Endogenous Spatial Attention Research Articles

Enhancing rhythmic temporal expectations: The dominance of auditory modality under spatial uncertainty.

To effectively process the most relevant information, the brain anticipates the optimal timing for allocating attentional resources. Behavior can be optimized by automatically aligning attention with external rhythmic structures, whether visual or auditory. Although the auditory modality is known for its efficacy in representing temporal information, the current body of research has not conclusively determined whether visual or auditory rhythmic presentations have a definitive advantage in entraining temporal attention. The present study directly examined the effects of auditory and visual rhythmic cues on the discrimination of visual targets in Experiment 1 and on auditory targets in Experiment 2. Additionally, the role of endogenous spatial attention was also considered. When and where the target was the most likely to occur were cued by unimodal (visual or auditory) and bimodal (audiovisual) signals. A sequence of salient events was employed to elicit rhythm-based temporal expectations and a symbolic predictive cue served to orient spatial attention. The results suggest a superiority of auditory over visual rhythms, irrespective of spatial attention, whether the spatial cue and rhythm converge or not (unimodal or bimodal), and regardless of the target modality (visual or auditory). These findings are discussed in terms of a modality-specific rhythmic orienting, while considering a single, supramodal system operating in a top-down manner for endogenous spatial attention.

On the relationship between rhythm-based temporal expectations and endogenous spatial attention in simple reaction-time tasks

IntroductionRecent evidence suggests that the temporal expectations afforded by a regular rhythmic structure operate independently from endogenous spatial attention in simple reaction-time tasks. The most common manipulation followed in previous studies consisted of presenting a target stimulus either “in-time” or “out-of-time” (earlier or later) with a preceding rhythm. However, contrary to the proposal of entrainment models predicting a behavioral advantage for in-time compared to both early and late targets, responses were still faster for late targets, according to the so-called “foreperiod effect”. This finding makes it difficult to fully disentangle the impact of rhythm and the benefits afforded by the passage of time on the relationship between rhythm-based temporal and endogenous spatial attention.MethodsTo shed more light on this issue, we combined a spatial orienting task with a rhythmic manipulation, in which two placeholders flanking the fixation cross flickered at either a regular or irregular pace. Spatial orienting to the target location was deployed by symbolic color cues that were displayed independently of the rhythm (independent cues) or were integrated with the temporal rhythmic cues (integrated cues). Crucially, for both independent and integrated cues, and for regular and irregular rhythms, the interval between the rhythm and the target (i.e., the foreperiod) was kept fixed in Experiments 1–3 to control for the foreperiod effect, while the effect of foreperiod was explored in Experiment 4.ResultsResults showed a more beneficial effect of rhythms with independent cues as compared to integrated cues. Additionally, the benefit of rhythms was slightly but significantly larger at valid, compared to invalid, spatial locations, regardless of the foreperiod.DiscussionOur results extend previous studies by showing that interactive effects of rhythms and endogenous spatial attention may emerge in low-demanding detection tasks.

Somatosensory Event-Related Potential as an Electrophysiological Correlate of Endogenous Spatial Tactile Attention: Prospects for Electrotactile Brain-Computer Interface for Sensory Training.

Tactile attention tasks are used in the diagnosis and treatment of neurological and sensory processing disorders, while somatosensory event-related potentials (ERP) measured by electroencephalography (EEG) are used as neural correlates of attention processes. Brain-computer interface (BCI) technology provides an opportunity for the training of mental task execution via providing online feedback based on ERP measures. Our recent work introduced a novel electrotactile BCI for sensory training, based on somatosensory ERP; however, no previous studies have addressed specific somatosensory ERP morphological features as measures of sustained endogenous spatial tactile attention in the context of BCI control. Here we show the morphology of somatosensory ERP responses induced by a novel task introduced within our electrotactile BCI platform i.e., the sustained endogenous spatial electrotactile attention task. By applying pulsed electrical stimuli to the two proximal stimulation hotspots at the user's forearm, stimulating sequentially the mixed branches of radial and median nerves with equal probability of stimuli occurrence, we successfully recorded somatosensory ERPs for both stimulation locations, in the attended and unattended conditions. Waveforms of somatosensory ERP responses for both mixed nerve branches showed similar morphology in line with previous reports on somatosensory ERP components obtained by stimulation of exclusively sensory nerves. Moreover, we found statistically significant increases in ERP amplitude on several components, at both stimulation hotspots, while sustained endogenous spatial electrotactile attention task is performed. Our results revealed the existence of general ERP windows of interest and signal features that can be used to detect sustained endogenous tactile attention and classify between spatial attention locations in 11 healthy subjects. The current results show that features of N140, P3a and P3b somatosensory ERP components are the most prominent global markers of sustained spatial electrotactile attention, over all subjects, within our novel electrotactile BCI task/paradigm, and this work proposes the features of those components as markers of sustained endogenous spatial tactile attention in online BCI control. Immediate implications of this work are the possible improvement of online BCI control within our novel electrotactile BCI system, while these finding can be used for other tactile BCI applications in the diagnosis and treatment of neurological disorders by employing mixed nerve somatosensory ERPs and sustained endogenous electrotactile attention task as control paradigms.

