predicting effects of endogenous and exogenous attention on perception across the visual field.

Children with dyslexia show no deficit in exogenous spatial attention but show differences in visual encoding.

In the search for mechanisms that contribute to dyslexia, the term "attention" has been invoked to explain performance in a variety of tasks, creating confusion since all tasks do, indeed, demand "attention." Many studies lack an experimental manipulation of attention that would be necessary to determine its influence on task performance. Nonetheless, an emerging view is that children with dyslexia have an impairment in the exogenous (automatic/reflexive) orienting of spatial attention. Here we investigated the link between exogenous attention and reading ability by presenting exogenous spatial cues in the multi-letter processing task-a task relevant for reading. The task was gamified and administered online to a large sample of children (N=187) between 6 and 17 years. Children with dyslexia performed worse overall at rapidly recognizing and reporting strings of letters. However, we found no evidence for a difference in the utilization of exogenous spatial cues, resolving two decades of ambiguity in the field. Previous studies that claimed otherwise may have failed to distinguish attention effects from overall task performance or found spurious group differences in small samples. RESEARCH HIGHLIGHTS: We manipulated exogenous visual spatial attention using pre-cues in a task that is relevant for reading and we see robust task effects of exogenous attention. We found no evidence for a deficit in utilizing exogenous spatial pre-cues in children with dyslexia. However, children with dyslexia showed reduced recognition ability for all letter positions. Children with dyslexia were just as likely to make letter transposition errors as typical readers.

Crossmodal interference on counting performance: Evidence for shared attentional resources.

During the act of counting, our perceptual system may rely on information coming from different sensory channels. However, when the information coming from different sources is discordant, such as in the case of a de-synchronization between visual stimuli to be counted and irrelevant auditory stimuli, the performance in a sequential counting task might deteriorate. Such deterioration may originate from two different mechanisms, both linked to exogenous attention attracted by auditory stimuli. Indeed, exogenous auditory triggers may infiltrate our internal "counter", interfering with the counting process, resulting in an overcount; alternatively, the exogenous auditory triggers may disrupt the internal "counter" by deviating participants' attention from the visual stimuli, resulting in an undercount. We tested these hypotheses by asking participants to count visual discs sequentially appearing on the screen while listening to task-irrelevant sounds, in systematically varied conditions: visual stimuli could be synchronized or de-synchronized with sounds; they could feature regular or irregular pacing; and their speed presentation could be fast (approx. 3/sec), moderate (approx. 2/sec), or slow (approx. 1.5/sec). Our results support the second hypothesis since participants tend to undercount visual stimuli in all harder conditions (de-synchronized, irregular, fast sequences). We discuss these results in detail, adding novel elements to the study of crossmodal interference.