Covert Orienting Of Attention Research Articles

Representational coding of overt and covert orienting of visuospatial attention in the frontoparietal network

Orienting of visuospatial attention refers to reallocation of attentional focus from one target or location to another and can occur either with (overt) or without (covert) eye movement. Although it has been demonstrated that both types of orienting commonly involve frontal and parietal brain regions as the frontoparietal network (FPN), the underlying representational coding of these two types of orienting remains unclear. In this functional magnetic resonance imaging study, participants performed a task that elicited overt and covert orienting to endogenously or exogenously cued targets with eye-tracking to monitor eye movement. Although the FPN was commonly activated for both overt and covert orienting, multivariate patterns of the activation of voxels in the FPN accurately predicted whether eye movements were involved or not during orienting. These overt- and covert-preferred voxels were topologically distributed as distinct and interlaced clusters in a millimeter scale. Inclusion of the two types of clusters predicted orienting type more accurately than one type of clusters alone. These findings suggest that overt and covert orienting are represented by interdependent functional clusters of neuronal populations in regions of the FPN, which might reflect a generalizable principle in the nervous system for functional organization of closely associated processes.

Does age-related hearing loss deteriorate attentional resources?

ABSTRACT Recent work suggests that age-related hearing loss (HL) is a possible risk factor for cognitive decline in older adults. Resulting poor speech recognition negatively impacts cognitive, social and emotional functioning and may relate to dementia. However, little is known about the consequences of hearing loss on other non-linguistic domains of cognition. The aim of this study was to investigate the role of HL on covert orienting of attention, selective attention and executive control. We compared older adults with and without mild to moderate hearing loss (26–60 dB) performing (1) a spatial cueing task with uninformative central cues (social vs. nonsocial cues), (2) a flanker task and (3) a neuropsychological assessment of attention. The results showed that overall response times and flanker interference effects were comparable across groups. However, in spatial cueing of attention using social and nonsocial cues, hearing impaired individuals were characterized by reduced validity effects, though no additional group differences were found between social and nonsocial cues. Hearing impaired individuals also demonstrated diminished performance on the Montreal Cognitive Assessment (MoCA) and on tasks requiring divided attention and flexibility. This work indicates that while response speed and response inhibition appear to be preserved following mild-to-moderate acquired hearing loss, orienting of attention, divided attention and the ability to flexibly allocate attentional resources are more deteriorated in older adults with HL. This work suggests that hearing loss might exacerbate the detrimental influences of aging on visual attention.

Orienting of covert attention by neutral and emotional gaze cues appears to be unaffected by mild to moderate amblyopia.

Amblyopia is a developmental disorder of vision associated with higher-order visual attention deficits. We explored whether amblyopia affects the orienting of covert spatial attention by measuring the magnitude of the gaze cueing effect from emotional faces. Gaze and emotion cues are key components of social attention. Participants with normal vision (n = 30), anisometropic (n = 7) or strabismic/mixed (n = 5) amblyopia performed a cued peripheral target detection task under monocular and binocular viewing conditions. The cue consisted of a centrally presented face with left or right gaze (50% validity to target location) and a fearful, happy, or neutral expression. The magnitude of spatial cueing was computed as the reaction time difference between congruent and incongruent trials for each expression. Fearful facial expressions oriented spatial attention significantly more than happy or neutral expressions. The magnitude of the gaze cueing effect in our cohort of mild-to-moderate amblyopia was comparable to that in normal vision and was not correlated with the severity of amblyopia. There were no statistical group or amblyopia subtype differences for reaction time in any viewing condition. These results place constraints on the range of attentional mechanisms affected by amblyopia and possibly suggest normal covert processing of emotional face stimuli in mild and moderate amblyopia.

Eye movements are not mandatorily preceded by the N2pc component.