Visual-cortical enhancement by acoustic distractors: The effects of endogenous spatial attention and visual working memory load

Past work has shown that when a peripheral sound captures our attention, it activates the contralateral visual cortex as revealed by an event-related potential component labelled the auditory-evoked contralateral occipital positivity (ACOP). This cross-modal activation of the visual cortex has been observed even when the sounds were not relevant to the ongoing task (visual or auditory), suggesting that peripheral sounds automatically activate the visual cortex. However, it is unclear whether top-down factors such as visual working memory (VWM) load and endogenous attention, which modulate the impact of task-irrelevant information, may modulate this spatially-specific component. Here, we asked participants to perform a lateralized VWM task (change detection), whose performance is supported by both endogenous spatial attention and VWM storage. A peripheral sound that was unrelated to the ongoing task was delivered during the retention interval. The amplitude of sound-elicited ACOP was analyzed as a function of the spatial correspondence with the cued hemifield, and of the memory array set-size. The typical ACOP modulation was observed over parieto-occipital sites in the 280–500 ms time window after sound onset. Its amplitude was not affected by VWM load but was modulated when the location of the sound did not correspond to the hemifield (right or left) that was cued for the change detection task. Our results suggest that sound-elicited activation of visual cortices, as reflected in the ACOP modulation, is unaffected by visual working memory load. However, endogenous spatial attention affects the ACOP, challenging the hypothesis that it reflects an automatic process.

Exogenous and endogenous spatial attention in crows

Attention describes the ability to selectively process a particular aspect of the environment at the expense of others. Despite the significance of selective processing, the types and scopes of attentional mechanisms in nonprimate species remain underexplored. We trained four carrion crows in Posner spatial cueing tasks using two separate protocols where the attention-capturing cues are shown at different times before target onset at either the same or a different location as the impending target. To probe automatic bottom-up, or exogenous, attention, two naïve crows were tested with a cue that had no predictive value concerning the location of the subsequent target. To examine volitional top-down, or endogenous, attention, the other two crows were tested with the previously learned cues that predicted the impending target location. Comparing the performance for valid (cue and target at same location) and invalid (cue and target at opposing locations) cues in the nonpredictive cue condition showed a transient, mild reaction time advantage signifying exogenous attention. In contrast, there was a strong and long-lasting performance advantage for the valid conditions with predictive cues indicating endogenous attention. Together, these results demonstrate that crows possess two different attention mechanisms (exogenous and endogenous). These findings signify that crows possess a substantial attentional capacity and robust cognitive control over attention allocation.

Benefits of Endogenous Spatial Attention During Visual Double-Training in Cortically-Blinded Fields.

