Cognitive, Affective, & Behavioral Neuroscience
Reward associations are known to shape the brain's processing of visual stimuli, but relatively l... more Reward associations are known to shape the brain's processing of visual stimuli, but relatively less is known about how reward associations impact the processing of auditory stimuli. We leveraged the high-temporal resolution of electroencephalography (EEG) and event-related potentials (ERPs) to investigate the influence of low-and high-magnitude stimulus-reward associations in an auditory oddball task. We associated fast, correct detection of certain auditory target stimuli with larger monetary rewards, and other auditory targets with smaller rewards. We found enhanced attentional processing of the more highly rewarded target stimuli, as evidenced by faster behavioral detection of those stimuli compared with lower-rewarded stimuli. Neurally, higher-reward associations enhanced the early sensory processing of auditory targets. Targets associated with higher-magnitude rewards had higher amplitude N1 and mismatch negativity (MMN) ERP components than targets associated with lower-magnitude rewards. Reward did not impact the latency of these early components. Higher-reward magnitude also decreased the latency and increased the amplitude of the longer-latency P3 component, suggesting that reward also can enhance the final processing stages of auditory target stimuli. These results provide insight into how the sensory and attentional neural processing of auditory stimuli is modulated by stimulus-reward associations and the magnitude of those associations, with higher-magnitude reward associations yielding enhanced auditory processing at both early and late stages compared with lower-magnitude reward associations.
While it is broadly accepted that attention modulates memory, the contribution of specific rapid ... more While it is broadly accepted that attention modulates memory, the contribution of specific rapid attentional processes to successful encoding is largely unknown. To investigate this issue, we leveraged the high temporal resolution of electroencephalographic recordings to directly link a cascade of visuo-attentional neural processes to successful encoding: namely (1) the N2pc (peaking ~200 ms), which reflects stimulus-specific attentional orienting and allocation, (2) the sustained posterior-contralateral negativity (post-N2pc), which has been associated with sustained visual processing, (3) the contralateral reduction in oscillatory alpha power (contralateral reduction in alpha > 200 ms), which has also been independently related to attentionally sustained visual processing. Each of these visuo-attentional processes was robustly predictive of successful encoding, and, moreover, each enhanced memory independently of the classic, longer-latency, conceptually related, difference-due...
and it is a condition of accessing publications that users recognise and abide by the legal requi... more and it is a condition of accessing publications that users recognise and abide by the legal requirements associated with these rights. • Users may download and print one copy of any publication from the public portal for the purpose of private study or research. • You may not further distribute the material or use it for any profit-making activity or commercial gain • You may freely distribute the URL identifying the publication in the public portal ? Take down policy If you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediately and investigate your claim.
Here we describe an EEG study investigating the interactions between multisensory (audio-visual) ... more Here we describe an EEG study investigating the interactions between multisensory (audio-visual) integration and spatial attention, using oscillatory gamma-band responses (GBRs). The results include a comparison with previously reported event-related potential (ERP) findings from the same paradigm. Unisensory-auditory (A), unisensory-visual (V), and multisensory (AV) stimuli were presented to the left and right hemispaces while subjects attended to a designated side to detect deviant target stimuli in either sensory modality. For attended multisensory stimuli we observed larger evoked GBRs approximately 40-50 ms post-stimulus over medial-frontal brain areas compared with those same multisensory stimuli when unattended. Further analysis indicated that the integration effect and its attentional enhancement may be caused in part by a stimulus-triggered phase resetting of ongoing gamma-band responses. Interestingly, no such early interaction effects (<90 ms) could be found in the ERP waveforms, suggesting that oscillatory GBRs may be more sensitive than ERPs to these early latency attention effects. Moreover, no GBR attention effects could be found for the unisensory auditory or unisensory visual stimuli, suggesting that attention particularly affects the integrative processing of audiovisual stimuli at these early latencies.
Attention can be involuntarily biased toward reward-associated distractors (value-driven attentio... more Attention can be involuntarily biased toward reward-associated distractors (value-driven attentional capture, VDAC). Yet past work has primarily demonstrated this distraction phenomenon during a particular set of circumstances: transient attentional orienting to potentially relevant stimuli occurring in our visual environment. Consequently, it is not well-understood if reward-based attentional capture can occur under other circumstances, such as during sustained visuospatial attention. Using EEG, we investigated whether associating transient distractors with reward value would increase their distractibility and lead to greater decrements in concurrent sustained spatial attention directed elsewhere. Human participants learned to associate three differently colored, laterally presented squares with rewards of varying magnitude (zero, small, and large). These colored squares were then periodically reintroduced as distractors at the same lateral locations during a demanding sustained-at...