Researchers typically distinguish between two mechanisms of attentional selection in vision: overt and covert attention. A commonplace assumption is that overt eye movements are automatically preceded by shifts of covert attention during visual search. Although the N2pc component is a putative index of covert attentional orienting, little is currently known about its relationship with overt eye movements. This is because most previous studies of the N2pc component prohibit overt eye movements. The current study assessed this relationship by concurrently measuring covert attention (via the N2pc) and overt eye movements (via eye tracking). Participants searched displays for a lateralized target stimulus and were allowed to generate overt eye movements during the search. We then assessed whether overt eye movements were preceded by the N2pc component. The results indicated that saccades were preceded by an N2pc component, but only when participants were required to carefully inspect the target stimulus before initiating the eye movement. When participants were allowed to make naturalistic eye movements in service of visual search, there was no evidence of an N2pc component before eye movements. These findings suggest that the N2pc component does not always precede overt eye movements during visual search. Implications for understanding the relationship between covert and overt attention are discussed.

Decoding the Temporal Dynamics of Covert Spatial Attention Using Multivariate EEG Analysis: Contributions of Raw Amplitude and Alpha Power.

Attention can be oriented in space covertly without the need of eye movements. We used multivariate pattern classification analyses (MVPA) to investigate whether the time course of the deployment of covert spatial attention leading up to the observer’s perceptual decision can be decoded from both EEG alpha power and raw activity traces. Decoding attention from these signals can help determine whether raw EEG signals and alpha power reflect the same or distinct features of attentional selection. Using a classical cueing task, we showed that the orientation of covert spatial attention can be decoded by both signals. However, raw activity and alpha power may reflect different features of spatial attention, with alpha power more associated with the orientation of covert attention in space and raw activity with the influence of attention on perceptual processes.

Covert orienting of attention in 3-month-old infants: The case of biological motion

Humans attend to different positions in the space either by moving their eyes or by moving covertly their attention. The development of covert attention occurs during the first year of life. According to Colombo’s model of attention (2001), within the first years there is a significant change in infants’ visuo-spatial orienting mechanisms, from a predominantly overt form to a covert orienting starting from 4 to 5 months of life. The use of non-invasive brain imaging techniques can shed light on the origin of such mechanisms. In particular, EEG and ERP studies can directly investigate the neural correlates of covert attention in young infants. The present study investigated the neural correlates of covert attention employing a visuo-spatial cueing paradigm in 3-month-old infants. Infants were presented with a central point-light walker (PLD) followed by a single peripheral target. The target appeared randomly at a position either congruent or incongruent with the walking direction of the cue. We examined infants’ target-locked P1 component and the saccade latencies toward the peripheral target. Results showed that the P1 component was larger in response to congruent than to incongruent targets and saccade latencies were faster for congruent rather than incongruent trials. Moreover, the facilitation in processing sensory information (priming effects) presented at the cued spatial location occurs even before the onset of the oculomotor response, suggesting that covert attention is present before 4 months of age. Overall, this study highlights how ERPs method could help researchers at investigating the neural basis of attentional mechanisms in infants.

Exogenous Covert Orientation of Attention to the Center of Mass

Exogenous Covert Orientation of Attention to the Center of Mass

A Study of the Midbrain Network for Covert Attentional Orienting in Cervical Dystonia Patients using Dynamic Causal Modelling.

Understanding the neuronal network dynamics underlying the third most common movement disorder, cervical dystonia, can be achieved using dynamic causal modelling. Current literature establishes structures of the midbrain network for covert attentional orienting as dysfunctional in patients with cervical dystonia. One of these structures is the superior colliculus, for which it is hypothesised that deficient GABAergic activity therein causes cervical dystonia. To understand the role that this node plays in cervical dystonia, various connectivity models of the midbrain network were compared under the influence of a loom-recede visual stimulus fMRI paradigm. These models included the thalamus and striatum, crucial nodes in the direct/indirect pathways for motor movement and inhibition. The parametric empirical Bayes approach was used to quantify the difference in connection strengths across the winning models between patients and controls. Our findings demonstrated greater modulation by a looming stimulus event on the strength of connection from the striatum to the superior colliculus in patients. These results offer new means to understanding the pathophysiology of cervical dystonia.