Recovery of visual discrimination thresholds inside cortically-blinded (CB) fields is most commonly attained at a single, trained location at a time, with iterative progress deeper into the blind field as performance improves over several months. As such, training is slow, inefficient, burdensome, and often frustrating for patients. Here, we investigated whether double-location training, coupled with a covert spatial-attention (SA) pre-cue, could improve the efficiency of training. Nine CB participants completed a randomized, training assignment with either a spatial attention or neutral pre-cue. All trained for a similar length of time on a fine direction discrimination task at two blind field locations simultaneously. Training stimuli and tasks for both cohorts were identical, save for the presence of a central pre-cue, to manipulate endogenous (voluntary) SA, or a Neutral pre-cue. Participants in the SA training cohort demonstrated marked improvements in direction discrimination thresholds, albeit not to normal/intact-field levels; participants in the Neutral training cohort remained impaired. Thus, double-training within cortically blind fields, when coupled with SA pre-cues can significantly improve direction discrimination thresholds at two locations simultaneously, offering a new method to improve performance and reduce the training burden for CB patients. Double-training without SA pre-cues revealed a hitherto unrecognized limitation of cortically-blind visual systems’ ability to improve while processing two stimuli simultaneously. These data could potentially explain why exposure to the typically complex visual environments encountered in everyday life is insufficient to induce visual recovery in CB patients. It is hoped that these new insights will direct both research and therapeutic developments toward methods that can attain better, faster recovery of vision in CB fields.

Dynamic causal interactions between occipital and parietal cortex explain how endogenous spatial attention and stimulus-driven salience jointly shape the distribution of processing priorities in 2D visual space

Visuo-spatial attention prioritizes the processing of relevant inputs via different types of signals, including current goals and stimulus salience. Complex mixtures of these signals engage in everyday life situations, but little is known about how these signals jointly modulate distributed patterns of activity across the occipital regions that represent visual space. Here, we measured spatio-topic, quadrant-specific occipital activity during the processing of visual displays containing both task-relevant targets and salient color-singletons. We computed spatial bias vectors indexing the effect of attention in 2D space, as coded by distributed activity in the occipital cortex. We found that goal-directed spatial attention biased activity towards the target and that salience further modulated this endogenous effect: salient distractors decreased the spatial bias, while salient targets increased it. Analyses of effective connectivity revealed that the processing of salient distractors relied on the modulation of the bidirectional connectivity between the occipital and the posterior parietal cortex, as well as the modulation of the lateral interactions within the occipital cortex. These findings demonstrate that goal-directed attention and salience jointly contribute to shaping processing priorities in the occipital cortex and highlight that multiple functional paths determine how spatial information about these signals is distributed across occipital regions.

Effects of endogenous spatial attention and exogenous spatial attention on multisensory integration

Effects of endogenous spatial attention and exogenous spatial attention on multisensory integration

A Stable Population Code for Attention in Prefrontal Cortex Leads a Dynamic Attention Code in Visual Cortex.

Attention often requires maintaining a stable mental state over time while simultaneously improving perceptual sensitivity. These requirements place conflicting demands on neural populations, as sensitivity implies a robust response to perturbation by incoming stimuli, which is antithetical to stability. Functional specialization of cortical areas provides one potential mechanism to resolve this conflict. We reasoned that attention signals in executive control areas might be highly stable over time, reflecting maintenance of the cognitive state, thereby freeing up sensory areas to be more sensitive to sensory input (i.e., unstable), which would be reflected by more dynamic attention signals in those areas. To test these predictions, we simultaneously recorded neural populations in prefrontal cortex (PFC) and visual cortical area V4 in rhesus macaque monkeys performing an endogenous spatial selective attention task. Using a decoding approach, we found that the neural code for attention states in PFC was substantially more stable over time compared with the attention code in V4 on a moment-by-moment basis, in line with our guiding thesis. Moreover, attention signals in PFC predicted the future attention state of V4 better than vice versa, consistent with a top-down role for PFC in attention. These results suggest a functional specialization of attention mechanisms across cortical areas with a division of labor. PFC signals the cognitive state and maintains this state stably over time, whereas V4 responds to sensory input in a manner dynamically modulated by that cognitive state.SIGNIFICANCE STATEMENT Attention requires maintaining a stable mental state while simultaneously improving perceptual sensitivity. We hypothesized that these two demands (stability and sensitivity) are distributed between prefrontal and visual cortical areas, respectively. Specifically, we predicted attention signals in visual cortex would be less stable than in prefrontal cortex, and furthermore prefrontal cortical signals would predict attention signals in visual cortex in line with the hypothesized role of prefrontal cortex in top-down executive control. Our results are consistent with suggestions deriving from previous work using separate recordings in the two brain areas in different animals performing different tasks and represent the first direct evidence in support of this hypothesis with simultaneous multiarea recordings within individual animals.

Endogenous Spatial Attention Modulates the Magnitude of the Colavita Visual Dominance Effect.