Previous studies have indicated that both increased physical salience and increased reward-value ... more Previous studies have indicated that both increased physical salience and increased reward-value salience of a target improve behavioral measures of attentional selection. It is unclear, however, whether these two forms of salience interact with attentional networks through similar or different neural mechanisms, and what such differences might be. We examined this question by separately manipulating both the value-driven and physical salience of targets in a visual search task while recording response times (RTs) and event-related potentials, focusing on the attentional-orienting-sensitive N2pc event-related potential component. Human participants of both sexes searched arrays for targets of either a high-physical-salience color or one of two low-physicalsalience colors across three experimental phases. The first phase ("baseline") offered no rewards. RT and N2pc latencies were shorter for high-physical-salience targets, indicating faster attentional orienting. In the second phase ("equal-reward"), a low monetary reward was given for fast correct responses for all target types. This reward context improved overall performance, similarly shortening RTs and enhancing N2pc amplitudes for all target types, but with no change in N2pc latencies. In the third phase ("selective-reward"), the reward rate was made selectively higher for one of the two low-physical-salience colors, resulting in their RTs becoming as fast as the high-physical-salience targets. Despite the equally fast RTs, the N2pc's for these low-physical-salience, high-value targets remained later than for high-physical-salience targets, instead eliciting significantly larger N2pc's. These results suggest that enhanced physical salience leads to faster attentional orienting, but value-driven salience to stronger attentional orienting, underscoring the utilization of different underlying mechanisms.
Addiction to nicotine is extremely challenging to overcome, and the intense craving for the next ... more Addiction to nicotine is extremely challenging to overcome, and the intense craving for the next cigarette often leads to relapse in smokers who wish to quit. To dampen the urges of craving and inhibit unwanted behaviour, smokers must harness cognitive control, which is itself impaired in addiction. It is likely that craving may interact with cognitive control, and the present study sought to test the specificity of such interactions. To this end, data from 24 smokers were gathered using EEG and behavioural measures in a craving session (following a three‐hour nicotine abstention period) and a non‐craving session (having just smoked). In both sessions, participants performed a task probing various facets of cognitive control (response inhibition, task switching and conflict processing). Results showed that craving smokers were less flexible with the implementation of cognitive control, with demands of task switching and incongruency yielding greater deficits under conditions of crav...
The contents of working memory (WM) guide visual attention toward matching features, with visual ... more The contents of working memory (WM) guide visual attention toward matching features, with visual search being faster when the target and a feature of an item held in WM spatially overlap (validly cued) than when they occur at different locations (invalidly cued). Recent behavioral studies have indicated that attentional capture by WM content can be modulated by cognitive control: When WM cues are reliably helpful to visual search (predictably valid), capture is enhanced, but when reliably detrimental (predictably invalid), capture is attenuated. The neural mechanisms underlying this effect are not well understood, however. Here, we leveraged the high temporal resolution of ERPs time-locked to the onset of the search display to determine how and at what processing stage cognitive control modulates the search process. We manipulated predictability by grouping trials into unpredictable (50% valid/invalid) and predictable (100% valid, 100% invalid) blocks. Behavioral results confirmed t...
The Journal of neuroscience : the official journal of the Society for Neuroscience, 2015
Practice can improve performance on visual search tasks; the neural mechanisms underlying such im... more Practice can improve performance on visual search tasks; the neural mechanisms underlying such improvements, however, are not clear. Response time typically shortens with practice, but which components of the stimulus-response processing chain facilitate this behavioral change? Improved search performance could result from enhancements in various cognitive processing stages, including (1) sensory processing, (2) attentional allocation, (3) target discrimination, (4) motor-response preparation, and/or (5) response execution. We measured event-related potentials (ERPs) as human participants completed a five-day visual-search protocol in which they reported the orientation of a color popout target within an array of ellipses. We assessed changes in behavioral performance and in ERP components associated with various stages of processing. After practice, response time decreased in all participants (while accuracy remained consistent), and electrophysiological measures revealed modulatio...
Cerebral cortex (New York, N.Y. : 1991), Jan 19, 2014
Adaptive choice behavior depends critically on identifying and learning from outcome-predicting c... more Adaptive choice behavior depends critically on identifying and learning from outcome-predicting cues. We hypothesized that attention may be preferentially directed toward certain outcome-predicting cues. We studied this possibility by analyzing event-related potential (ERP) responses in humans during a probabilistic decision-making task. Participants viewed pairs of outcome-predicting visual cues and then chose to wager either a small (i.e., loss-minimizing) or large (i.e., gain-maximizing) amount of money. The cues were bilaterally presented, which allowed us to extract the relative neural responses to each cue by using a contralateral-versus-ipsilateral ERP contrast. We found an early lateralized ERP response, whose features matched the attention-shift-related N2pc component and whose amplitude scaled with the learned reward-predicting value of the cues as predicted by an attention-for-reward model. Consistently, we found a double dissociation involving the N2pc. Across participan...
Recently, attempts have been made to disentangle the neural underpinnings of preparatory processe... more Recently, attempts have been made to disentangle the neural underpinnings of preparatory processes related to reward and attention. Functional magnetic resonance imaging (fMRI) research showed that neural activity related to the anticipation of reward and to attentional demands invokes neural activity patterns featuring large-scale overlap, along with some differences and interactions. Due to the limited temporal resolution of fMRI, however, the temporal dynamics of these processes remain unclear. Here, we report an event-related potentials (ERP) study in which cued attentional demands and reward prospect were combined in a factorial design. Results showed that reward prediction dominated early cue processing, as well as the early and later parts of the contingent negative variation (CNV) slow-wave ERP component that has been associated with task-preparation processes. Moreover these reward-related electrophysiological effects correlated across participants with response-time speeding on reward-prospect trials. In contrast, cued attentional demands affected only the later part of the CNV, with the highest amplitudes following cues predicting high-difficulty potential-reward targets, thus suggesting maximal task preparation when the task requires it and entails reward prospect. Consequently, we suggest that task-preparation processes triggered by reward can arise earlier, and potentially more directly, than strategic top-down aspects of preparation based on attentional demands.
