Signed Reward Prediction Errors in the Ventral Striatum Drive Episodic Memory.

Recent behavioral evidence implicates reward prediction errors (RPEs) as a key factor in the acquisition of episodic memory. Yet, important neural predictions related to the role of RPEs in episodic memory acquisition remain to be tested. Humans (both sexes) performed a novel variable-choice task where we experimentally manipulated RPEs and found support for key neural predictions with fMRI. Our results show that in line with previous behavioral observations, episodic memory accuracy increases with the magnitude of signed (i.e., better/worse-than-expected) RPEs (SRPEs). Neurally, we observe that SRPEs are encoded in the ventral striatum (VS). Crucially, we demonstrate through mediation analysis that activation in the VS mediates the experimental manipulation of SRPEs on episodic memory accuracy. In particular, SRPE-based responses in the VS (during learning) predict the strength of subsequent episodic memory (during recollection). Furthermore, functional connectivity between task-relevant processing areas (i.e., face-selective areas) and hippocampus and ventral striatum increased as a function of RPE value (during learning), suggesting a central role of these areas in episodic memory formation. Our results consolidate reinforcement learning theory and striatal RPEs as key factors subtending the formation of episodic memory.SIGNIFICANCE STATEMENT Recent behavioral research has shown that reward prediction errors (RPEs), a key concept of reinforcement learning theory, are crucial to the formation of episodic memories. In this study, we reveal the neural underpinnings of this process. Using fMRI, we show that signed RPEs (SRPEs) are encoded in the ventral striatum (VS), and crucially, that SRPE VS activity is responsible for the subsequent recollection accuracy of one-shot learned episodic memory associations.

Reward does not modulate corticospinal excitability in anticipation of a Stroop trial.

Action preparation is associated with a transient decrease of corticospinal excitability just before target onset. We have previously shown that the prospect of reward modulates preparatory corticospinal excitability in a Simon task. While the conflict in the Simon task strongly implicates the motor system, it is unknown whether reward prospect modulates preparatory corticospinal excitability in tasks that implicate the motor system less directly. To that effect, we examined reward-modulated preparatory corticospinal excitability in the Stroop task. We administered a rewarded cue-target delay paradigm using Stroop stimuli that afforded a left or right index finger response. Single-pulse transcranial magnetic stimulation was administered over the left primary motor cortex and electromyography was obtained from the right first dorsal interosseous muscle. In line with previous findings, there was a preparatory decrease in corticospinal excitability during the delay period. In contrast to our previous study using the Simon task, preparatory corticospinal excitability was not modulated by reward. Our results indicate that reward-modulated changes in the motor system depend on specific task-demands, possibly related to varying degrees of motor conflict.

Dissociating Reward- and Attention-driven Biasing of Global Feature-based Selection in Human Visual Cortex.

Objects that promise rewards are prioritized for visual selection. The way this prioritization shapes sensory processing in visual cortex, however, is debated. It has been suggested that rewards motivate stronger attentional focusing, resulting in a modulation of sensory selection in early visual cortex. An open question is whether those reward-driven modulations would be independent of similar modulations indexing the selection of attended features that are not associated with reward. Here, we use magnetoencephalography in human observers to investigate whether the modulations indexing global color-based selection in visual cortex are separable for target- and (monetary) reward-defining colors. To assess the underlying global color-based activity modulation, we compare the event-related magnetic field response elicited by a color probe in the unattended hemifield drawn either in the target color, the reward color, both colors, or a neutral task-irrelevant color. To test whether target and reward relevance trigger separable modulations, we manipulate attention demands on target selection while keeping reward-defining experimental parameters constant. Replicating previous observations, we find that reward and target relevance produce almost indistinguishable gain modulations in ventral extratriate cortex contralateral to the unattended color probe. Importantly, increasing attention demands on target discrimination increases the response to the target-defining color, whereas the response to the rewarded color remains largely unchanged. These observations indicate that, although task relevance and reward influence the very same feature-selective area in extrastriate visual cortex, the associated modulations are largely independent.

Cortical and Subcortical Coordination of Visual Spatial Attention Revealed by Simultaneous EEG-fMRI Recording.