Visual attention is not limited to the oculomotor range

Both patients with eye movement disorders and healthy participants whose oculomotor range had been experimentally reduced have been reported to show attentional deficits at locations unreachable by their eyes. Whereas previous studies were mainly based on the evaluation of reaction times, we measured visual sensitivity before saccadic eye movements and during fixation at locations either within or beyond participants' oculomotor range. Participants rotated their heads to prevent them from performing large rightward saccades. In this posture, an attentional cue was presented inside or outside their oculomotor range. Participants either made a saccade to the cue or maintained fixation while they discriminated the orientation of a visual noise patch. In contrast to previous reports, we found that the cue attracted visual attention regardless of whether it was presented within or beyond participants' oculomotor range during both fixation and saccade preparation. Moreover, when participants aimed to look to a cue that they could not reach with their eyes, we observed no benefit at their actual saccade endpoint. This demonstrates that spatial attention is not coupled to the executed oculomotor program but instead can be deployed unrestrictedly also toward locations to which no saccade can be executed. Our results are compatible with the view that covert and overt attentional orienting are guided by feedback projections of visual and visuomotor neurons of the gaze control system, irrespective of oculomotor limitations.

The association between reading abilities and visual-spatial attention in Hong Kong Chinese children.

A growing body of research suggests that visual-spatial attention is important for reading achievement. However, few studies have been conducted in non-alphabetic orthographies. This study extended the current research to reading development in Chinese, a logographic writing system known for its visual complexity. Eighty Hong Kong Chinese children were selected and divided into poor reader and typical reader groups, based on their performance on the measures of reading fluency, Chinese character reading, and reading comprehension. The poor and typical readers were matched on age and nonverbal intelligence. A Posner's spatial cueing task was adopted to measure the exogenous and endogenous orienting of visual-spatial attention. Although the typical readers showed the cueing effect in the central cue condition (i.e., responses to targets following valid cues were faster than those to targets following invalid cues), the poor readers did not respond differently in valid and invalid conditions, suggesting an impairment of the endogenous orienting of attention. The two groups, however, showed a similar cueing effect in the peripheral cue condition, indicating intact exogenous orienting in the poor readers. These findings generally supported a link between the orienting of covert attention and Chinese reading, providing evidence for the attentional-deficit theory of dyslexia.

The effect of offset cues on saccade programming and covert attention.

Salient peripheral events trigger fast, “exogenous” covert orienting. The influential premotor theory of attention argues that covert orienting of attention depends upon planned but unexecuted eye-movements. One problem with this theory is that salient peripheral events, such as offsets, appear to summon attention when used to measure covert attention (e.g., the Posner cueing task) but appear not to elicit oculomotor preparation in tasks that require overt orienting (e.g., the remote distractor paradigm). Here, we examined the effects of peripheral offsets on covert attention and saccade preparation. Experiment 1 suggested that transient offsets summoned attention in a manual detection task without triggering motor preparation planning in a saccadic localisation task, although there were a high proportion of saccadic capture errors on “no-target” trials, where a cue was presented but no target appeared. In Experiment 2, “no-target” trials were removed. Here, transient offsets produced both attentional facilitation and faster saccadic responses on valid cue trials. A third experiment showed that the permanent disappearance of an object also elicited attentional facilitation and faster saccadic reaction times. These experiments demonstrate that offsets trigger both saccade programming and covert attentional orienting, consistent with the idea that exogenous, covert orienting is tightly coupled with oculomotor activation. The finding that no-go trials attenuates oculomotor priming effects offers a way to reconcile the current findings with previous claims of a dissociation between covert attention and oculomotor control in paradigms that utilise a high proportion of catch trials.

Measurement & Analysis of the Temporal Discrimination Threshold Applied to Cervical Dystonia.