Multisensory integration has often been characterized as an automatic process. Recent findings su... more Multisensory integration has often been characterized as an automatic process. Recent findings suggest that multisensory integration can occur across various stages of stimulus processing that are linked to, and can be modulated by, attention. Stimulus-driven, bottom-up mechanisms induced by cross-modal interactions can automatically capture attention towards multisensory events, particularly when competition to focus elsewhere is relatively low. Conversely, top-down attention can facilitate the integration of multisensory inputs and lead to a spread of attention across sensory modalities. These findings point to a more intimate and multifaceted interplay between attention and multisensory integration than was previously thought. We review developments in our understanding of the interactions between attention and multisensory processing, and propose a framework that unifies previous, apparently discordant findings. Bidirectional Influences of Multisensory Integration and Attention Our brains are continually inundated with stimulus input arriving through our various sensory pathways. The processes involved in synthesizing and organizing this multisensory deluge of inputs are fundamental to effective perception and cognitive functioning. Although the combination of information across the senses has been investigated since psychology became an experimental discipline [1], the last decade has seen a sharp increase of interest in this question. Within this context, the issue of how attentional shifts in one modality can affect orienting in other modalities has inspired a great deal of this recent research [2], but, perhaps surprisingly, the role that attention plays during multisensory integration itself has been largely overlooked. Multisensory integration generally refers to the set of processes by which information arriving from different sensory modalities (e.g., vision, audition, touch) interacts and
The integration of multisensory information has been shown to be guided by spatial and temporal p... more The integration of multisensory information has been shown to be guided by spatial and temporal proximity, as well as to be influenced by attention. Here we used neural measures of the multisensory spread of attention to investigate the spatial and temporal linking of synchronous versus near-synchronous auditory and visual events. Human participants attended selectively to one of two lateralized visual-stimulus streams while task-irrelevant tones were presented centrally. Electrophysiological measures of brain activity showed that tones occurring simultaneously or delayed by 100 ms were temporally linked to an attended visual stimulus, as reflected by robust cross-modal spreading-of-attention activity, but not when delayed by 300 ms. The neural data also indicated a ventriloquist-like spatial linking of the auditory to the attended visual stimuli, but only when occurring simultaneously. These neurophysiological results thus provide unique insight into the temporal and spatial princi...
Proceedings of the National Academy of Sciences, 2005
Attending to a stimulus is known to enhance the neural responses to that stimulus. Recent experim... more Attending to a stimulus is known to enhance the neural responses to that stimulus. Recent experiments on visual attention have shown that this modulation can have object-based characteristics, such that, when certain parts of a visual object are attended, other parts automatically also receive enhanced processing. Here, we investigated whether visual attention can modulate neural responses to other components of a multisensory object defined by synchronous, but spatially disparate, auditory and visual stimuli. The audiovisual integration of such multisensory stimuli typically leads to mislocalization of the sound toward the visual stimulus (ventriloquism illusion). Using event-related potentials and functional MRI, we found that the brain's response to task-irrelevant sounds occurring synchronously with a visual stimulus from a different location was larger when that accompanying visual stimulus was attended versus unattended. The event-related potential effect consisted of sust...
Associating stimuli with the prospect of reward typically facilitates responses to those stimuli ... more Associating stimuli with the prospect of reward typically facilitates responses to those stimuli due to an enhancement of attentional and cognitive-control processes. Such reward-induced facilitation might be especially helpful when cognitivecontrol mechanisms are challenged, as when one must overcome interference from irrelevant inputs. Here, we investigated the neural dynamics of reward effects in a color-naming Stroop task by employing event-related potentials (ERPs). We found that behavioral facilitation in potential-reward trials, as compared to no-reward trials, was paralleled by early ERP modulations likely indexing increased attention to the reward-predictive stimulus. Moreover, reward changed the temporal dynamics of conflict-related ERP components, which may be a consequence of an early access to the various stimulus features and their relationships. Finally, although word meanings referring to potential-reward colors were always taskirrelevant, they caused greater interference compared to words referring to no-reward colors, an effect that was accompanied by a relatively early fronto-central ERP modulation. This latter observation suggests that task-irrelevant reward information can undermine goal-directed behavior at an early processing stage, presumably reflecting priming of a goalincompatible response. Yet, these detrimental effects of incongruent reward-related words were absent in potential-reward trials, apparently due to the prioritized processing of task-relevant reward information. Taken together, the present data demonstrate that reward associations can influence conflict processing by changing the temporal dynamics of stimulus processing and subsequent cognitive-control mechanisms.