Visual spatial attention has been studied in humans with both electroencephalography (EEG) and functional magnetic resonance imaging (fMRI) individually. However, due to the intrinsic limitations of each of these methods used alone, our understanding of the systems-level mechanisms underlying attentional control remains limited. Here, we examined trial-to-trial covariations of concurrently recorded EEG and fMRI in a cued visual spatial attention task in humans, which allowed delineation of both the generators and modulators of the cue-triggered event-related oscillatory brain activity underlying attentional control function. The fMRI activity in visual cortical regions contralateral to the cued direction of attention covaried positively with occipital gamma-band EEG, consistent with activation of cortical regions representing attended locations in space. In contrast, fMRI activity in ipsilateral visual cortical regions covaried inversely with occipital alpha-band oscillations, consistent with attention-related suppression of the irrelevant hemispace. Moreover, the pulvinar nucleus of the thalamus covaried with both of these spatially specific, attention-related, oscillatory EEG modulations. Because the pulvinar's neuroanatomical geometry makes it unlikely to be a direct generator of the scalp-recorded EEG, these covariational patterns appear to reflect the pulvinar's role as a regulatory control structure, sending spatially specific signals to modulate visual cortex excitability proactively. Together, these combined EEG/fMRI results illuminate the dynamically interacting cortical and subcortical processes underlying spatial attention, providing important insight not realizable using either method alone.SIGNIFICANCE STATEMENT Noninvasive recordings of changes in the brain's blood flow using functional magnetic resonance imaging and electrical activity using electroencephalography in humans have individually shown that shifting attention to a location in space produces spatially specific changes in visual cortex activity in anticipation of a stimulus. The mechanisms controlling these attention-related modulations of sensory cortex, however, are poorly understood. Here, we recorded these two complementary measures of brain activity simultaneously and examined their trial-to-trial covariations to gain insight into these attentional control mechanisms. This multi-methodological approach revealed the attention-related coordination of visual cortex modulation by the subcortical pulvinar nucleus of the thalamus while also disentangling the mechanisms underlying the attentional enhancement of relevant stimulus input and those underlying the concurrent suppression of irrelevant input.

Neural conflict-control mechanisms improve memory for target stimuli.

According to conflict-monitoring models, conflict serves as an internal signal for reinforcing top-down attention to task-relevant information. While evidence based on measures of ongoing task performance supports this idea, implications for long-term consequences, that is, memory, have not been tested yet. Here, we evaluated the prediction that conflict-triggered attentional enhancement of target-stimulus processing should be associated with superior subsequent memory for those stimuli. By combining functional magnetic resonance imaging (fMRI) with a novel variant of a face-word Stroop task that employed trial-unique face stimuli as targets, we were able to assess subsequent (incidental) memory for target faces as a function of whether a given face had previously been accompanied by congruent, neutral, or incongruent (conflicting) distracters. In line with our predictions, incongruent distracters not only induced behavioral conflict, but also gave rise to enhanced memory for target faces. Moreover, conflict-triggered neural activity in prefrontal and parietal regions was predictive of subsequent retrieval success, and displayed conflict-enhanced functional coupling with medial-temporal lobe regions. These data provide support for the proposal that conflict evokes enhanced top-down attention to task-relevant stimuli, thereby promoting their encoding into long-term memory. Our findings thus delineate the neural mechanisms of a novel link between cognitive control and memory.

Determinants of Global Color-Based Selection in Human Visual Cortex.

Feature attention operates in a spatially global way, with attended feature values being prioritized for selection outside the focus of attention. Accounts of global feature attention have emphasized feature competition as a determining factor. Here, we use magnetoencephalographic recordings in humans to test whether competition is critical for global feature selection to arise. Subjects performed a color/shape discrimination task in one visual field (VF), while irrelevant color probes were presented in the other unattended VF. Global effects of color attention were assessed by analyzing the response to the probe as a function of whether or not the probe's color was a target-defining color. We find that global color selection involves a sequence of modulations in extrastriate cortex, with an initial phase in higher tier areas (lateral occipital complex) followed by a later phase in lower tier retinotopic areas (V3/V4). Importantly, these modulations appeared with and without color competition in the focus of attention. Moreover, early parts of the modulation emerged for a task-relevant color not even present in the focus of attention. All modulations, however, were eliminated during simple onset-detection of the colored target. These results indicate that global color-based attention depends on target discrimination independent of feature competition in the focus of attention.

The role of the striatum in effort-based decision-making in the absence of reward.