The temporal discrimination threshold (TDT) is the shortest time interval at which an observer can discriminate two sequential stimuli as being asynchronous (typically 30-50 ms). It has been shown to be abnormal (prolonged) in neurological disorders, including cervical dystonia, a phenotype of adult onset idiopathic isolated focal dystonia. The TDT is a quantitative measure of the ability to perceive rapid changes in the environment and is considered indicative of the behavior of the visual neurons in the superior colliculus, a key node in covert attentional orienting. This article sets out methods for measuring the TDT (including two hardware options and two modes of stimuli presentation). We also explore two approaches of data analysis and TDT calculation. The application of the assessment of temporal discrimination to the understanding of the pathogenesis of cervical dystonia and adult onset idiopathic isolated focal dystonia is also discussed.

Where Is Your Attention? Assessing Individual Instances of Covert Attentional Orienting in Response to Gaze and Arrow Cues.

Humans spontaneously follow where others are looking. However, recent investigations suggest such gaze-following behavior during natural interactions occurs relatively infrequently, only in about a third of available instances. Here we investigated if a similar frequency of orienting is also found in laboratory tasks that measure covert attentional orienting using manual responses. To do so, in two experiments, we analyzed responses from a classic gaze cuing task, with arrow cues serving as control stimuli. We reasoned that the proportions of attentional benefits and costs, defined as responses falling outside of 1 standard deviation of the average performance for the neutral condition, would provide a good approximation of individual instances of attentional shifts. We found that although benefits and costs occurred in less than half of trials, benefits emerged on a greater proportion of validly cued relative to invalidly cued trials. This pattern of data held across two different measures of neutral performance, as assessed by Experiments 1 and 2, as well as across the two cue types. These results suggest that similarly to gaze-following in naturalistic settings, covert orienting within the cuing task also appears to occur relatively infrequently.

Exercise-Induced Fatigue and Caffeine Supplementation Affect Psychomotor Performance but Not Covert Visuo-Spatial Attention.

Fatigue resulting from strenuous exercise can impair cognition and oculomotor control. These impairments can be prevented by administering psychostimulants such as caffeine. This study used two experiments to explore the influence of caffeine administered at rest and during fatiguing physical exercise on spatial attention—a cognitive function that is crucial for task-based visually guided behavior. In independent placebo-controlled studies, cohorts of 12 healthy participants consumed caffeine and rested or completed 180 min of stationary cycling. Covert attentional orienting was measured in both experiments using a spatial cueing paradigm. We observed no alterations in attentional facilitation toward spatial cues suggesting that covert attentional orienting is not influenced by exercise fatigue or caffeine supplementation. Response times were increased (impaired) after exercise and this deterioration was prevented by caffeine supplementation. In the resting experiment, response times across all conditions and cues were decreased (improved) with caffeine. Covert spatial attention was not influenced by caffeine. Together, the results of these experiments suggest that covert attentional orienting is robust to the effects of fatiguing exercise and not influenced by caffeine. However, exercise fatigue impairs response times, which can be prevented by caffeine, suggesting that pre-motor planning and execution of the motor responses required for performance of the cueing task are sensitive to central nervous system fatigue. Caffeine improves response time in both fatigued and fresh conditions, most likely through action on networks controlling motor function.

Covert preparation of a manual response in a 'go'/'no-go' saccadic task is driven by execution of the eye movement and not by visual stimulus occurrence.

It has been recently demonstrated that visually guided saccades are linked to changes in muscle excitability in the relaxed upper limb, which are compatible with a covert motor plan encoding a hand movement toward the gaze target. In this study we investigated whether these excitability changes are time locked to the visual stimulus, as predicted by influential attention models, or are strictly dependent on saccade execution. Single-pulse transcranial magnetic stimulation was applied to the motor cortex at eight different time delays during a ‘go’/‘no-go’ task, which involved overt or covert orienting of attention. By analyzing the time course of excitability in three hand muscles, synchronized with the onset of either the attentional cue or the eye movement, we demonstrated that side- and muscle-specific excitability changes were strictly time locked to the saccadic response and were not correlated to the onset of the visual attentive stimulus. Furthermore, muscle excitability changes were absent following a covert shift of attention. We conclude that a sub-threshold manual motor plan is automatically activated by the saccade decision-making process, as part of a covert eye-hand coordination program. We found no evidence for a representation of spatial attention within the upper limb motor map.