Recent brain imaging studies using functional magnetic resonance imaging (fMRI) have implicated a... more Recent brain imaging studies using functional magnetic resonance imaging (fMRI) have implicated a frontal-parietal network in the top-down control of attention. However, little is known about the timing and sequence of activations within this network. To investigate these timing questions, we used event-related electrical brain potentials (ERPs) and a specially designed visual-spatial attentional-cueing paradigm, which were applied as part of a multi-methodological approach that included a closely corresponding event-related fMRI study using an identical paradigm. In the first 400 ms post cue, attention-directing and control cues elicited similar general cue-processing activity, corresponding to the more lateral subregions of the frontal-parietal network identified with the fMRI. Following this, the attention-directing cues elicited a sustained negative-polarity brain wave that was absent for control cues. This activity could be linked to the more medial frontal-parietal subregions similarly identified in the fMRI as specifically involved in attentional orienting. Critically, both the scalp ERPs and the fMRI-seeded source modeling for this orienting-related activity indicated an earlier onset of frontal versus parietal contribution (;400 versus ;700 ms). This was then followed (;800-900 ms) by pretarget biasing activity in the region-specific visual-sensory occipital cortex. These results indicate an activation sequence of key components of the attentional-control brain network, providing insight into their functional roles. More specifically, these results suggest that voluntary attentional orienting is initiated by medial portions of frontal cortex, which then recruit medial parietal areas. Together, these areas then implement biasing of region-specific visual-sensory cortex to facilitate the processing of upcoming visual stimuli.
The synchronous occurrence of the unisensory components of a multisensory stimulus contributes to... more The synchronous occurrence of the unisensory components of a multisensory stimulus contributes to their successful merging into a coherent perceptual representation. Oscillatory gamma-band responses (GBRs, 30-80 Hz) have been linked to feature integration mechanisms and to multisensory processing, suggesting they may also be sensitive to the temporal alignment of multisensory stimulus components. Here we examined the effects on early oscillatory GBR brain activity of varying the precision of the temporal synchrony of the unisensory components of an audiovisual stimulus. Audiovisual stimuli were presented with stimulus onset asynchronies ranging from −125 to +125 ms. Randomized streams of auditory (A), visual (V), and audiovisual (AV) stimuli were presented centrally while subjects attended to either the auditory or visual modality to detect occasional targets. GBRs to auditory and visual components of multisensory AV stimuli were extracted for five subranges of asynchrony (e.g., A preceded by V by 100 ± 25 ms, by 50 ± 25 ms, etc.) and compared with GBRs to unisensory control stimuli. Robust multisensory interactions were observed in the early GBRs when the auditory and visual stimuli were presented with the closest synchrony. These effects were found over medial-frontal brain areas after 30-80 ms and over occipital brain areas after 60-120 ms. A second integration effect, possibly reflecting the perceptual separation of the two sensory inputs, was found over occipital areas when auditory inputs preceded visual by 100 ± 25 ms. No significant interactions were observed for the other subranges of asynchrony. These results show that the precision of temporal synchrony can have an impact on early cross-modal interactions in human cortex.
The extent of visual perceptual processing that occurs in the absence of awareness is as yet uncl... more The extent of visual perceptual processing that occurs in the absence of awareness is as yet unclear. Here we examined event-related-potential (ERP) indices of visual and cognitive processes as awareness was manipulated through object-substitution masking (OSM), an awarenessdisrupting effect that has been hypothesized to result from the disruption of reentrant signaling to low-level visual cortical areas. In OSM, a visual stimulus array is briefly presented that includes a parafoveal visual target denoted by a cue, typically consisting of several surrounding dots. When the offset of the target-surrounding cue dots are delayed relative to the rest of the array, a striking reduction in the perception of the target image surrounded by the dots is observed. Using faces and houses as the target stimuli, we found that successful OSM reduced or eliminated all the measured electrophysiological indices of visual processing stages after 130 ms post-stimulus. More specifically, when targets were missed within the masked condition (i.e., on trials with effective OSM that disrupted awareness), we observed fully intact early feed-forward processing up through the visual extrastriate P1 ERP component peaking at 100 ms, followed by reduced low-level activity over the occipital pole 130-170 ms post-stimulus, reduced ERP indices of lateralized shifts of attention toward the parafoveal target, reduced object-generic visual processing, abolished object-category-specific (face-specific) processing, and reduced late visual short-termmemory processing activity. The results provide a comprehensive electrophysiological account of the neurocognitive underpinnings of effective OSM of visual-object images, including evidence for central roles of early reentrant signal disruption and inadequate visual attentional deployment.
Humans are constantly confronted with environmental stimuli that conflict with task goals and can... more Humans are constantly confronted with environmental stimuli that conflict with task goals and can interfere with successful behavior. Prevailing theories propose the existence of cognitive control mechanisms that can suppress the processing of conflicting input and enhance that of the relevant input. However, the temporal cascade of brain processes invoked in response to conflicting stimuli remains poorly understood. By examining evoked electrical brain responses in a novel, hemifield-specific, visual-flanker task, we demonstrate that task-irrelevant conflicting stimulus input is quickly detected in higher level executive regions while simultaneously inducing rapid, recurrent modulation of sensory processing in the visual cortex. Importantly, however, both of these effects are larger for individuals with greater incongruency-related RT slowing. The combination of neural activation patterns and behavioral interference effects suggest that this initial sensory modulation induced by co...