The Colavita effect refers to the phenomenon wherein people tend to not respond to an auditory stimulus when a visual stimulus is simultaneously presented. Although previous studies have shown that endogenous modality attention influences the Colavita effect, whether the Colavita effect is influenced by endogenous spatial attention remains unknown. In the present study, we established endogenous spatial cues to investigate whether the size of the Colavita effect changes under visual or auditory cues. We measured three indexes to investigate the effect of endogenous spatial attention on the size of the Colavita effect. These three indexes were developed based on the following observations in bimodal trials: (a) The proportion of the “only vision” response was significantly higher than that of the “only audition” response; (b) the proportion of the “vision precedes audition” response was significantly higher than that of the “audition precedes vision” response; and (c) the reaction time difference of the “vision precedes audition” response was significantly higher than that of the “audition precedes vision” response. Our results showed that the Colavita effect was always influenced by endogenous spatial attention and that its size was larger at the cued location than at the uncued location; the cue modality (visual vs. auditory) had no effect on the size of the Colavita effect. Taken together, the present results shed light on how endogenous spatial attention affects the Colavita effect.

Voluntary attention improves performance similarly around the visual field.

Performance as a function of polar angle at isoeccentric locations across the visual field is known as a performance field (PF) and is characterized by two asymmetries: the HVA (horizontal-vertical anisotropy) and VMA (vertical meridian asymmetry). Exogenous (involuntary) spatial attention does not affect the shape of the PF, improving performance similarly across polar angle. Here we investigated whether endogenous (voluntary) spatial attention, a flexible mechanism, can attenuate these perceptual asymmetries. Twenty participants performed an orientation discrimination task while their endogenous attention was either directed to the target location or distributed across all possible locations. The effects of attention were assessed either using the same stimulus contrast across locations or equating difficulty across locations using individually titrated contrast thresholds. In both experiments, endogenous attention similarly improved performance at all locations, maintaining the canonical PF shape. Thus, despite its voluntary nature, like exogenous attention, endogenous attention cannot alleviate perceptual asymmetries at isoeccentric locations.

Incentive value and spatial certainty combine additively to determine visual priorities

How does the brain combine information predictive of the value of a visually guided task (incentive value) with information predictive of where task-relevant stimuli may occur (spatial certainty)? Human behavioural evidence indicates that these two predictions may be combined additively to bias visual selection (Additive Hypothesis), whereas neuroeconomic studies posit that they may be multiplicatively combined (Expected Value Hypothesis). We sought to adjudicate between these two alternatives. Participants viewed two coloured placeholders that specified the potential value of correctly identifying an imminent letter target if it appeared in that placeholder. Then, prior to the target’s presentation, an endogenous spatial cue was presented indicating the target’s more likely location. Spatial cues were parametrically manipulated with regard to the information gained (in bits). Across two experiments, performance was better for targets appearing in high versus low value placeholders and better when targets appeared in validly cued locations. Interestingly, as shown with a Bayesian model selection approach, these effects did not interact, clearly supporting the Additive Hypothesis. Even when conditions were adjusted to increase the optimality of a multiplicative operation, support for it remained. These findings refute recent theories that expected value computations are the singular mechanism driving the deployment of endogenous spatial attention. Instead, incentive value and spatial certainty seem to act independently to influence visual selection.

The coordinate system of endogenous spatial attention during smooth pursuit.

A central question in vision is whether spatial attention is represented in an eye-centered (retinotopic) or world-centered (spatiotopic) reference-frame. Most previous studies on this question focused on how coordinates are modulated across saccades. In the present study, we investigated the reference-frame of attention across smooth pursuit eye-movements using a goal-directed saccade task. In two experiments, participants were asked to pursue a moving target while attending to one or two grating stimuli. On each trial, one stimulus was constant in its retinal position and the other was constant in its spatial position. Upon detection of a slight change in stimulus orientation, participants were asked to stop pursuing and perform a fast saccade toward the modified stimulus. In the focused attention condition, they attended one, predefined, stimulus, and in the divided attention condition they attended both. In Experiment 1 the angle of the orientation change marking the target event was constant across participants and conditions. In Experiment 2, the angle was individually adapted to equate performance across participants and conditions. Findings of the two experiments were consistent and showed that the enhancement of mean visual sensitivity in the focused relative to the divided attention condition was similar in magnitude for both retinotopic and spatiotopic targets. This indicates that during smooth pursuit, endogenous attention was proportionally divided between targets in retinotopic and spatiotopic frames of reference.