Effects of social stimuli on covert attentional orienting and saccaddic eye-movements during visual search

Effects of social stimuli on covert attentional orienting and saccaddic eye-movements during visual search

Differential Effects of Lowered Arousal on Covert and Overt Shifts of Attention.

Based on previous studies demonstrating detrimental effects of reduced alertness on attentional orienting our study seeks to examine covert and overt attentional orienting in different arousal states. We hypothesized an attentional asymmetry with increasing reaction times to stimuli presented to the left visual field in a state of maximally reduced arousal. Eleven healthy participants underwent sleep deprivation and were examined repeatedly every 4 hr over 28 hr in total with two tasks measuring covert and overt orienting of attention. Contrary to our hypothesis, a reduction of arousal did not induce any asymmetry of overt orienting. Even in participants with profound and significant attentional asymmetries in covert orienting no substantial reaction time differences between left- and right-sided targets in the overt orienting task could be observed. This result is not in agreement with assumptions of a tight coupling of covert and overt attentional processes. In conclusion, we found differential effects of lowered arousal induced by sleep deprivation on covert and overt orienting of attention. This pattern of results points to a neuronal non-overlap of brain structures subserving these functions and a differential influence of the norepinephrine system on these modes of spatial attention.

Neural correlates of the preserved inhibition of return in schizophrenia.

Inhibition of return (IOR) is an attentional mechanism that previously has been reported to be either intact or blunted in subjects with schizophrenia (SCZ). In the present study, we explored the neural mechanism of IOR in SCZ by comparing the target-locked N1 and P1 activity evoked by valid-cued trials with that evoked by invalid-cued trials. Twenty-seven schizophrenia patients and nineteen healthy controls participated in a task involving covert orienting of attention with two stimulus onset asynchronies (SOAs: 700 ms and 1200 ms) during which 64-channel EEG data were recorded. Behavioral reaction times (RTs) were longer in response to valid-cued trials than to invalid-cued ones, suggesting an intact IOR in SCZ. However, reduced N1 amplitude elicited by valid-cued trials suggested a stronger inhibition of attention from being oriented to a previously cued location, and therefore a relative inhibition of perceptual processing at that location in SCZ. These results indicate that altered N1 activity is associated with the preservation of IOR in SCZ and could be a sensitive marker to track the IOR effect.