Cognitive, Affective, & Behavioral Neuroscience
Reward associations are known to shape the brain's processing of visual stimuli, but relatively l... more Reward associations are known to shape the brain's processing of visual stimuli, but relatively less is known about how reward associations impact the processing of auditory stimuli. We leveraged the high-temporal resolution of electroencephalography (EEG) and event-related potentials (ERPs) to investigate the influence of low-and high-magnitude stimulus-reward associations in an auditory oddball task. We associated fast, correct detection of certain auditory target stimuli with larger monetary rewards, and other auditory targets with smaller rewards. We found enhanced attentional processing of the more highly rewarded target stimuli, as evidenced by faster behavioral detection of those stimuli compared with lower-rewarded stimuli. Neurally, higher-reward associations enhanced the early sensory processing of auditory targets. Targets associated with higher-magnitude rewards had higher amplitude N1 and mismatch negativity (MMN) ERP components than targets associated with lower-magnitude rewards. Reward did not impact the latency of these early components. Higher-reward magnitude also decreased the latency and increased the amplitude of the longer-latency P3 component, suggesting that reward also can enhance the final processing stages of auditory target stimuli. These results provide insight into how the sensory and attentional neural processing of auditory stimuli is modulated by stimulus-reward associations and the magnitude of those associations, with higher-magnitude reward associations yielding enhanced auditory processing at both early and late stages compared with lower-magnitude reward associations.
While it is broadly accepted that attention modulates memory, the contribution of specific rapid ... more While it is broadly accepted that attention modulates memory, the contribution of specific rapid attentional processes to successful encoding is largely unknown. To investigate this issue, we leveraged the high temporal resolution of electroencephalographic recordings to directly link a cascade of visuo-attentional neural processes to successful encoding: namely (1) the N2pc (peaking ~200 ms), which reflects stimulus-specific attentional orienting and allocation, (2) the sustained posterior-contralateral negativity (post-N2pc), which has been associated with sustained visual processing, (3) the contralateral reduction in oscillatory alpha power (contralateral reduction in alpha > 200 ms), which has also been independently related to attentionally sustained visual processing. Each of these visuo-attentional processes was robustly predictive of successful encoding, and, moreover, each enhanced memory independently of the classic, longer-latency, conceptually related, difference-due...
and it is a condition of accessing publications that users recognise and abide by the legal requi... more and it is a condition of accessing publications that users recognise and abide by the legal requirements associated with these rights. • Users may download and print one copy of any publication from the public portal for the purpose of private study or research. • You may not further distribute the material or use it for any profit-making activity or commercial gain • You may freely distribute the URL identifying the publication in the public portal ? Take down policy If you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediately and investigate your claim.
Here we describe an EEG study investigating the interactions between multisensory (audio-visual) ... more Here we describe an EEG study investigating the interactions between multisensory (audio-visual) integration and spatial attention, using oscillatory gamma-band responses (GBRs). The results include a comparison with previously reported event-related potential (ERP) findings from the same paradigm. Unisensory-auditory (A), unisensory-visual (V), and multisensory (AV) stimuli were presented to the left and right hemispaces while subjects attended to a designated side to detect deviant target stimuli in either sensory modality. For attended multisensory stimuli we observed larger evoked GBRs approximately 40-50 ms post-stimulus over medial-frontal brain areas compared with those same multisensory stimuli when unattended. Further analysis indicated that the integration effect and its attentional enhancement may be caused in part by a stimulus-triggered phase resetting of ongoing gamma-band responses. Interestingly, no such early interaction effects (<90 ms) could be found in the ERP waveforms, suggesting that oscillatory GBRs may be more sensitive than ERPs to these early latency attention effects. Moreover, no GBR attention effects could be found for the unisensory auditory or unisensory visual stimuli, suggesting that attention particularly affects the integrative processing of audiovisual stimuli at these early latencies.
Attention can be involuntarily biased toward reward-associated distractors (value-driven attentio... more Attention can be involuntarily biased toward reward-associated distractors (value-driven attentional capture, VDAC). Yet past work has primarily demonstrated this distraction phenomenon during a particular set of circumstances: transient attentional orienting to potentially relevant stimuli occurring in our visual environment. Consequently, it is not well-understood if reward-based attentional capture can occur under other circumstances, such as during sustained visuospatial attention. Using EEG, we investigated whether associating transient distractors with reward value would increase their distractibility and lead to greater decrements in concurrent sustained spatial attention directed elsewhere. Human participants learned to associate three differently colored, laterally presented squares with rewards of varying magnitude (zero, small, and large). These colored squares were then periodically reintroduced as distractors at the same lateral locations during a demanding sustained-at...