Hemisphere-specific, differential effects of lateralized, occipital\u2013parietal \u03b1- versus \u03b3-tACS on endogenous but not exogenous visual-spatial attention

Orienting spatial attention has been associated with interhemispheric asymmetry of power in the α- and γ-band. Specifically, increased α-power has been linked to the inhibition of unattended sensory streams (e.g. the unattended visual field), while increased γ-power is associated with active sensory processing. Here, we aimed to differentially modulate endogenous and exogenous visual-spatial attention using transcranial alternating current stimulation (tACS). In a single-blind, within-subject design, participants performed several blocks of a spatial cueing task comprised of endogenous and exogenous cues while receiving lateralized α- or γ-tACS or no stimulation over left or right occipital-parietal areas. We found a significant, differential effect of α- and γ-tACS on endogenous (top-down) spatial attention but not on exogenous (bottom-up) attention. The effect was specific to tACS applied to the left hemisphere and driven by a modulation of attentional disengagement and re-orientation as measured during invalid trials. Our results indicate a causal role of α-/γ-oscillations for top-down (endogenous) attention. They may further suggest a left hemispheric dominance in controlling interhemispheric α-/γ-power asymmetry. The absence of an effect on exogenous attention may be indicative of a differential role of α-/γ-oscillations during different attention types or spatially distinct attentional subsystems.

Exogenous attention facilitates perceptual learning in visual acuity to untrained stimulus locations and features.

Visual perceptual learning (VPL) refers to the improvement in performance on a visual task due to practice. A hallmark of VPL is specificity, as improvements are often confined to the trained retinal locations or stimulus features. We have previously found that exogenous (involuntary, stimulus-driven) and endogenous (voluntary, goal-driven) spatial attention can facilitate the transfer of VPL across locations in orientation discrimination tasks mediated by contrast sensitivity. Here, we investigated whether exogenous spatial attention can facilitate such transfer in acuity tasks that have been associated with higher specificity. We trained observers for 3 days (days 2–4) in a Landolt acuity task (Experiment 1) or a Vernier hyperacuity task (Experiment 2), with either exogenous precues (attention group) or neutral precues (neutral group). Importantly, during pre-tests (day 1) and post-tests (day 5), all observers were tested with neutral precues; thus, groups differed only in their attentional allocation during training. For the Landolt acuity task, we found evidence of location transfer in both the neutral and attention groups, suggesting weak location specificity of VPL. For the Vernier hyperacuity task, we found evidence of location and feature specificity in the neutral group, and learning transfer in the attention group—similar improvement at trained and untrained locations and features. Our results reveal that, when there is specificity in a perceptual acuity task, exogenous spatial attention can overcome that specificity and facilitate learning transfer to both untrained locations and features simultaneously with the same training. Thus, in addition to improving performance, exogenous attention generalizes perceptual learning across locations and features.

Attribute amnesia can be modulated by foveal presentation and the pre-allocation of endogenous spatial attention.

Even in sparse visual environments, observers may not be able to report features of objects they have just encountered in a surprise question. Attribute amnesia and seeing without knowing describe report failures for irrelevant features of objects that have been processed to some extent in the primary task. Both phenomena are attributed to the exclusive selection of relevant information for memory consolidation or for awareness, respectively. While attribute amnesia was found even for irrelevant attributes of the target in the primary task, seeing without knowing was not observed when a single object was presented foveally. To elucidate this discrepancy, we examined report failures for irrelevant attributes of single target objects, which were presented either in the fovea or in the periphery, and either at cued or uncued locations. On a surprise trial, observers were able to report the irrelevant shape and color of the target object when it was presented foveally. However, presenting the same object just slightly away from the fovea led to report failures for shape. Introducing a valid peripheral cue prior to target presentation reduced report failures for shape when the cue was predictive of the target location, suggesting that the pre-allocation of endogenous spatial attention promoted the processing of irrelevant shape information. In accordance with previous research, we suggest that these modulations are due to differences in late selection for conscious awareness or consolidation in working memory.

On the link between attentional search and the oculomotor system: Is preattentive search restricted to the range of eye movements?