Modulation of time perception by eye movements

Event Abstract Back to Event Modulation of time perception by eye movements Cheng Xiaoqin1, 2 and Trevor B. Penney2, 3* 1 National University of Singapore, Graduate School for Integrative Sciences and Engineering, Singapore 2 National University of Singapore, LSI Programme in Neurobiology and Aging, Singapore 3 National University of Singapore, Psychology, Singapore The perception of time is intricately linked to attention. Overt shifting of one’s attention involves foveating to the area of interest through ballistic eye movements known as saccades. However, the direction of gaze does not always indicate one’s attentional focus because attention shifts also occur covertly in the absence of saccades. Microsaccades are small eye movements that occur during eye fixations and underlie covert attention shifts (Engbert and Kliegl, 2003). In the present study, we investigated whether saccades and microsaccades impact interval timing. Participants were asked to memorise the duration of a six-second visual (Experiment 1) or auditory (Experiment 2) stimulus and to then reproduce that duration in subsequent test trials. In three of the four test conditions, the interval timing trials included a secondary cue discrimination task. Participants were required to foveate to stimuli appearing in the periphery (Saccade-Discrimination), maintain fixation in the center despite appearance of peripheral stimuli (No Saccade-Peripheral Discrimination), or maintain central fixation with stimuli appearing in the center (No Saccade-Central Discrimination). In the fourth condition, participants did not perform saccades or the discrimination task (No Saccade-No Discrimination). Using mixed effects modelling, we examined the relationship between saccade duration and reproduced duration in the Saccade-Discrimination condition and between microsaccade duration and reproduced duration in all No Saccade conditions. Saccade duration positively predicted reproduced duration in both experiments. In addition, microsaccade duration positively predicted reproduced duration for all No Saccade conditions in Experiment 2. However, this relationship was only found in the No Saccade-Central Discrimination and the No Saccade-No Discrimination condition for Experiment 1. Taken together, the results suggest a role of both saccades and microsaccades in visual attention and time perception. References Engbert, R., and Kliegl, R. (2003). Microsaccades uncover the orientation of covert attention. Vision Research, 43, 1035-1045. Keywords: Time Perception, microsaccades, Saccades, Time distortion, Eye Movements, Seconds range Conference: ASP2015 - 25th Annual Conference of the Australasian Society for Psychophysiology, Sydney, Australia, 2 Dec - 4 Dec, 2015. Presentation Type: Oral Presentation Topic: Psychophysiology Citation: Xiaoqin C and Penney TB (2015). Modulation of time perception by eye movements. Conference Abstract: ASP2015 - 25th Annual Conference of the Australasian Society for Psychophysiology. doi: 10.3389/conf.fnhum.2015.219.00049 Copyright: The abstracts in this collection have not been subject to any Frontiers peer review or checks, and are not endorsed by Frontiers. They are made available through the Frontiers publishing platform as a service to conference organizers and presenters. The copyright in the individual abstracts is owned by the author of each abstract or his/her employer unless otherwise stated. Each abstract, as well as the collection of abstracts, are published under a Creative Commons CC-BY 4.0 (attribution) licence (https://creativecommons.org/licenses/by/4.0/) and may thus be reproduced, translated, adapted and be the subject of derivative works provided the authors and Frontiers are attributed. For Frontiers’ terms and conditions please see https://www.frontiersin.org/legal/terms-and-conditions. Received: 25 Oct 2015; Published Online: 30 Nov 2015. * Correspondence: Dr. Trevor B Penney, National University of Singapore, LSI Programme in Neurobiology and Aging, Singapore, Singapore, penney@cuhk.edu.hk Login Required This action requires you to be registered with Frontiers and logged in. To register or login click here. Abstract Info Abstract The Authors in Frontiers Cheng Xiaoqin Trevor B Penney Google Cheng Xiaoqin Trevor B Penney Google Scholar Cheng Xiaoqin Trevor B Penney PubMed Cheng Xiaoqin Trevor B Penney Related Article in Frontiers Google Scholar PubMed Abstract Close Back to top Javascript is disabled. Please enable Javascript in your browser settings in order to see all the content on this page.

The role of the collicular pathway in the salience-based progression of visual attention

Visual attention has been shown to progress from the most to the least salient item in a given scene. Cognitive and physiological models assume that this orienting of covert attention relies on the collicular pathway, involving the superior colliculus and the pulvinar. Recent studies questioned this statement as they described attentional capture by visual items invisible to the superior colliculus. Electrophysiological studies shown that there is no direct projections from short-wave receptors to the superior colliculus. S-cone stimuli can thus be employed to assess visual processing without the involvement of the collicular pathway. We have attempted to investigate whether this pathway is involved in the salience-based orientation of attention by presenting S-cone stimuli. Volunteers were asked to make a judgment regarding a target among two distractors (all items of unequal sizes). Items’ location and size varied randomly, as well as color, randomly black or calibrated for each subject to activate exclusively S-cones. The hierarchical pattern testifying of the salience-based orientation of attention was only found with black stimuli, arguing in favor of an implication of the collicular pathway in salience. In a second experiment, one item was presented at a time in order to test the item-multiplicity effect by comparing experiments. Performance was the most penalized when presenting multiple stimuli in the black condition. Results were interpreted in terms of distinct modes of processing by the collicular and geniculate pathways. The establishment of salience that determines attentional progression appeared to be only possible when the collicular pathway was solicited.