Previous studies have indicated that both increased physical salience and increased reward-value ... more Previous studies have indicated that both increased physical salience and increased reward-value salience of a target improve behavioral measures of attentional selection. It is unclear, however, whether these two forms of salience interact with attentional networks through similar or different neural mechanisms, and what such differences might be. We examined this question by separately manipulating both the value-driven and physical salience of targets in a visual search task while recording response times (RTs) and event-related potentials, focusing on the attentional-orienting-sensitive N2pc event-related potential component. Human participants of both sexes searched arrays for targets of either a high-physical-salience color or one of two low-physicalsalience colors across three experimental phases. The first phase ("baseline") offered no rewards. RT and N2pc latencies were shorter for high-physical-salience targets, indicating faster attentional orienting. In the second phase ("equal-reward"), a low monetary reward was given for fast correct responses for all target types. This reward context improved overall performance, similarly shortening RTs and enhancing N2pc amplitudes for all target types, but with no change in N2pc latencies. In the third phase ("selective-reward"), the reward rate was made selectively higher for one of the two low-physical-salience colors, resulting in their RTs becoming as fast as the high-physical-salience targets. Despite the equally fast RTs, the N2pc's for these low-physical-salience, high-value targets remained later than for high-physical-salience targets, instead eliciting significantly larger N2pc's. These results suggest that enhanced physical salience leads to faster attentional orienting, but value-driven salience to stronger attentional orienting, underscoring the utilization of different underlying mechanisms.
Addiction to nicotine is extremely challenging to overcome, and the intense craving for the next ... more Addiction to nicotine is extremely challenging to overcome, and the intense craving for the next cigarette often leads to relapse in smokers who wish to quit. To dampen the urges of craving and inhibit unwanted behaviour, smokers must harness cognitive control, which is itself impaired in addiction. It is likely that craving may interact with cognitive control, and the present study sought to test the specificity of such interactions. To this end, data from 24 smokers were gathered using EEG and behavioural measures in a craving session (following a three‐hour nicotine abstention period) and a non‐craving session (having just smoked). In both sessions, participants performed a task probing various facets of cognitive control (response inhibition, task switching and conflict processing). Results showed that craving smokers were less flexible with the implementation of cognitive control, with demands of task switching and incongruency yielding greater deficits under conditions of crav...
The contents of working memory (WM) guide visual attention toward matching features, with visual ... more The contents of working memory (WM) guide visual attention toward matching features, with visual search being faster when the target and a feature of an item held in WM spatially overlap (validly cued) than when they occur at different locations (invalidly cued). Recent behavioral studies have indicated that attentional capture by WM content can be modulated by cognitive control: When WM cues are reliably helpful to visual search (predictably valid), capture is enhanced, but when reliably detrimental (predictably invalid), capture is attenuated. The neural mechanisms underlying this effect are not well understood, however. Here, we leveraged the high temporal resolution of ERPs time-locked to the onset of the search display to determine how and at what processing stage cognitive control modulates the search process. We manipulated predictability by grouping trials into unpredictable (50% valid/invalid) and predictable (100% valid, 100% invalid) blocks. Behavioral results confirmed t...
The Journal of neuroscience : the official journal of the Society for Neuroscience, 2015
Practice can improve performance on visual search tasks; the neural mechanisms underlying such im... more Practice can improve performance on visual search tasks; the neural mechanisms underlying such improvements, however, are not clear. Response time typically shortens with practice, but which components of the stimulus-response processing chain facilitate this behavioral change? Improved search performance could result from enhancements in various cognitive processing stages, including (1) sensory processing, (2) attentional allocation, (3) target discrimination, (4) motor-response preparation, and/or (5) response execution. We measured event-related potentials (ERPs) as human participants completed a five-day visual-search protocol in which they reported the orientation of a color popout target within an array of ellipses. We assessed changes in behavioral performance and in ERP components associated with various stages of processing. After practice, response time decreased in all participants (while accuracy remained consistent), and electrophysiological measures revealed modulatio...
Cerebral cortex (New York, N.Y. : 1991), Jan 19, 2014
Adaptive choice behavior depends critically on identifying and learning from outcome-predicting c... more Adaptive choice behavior depends critically on identifying and learning from outcome-predicting cues. We hypothesized that attention may be preferentially directed toward certain outcome-predicting cues. We studied this possibility by analyzing event-related potential (ERP) responses in humans during a probabilistic decision-making task. Participants viewed pairs of outcome-predicting visual cues and then chose to wager either a small (i.e., loss-minimizing) or large (i.e., gain-maximizing) amount of money. The cues were bilaterally presented, which allowed us to extract the relative neural responses to each cue by using a contralateral-versus-ipsilateral ERP contrast. We found an early lateralized ERP response, whose features matched the attention-shift-related N2pc component and whose amplitude scaled with the learned reward-predicting value of the cues as predicted by an attention-for-reward model. Consistently, we found a double dissociation involving the N2pc. Across participan...
Recently, attempts have been made to disentangle the neural underpinnings of preparatory processe... more Recently, attempts have been made to disentangle the neural underpinnings of preparatory processes related to reward and attention. Functional magnetic resonance imaging (fMRI) research showed that neural activity related to the anticipation of reward and to attentional demands invokes neural activity patterns featuring large-scale overlap, along with some differences and interactions. Due to the limited temporal resolution of fMRI, however, the temporal dynamics of these processes remain unclear. Here, we report an event-related potentials (ERP) study in which cued attentional demands and reward prospect were combined in a factorial design. Results showed that reward prediction dominated early cue processing, as well as the early and later parts of the contingent negative variation (CNV) slow-wave ERP component that has been associated with task-preparation processes. Moreover these reward-related electrophysiological effects correlated across participants with response-time speeding on reward-prospect trials. In contrast, cued attentional demands affected only the later part of the CNV, with the highest amplitudes following cues predicting high-difficulty potential-reward targets, thus suggesting maximal task preparation when the task requires it and entails reward prospect. Consequently, we suggest that task-preparation processes triggered by reward can arise earlier, and potentially more directly, than strategic top-down aspects of preparation based on attentional demands.