It has been proposed that covert visual search can be fast, efficient, and stimulus driven, particularly when the target is defined by a salient single feature, or slow, inefficient, and effortful when the target is defined by a nonsalient conjunction of features. This distinction between fast, stimulus-driven orienting and slow, effortful orienting can be related to the distinction between exogenous spatial attention and endogenous spatial attention. Several studies have shown that exogenous, covert orienting is limited to the range of saccadic eye movements, whereas covert endogenous orienting is independent of the range of saccadic eye movements. The current study examined whether covert visual search is affected in a similar way. Experiment 1 showed that covert visual search for feature singletons was impaired when stimuli were presented beyond the range of saccadic eye movements, whereas conjunction search was unaffected by array position. Experiment 2 replicated and extended this effect by measuring search times at 6 eccentricities. The impairment in covert feature search emerged only when stimuli crossed the effective oculomotor range and remained stable for locations further into the periphery, ruling out the possibility that the results of Experiment 1 were due to a failure to fully compensate for the effects of cortical magnification. The findings are interpreted in terms of biased competition and oculomotor theories of spatial attention. It is concluded that, as with covert exogenous orienting, biological constraints on overt orienting in the oculomotor system constrain covert, preattentive search.

P.443 Does alcohol use provoke relapse in patients with opioid use disorder stabilized on naltrexone?

Attention resources can be allocated in both space and time. Exogenous temporal attention can be driven by rhythmic events in our environment which automatically entrain periods of attention. Temporal expectancies can also be generated by the elapse of time, leading to foreperiod effects (the longer between a cue and imperative target, the faster the response). This study investigates temporal attention in touch and the influence of spatial orienting. In experiment 1, participants used bilateral tactile cues to orient endogenous spatial attention to the left or right hand where a unilateral tactile target was presented. This facilitated response times for attended over unattended targets. In experiment 2, the cue was unilateral and non-predictive of the target location resulting in inhibition of return. Importantly, the cue was rhythmic and targets were presented early, in synchrony or late in relation to the rhythmic cue. A foreperiod effect was observed in experiment 1 that was independent from any spatial attention effects. In experiment 2, in synchrony were slower compared to out of synchrony targets but only for cued and not uncued targets, suggesting the rhythm generates periods of exogenous inhibition. Taken together, temporal and spatial attention interact in touch, but only when both types of attention are exogenous. If the task requires endogenous spatial orienting, space and time are independent.

How exogenous spatial attention affects visual representation.

Orienting covert spatial attention to a target location enhances visual sensitivity and benefits performance in many visual tasks. How these attention-related improvements in performance affect the underlying visual representation of low-level visual features is not fully understood. Here we focus on characterizing how exogenous spatial attention affects the feature representations of orientation and spatial frequency. We asked observers to detect a vertical grating embedded in noise and performed psychophysical reverse correlation. Doing so allowed us to make comparisons with previous studies that utilized the same task and analysis to assess how endogenous attention and presaccadic modulations affect visual representations. We found that exogenous spatial attention improved performance and enhanced the gain of the target orientation without affecting orientation tuning width. Moreover, we found no change in spatial frequency tuning. We conclude that covert exogenous spatial attention alters performance by strictly boosting gain of orientation-selective filters, much like covert endogenous spatial attention.

Mathematics students demonstrate superior visuo-spatial working memory to humanities students under conditions of low central executive processing load

Previous research has demonstrated that working memory performance is linked to mathematics achievement. Most previous studies have involved children and arithmetic rather than more advanced forms of mathematics. This study compared the performance of groups of adult mathematics and humanities students. Experiment 1 employed verbal and visuo-spatial working memory span tasks using a novel face-matching processing element. Results showed that mathematics students had greater working memory capacity in the visuo-spatial domain only. Experiment 2 replicated this and demonstrated that neither visuo-spatial short-term memory nor endogenous spatial attention explained the visuo-spatial working memory differences. Experiment 3 used working memory span tasks with more traditional verbal or visuo-spatial processing elements to explore the effect of processing type. In this study mathematics students showed superior visuo-spatial working memory capacity only when the processing involved had a comparatively low level of central executive involvement. Both visuo-spatial working memory capacity and general visuo-spatial skills predicted mathematics achievement.