Multisensory integration has often been characterized as an automatic process. Recent findings su... more Multisensory integration has often been characterized as an automatic process. Recent findings suggest that multisensory integration can occur across various stages of stimulus processing that are linked to, and can be modulated by, attention. Stimulus-driven, bottom-up mechanisms induced by cross-modal interactions can automatically capture attention towards multisensory events, particularly when competition to focus elsewhere is relatively low. Conversely, top-down attention can facilitate the integration of multisensory inputs and lead to a spread of attention across sensory modalities. These findings point to a more intimate and multifaceted interplay between attention and multisensory integration than was previously thought. We review developments in our understanding of the interactions between attention and multisensory processing, and propose a framework that unifies previous, apparently discordant findings. Bidirectional Influences of Multisensory Integration and Attention Our brains are continually inundated with stimulus input arriving through our various sensory pathways. The processes involved in synthesizing and organizing this multisensory deluge of inputs are fundamental to effective perception and cognitive functioning. Although the combination of information across the senses has been investigated since psychology became an experimental discipline [1], the last decade has seen a sharp increase of interest in this question. Within this context, the issue of how attentional shifts in one modality can affect orienting in other modalities has inspired a great deal of this recent research [2], but, perhaps surprisingly, the role that attention plays during multisensory integration itself has been largely overlooked. Multisensory integration generally refers to the set of processes by which information arriving from different sensory modalities (e.g., vision, audition, touch) interacts and
The integration of multisensory information has been shown to be guided by spatial and temporal p... more The integration of multisensory information has been shown to be guided by spatial and temporal proximity, as well as to be influenced by attention. Here we used neural measures of the multisensory spread of attention to investigate the spatial and temporal linking of synchronous versus near-synchronous auditory and visual events. Human participants attended selectively to one of two lateralized visual-stimulus streams while task-irrelevant tones were presented centrally. Electrophysiological measures of brain activity showed that tones occurring simultaneously or delayed by 100 ms were temporally linked to an attended visual stimulus, as reflected by robust cross-modal spreading-of-attention activity, but not when delayed by 300 ms. The neural data also indicated a ventriloquist-like spatial linking of the auditory to the attended visual stimuli, but only when occurring simultaneously. These neurophysiological results thus provide unique insight into the temporal and spatial princi...
Proceedings of the National Academy of Sciences, 2005
Attending to a stimulus is known to enhance the neural responses to that stimulus. Recent experim... more Attending to a stimulus is known to enhance the neural responses to that stimulus. Recent experiments on visual attention have shown that this modulation can have object-based characteristics, such that, when certain parts of a visual object are attended, other parts automatically also receive enhanced processing. Here, we investigated whether visual attention can modulate neural responses to other components of a multisensory object defined by synchronous, but spatially disparate, auditory and visual stimuli. The audiovisual integration of such multisensory stimuli typically leads to mislocalization of the sound toward the visual stimulus (ventriloquism illusion). Using event-related potentials and functional MRI, we found that the brain's response to task-irrelevant sounds occurring synchronously with a visual stimulus from a different location was larger when that accompanying visual stimulus was attended versus unattended. The event-related potential effect consisted of sust...
Associating stimuli with the prospect of reward typically facilitates responses to those stimuli ... more Associating stimuli with the prospect of reward typically facilitates responses to those stimuli due to an enhancement of attentional and cognitive-control processes. Such reward-induced facilitation might be especially helpful when cognitivecontrol mechanisms are challenged, as when one must overcome interference from irrelevant inputs. Here, we investigated the neural dynamics of reward effects in a color-naming Stroop task by employing event-related potentials (ERPs). We found that behavioral facilitation in potential-reward trials, as compared to no-reward trials, was paralleled by early ERP modulations likely indexing increased attention to the reward-predictive stimulus. Moreover, reward changed the temporal dynamics of conflict-related ERP components, which may be a consequence of an early access to the various stimulus features and their relationships. Finally, although word meanings referring to potential-reward colors were always taskirrelevant, they caused greater interference compared to words referring to no-reward colors, an effect that was accompanied by a relatively early fronto-central ERP modulation. This latter observation suggests that task-irrelevant reward information can undermine goal-directed behavior at an early processing stage, presumably reflecting priming of a goalincompatible response. Yet, these detrimental effects of incongruent reward-related words were absent in potential-reward trials, apparently due to the prioritized processing of task-relevant reward information. Taken together, the present data demonstrate that reward associations can influence conflict processing by changing the temporal dynamics of stimulus processing and subsequent cognitive-control mechanisms.
Recent brain imaging studies using functional magnetic resonance imaging (fMRI) have implicated a... more Recent brain imaging studies using functional magnetic resonance imaging (fMRI) have implicated a frontal-parietal network in the top-down control of attention. However, little is known about the timing and sequence of activations within this network. To investigate these timing questions, we used event-related electrical brain potentials (ERPs) and a specially designed visual-spatial attentional-cueing paradigm, which were applied as part of a multi-methodological approach that included a closely corresponding event-related fMRI study using an identical paradigm. In the first 400 ms post cue, attention-directing and control cues elicited similar general cue-processing activity, corresponding to the more lateral subregions of the frontal-parietal network identified with the fMRI. Following this, the attention-directing cues elicited a sustained negative-polarity brain wave that was absent for control cues. This activity could be linked to the more medial frontal-parietal subregions similarly identified in the fMRI as specifically involved in attentional orienting. Critically, both the scalp ERPs and the fMRI-seeded source modeling for this orienting-related activity indicated an earlier onset of frontal versus parietal contribution (;400 versus ;700 ms). This was then followed (;800-900 ms) by pretarget biasing activity in the region-specific visual-sensory occipital cortex. These results indicate an activation sequence of key components of the attentional-control brain network, providing insight into their functional roles. More specifically, these results suggest that voluntary attentional orienting is initiated by medial portions of frontal cortex, which then recruit medial parietal areas. Together, these areas then implement biasing of region-specific visual-sensory cortex to facilitate the processing of upcoming visual stimuli.
The synchronous occurrence of the unisensory components of a multisensory stimulus contributes to... more The synchronous occurrence of the unisensory components of a multisensory stimulus contributes to their successful merging into a coherent perceptual representation. Oscillatory gamma-band responses (GBRs, 30-80 Hz) have been linked to feature integration mechanisms and to multisensory processing, suggesting they may also be sensitive to the temporal alignment of multisensory stimulus components. Here we examined the effects on early oscillatory GBR brain activity of varying the precision of the temporal synchrony of the unisensory components of an audiovisual stimulus. Audiovisual stimuli were presented with stimulus onset asynchronies ranging from −125 to +125 ms. Randomized streams of auditory (A), visual (V), and audiovisual (AV) stimuli were presented centrally while subjects attended to either the auditory or visual modality to detect occasional targets. GBRs to auditory and visual components of multisensory AV stimuli were extracted for five subranges of asynchrony (e.g., A preceded by V by 100 ± 25 ms, by 50 ± 25 ms, etc.) and compared with GBRs to unisensory control stimuli. Robust multisensory interactions were observed in the early GBRs when the auditory and visual stimuli were presented with the closest synchrony. These effects were found over medial-frontal brain areas after 30-80 ms and over occipital brain areas after 60-120 ms. A second integration effect, possibly reflecting the perceptual separation of the two sensory inputs, was found over occipital areas when auditory inputs preceded visual by 100 ± 25 ms. No significant interactions were observed for the other subranges of asynchrony. These results show that the precision of temporal synchrony can have an impact on early cross-modal interactions in human cortex.
The extent of visual perceptual processing that occurs in the absence of awareness is as yet uncl... more The extent of visual perceptual processing that occurs in the absence of awareness is as yet unclear. Here we examined event-related-potential (ERP) indices of visual and cognitive processes as awareness was manipulated through object-substitution masking (OSM), an awarenessdisrupting effect that has been hypothesized to result from the disruption of reentrant signaling to low-level visual cortical areas. In OSM, a visual stimulus array is briefly presented that includes a parafoveal visual target denoted by a cue, typically consisting of several surrounding dots. When the offset of the target-surrounding cue dots are delayed relative to the rest of the array, a striking reduction in the perception of the target image surrounded by the dots is observed. Using faces and houses as the target stimuli, we found that successful OSM reduced or eliminated all the measured electrophysiological indices of visual processing stages after 130 ms post-stimulus. More specifically, when targets were missed within the masked condition (i.e., on trials with effective OSM that disrupted awareness), we observed fully intact early feed-forward processing up through the visual extrastriate P1 ERP component peaking at 100 ms, followed by reduced low-level activity over the occipital pole 130-170 ms post-stimulus, reduced ERP indices of lateralized shifts of attention toward the parafoveal target, reduced object-generic visual processing, abolished object-category-specific (face-specific) processing, and reduced late visual short-termmemory processing activity. The results provide a comprehensive electrophysiological account of the neurocognitive underpinnings of effective OSM of visual-object images, including evidence for central roles of early reentrant signal disruption and inadequate visual attentional deployment.
Humans are constantly confronted with environmental stimuli that conflict with task goals and can... more Humans are constantly confronted with environmental stimuli that conflict with task goals and can interfere with successful behavior. Prevailing theories propose the existence of cognitive control mechanisms that can suppress the processing of conflicting input and enhance that of the relevant input. However, the temporal cascade of brain processes invoked in response to conflicting stimuli remains poorly understood. By examining evoked electrical brain responses in a novel, hemifield-specific, visual-flanker task, we demonstrate that task-irrelevant conflicting stimulus input is quickly detected in higher level executive regions while simultaneously inducing rapid, recurrent modulation of sensory processing in the visual cortex. Importantly, however, both of these effects are larger for individuals with greater incongruency-related RT slowing. The combination of neural activation patterns and behavioral interference effects suggest that this initial sensory modulation induced by co...
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Papers by Marty Woldorff