Decision-making involves weighing costs against benefits, for instance, in terms of the effort it takes to obtain a reward of a given magnitude. This evaluation process has been linked to the dorsal anterior cingulate cortex (dACC) and the striatum, with activation in these brain structures reflecting the discounting effect of effort on reward. Here, we investigate how cognitive effort influences neural choice processes in the absence of an extrinsic reward. Using functional magnetic resonance imaging in humans, we used an effort-based decision-making task in which participants were required to choose between two options for a subsequent flanker task that differed in the amount of cognitive effort. Cognitive effort was manipulated by varying the proportion of incongruent trials associated with each choice option. Choice-locked activation in the striatum was higher when participants chose voluntarily for the more effortful alternative but displayed the opposite trend on forced-choice trials. The dACC revealed a similar, yet only trend-level significant, activation pattern. Our results imply that activation levels in the striatum reflect a cost-benefit analysis, in which a balance is made between effort discounting and the intrinsic motivation to choose a cognitively challenging task. Moreover, our findings indicate that it matters whether this challenge is voluntarily chosen or externally imposed. As such, the present findings contrast with classical findings on effort discounting that found reductions in striatum activation for higher effort by finding enhancements of the same neural circuits when a cognitively challenging task is voluntarily selected and does not entail the danger of losing reward.

Reward- and attention-related biasing of sensory selection in visual cortex.

Attention to task-relevant features leads to a biasing of sensory selection in extrastriate cortex. Features signaling reward seem to produce a similar bias, but how modulatory effects due to reward and attention relate to each other is largely unexplored. To address this issue, it is critical to separate top-down settings defining reward relevance from those defining attention. To this end, we used a visual search paradigm in which the target's definition (attention to color) was dissociated from reward relevance by delivering monetary reward on search frames where a certain task-irrelevant color was combined with the target-defining color to form the target object. We assessed the state of neural biasing for the attended and reward-relevant color by analyzing the neuromagnetic brain response to asynchronously presented irrelevant distractor probes drawn in the target-defining color, the reward-relevant color, and a completely irrelevant color as a reference. We observed that for the prospect of moderate rewards, the target-defining color but not the reward-relevant color produced a selective enhancement of the neuromagnetic response between 180 and 280 msec in ventral extrastriate visual cortex. Increasing reward prospect caused a delayed attenuation (220-250 msec) of the response to reward probes, which followed a prior (160-180 msec) response enhancement in dorsal ACC. Notably, shorter latency responses in dorsal ACC were associated with stronger attenuation in extrastriate visual cortex. Finally, an analysis of the brain response to the search frames revealed that the presence of the reward-relevant color in search distractors elicited an enhanced response that was abolished after increasing reward size. The present data together indicate that when top-down definitions of reward relevance and attention are separated, the behavioral significance of reward-associated features is still rapidly coded in higher-level cortex areas, thereby commanding effective top-down inhibitory control to counter a selection bias for those features in extrastriate visual cortex.

Response inhibition and its relation to multidimensional impulsivity.

Impulsivity is a multidimensional construct that has been suggested as a vulnerability factor for several psychiatric disorders, especially addiction disorders. Poor response inhibition may constitute one facet of impulsivity. Trait impulsivity can be assessed by self-report questionnaires such as the widely used Barratt Impulsiveness Scale (BIS-11). However, regarding the multidimensionality of impulsivity different concepts have been proposed, in particular the UPPS self-report questionnaire ('Urgency', 'Lack of Premeditation', 'Lack of Perseverance', 'Sensation Seeking') that is based on a factor analytic approach. The question as to which aspects of trait impulsivity map on individual differences of the behavioral and neural correlates of response inhibition so far remains unclear. In the present study, we investigated 52 healthy individuals that scored either very high or low on the BIS-11 and underwent a reward-modulated Stop-signal task during fMRI. Neither behavioral nor neural differences were observed with respect to high- and low-BIS groups. In contrast, UPPS subdomain Urgency best explained inter-individual variability in SSRT scores and was further negatively correlated to right IFG/aI activation in 'Stop>Go' trials - a key region for response inhibition. Successful response inhibition in rewarded compared to nonrewarded stop trials yielded ventral striatal (VS) activation which might represent a feedback signal. Interestingly, only participants with low Urgency scores were able to use this VS feedback signal for better response inhibition. Our findings indicate that the relationship of impulsivity and response inhibition has to be treated carefully. We propose Urgency as an important subdomain that might be linked to response inhibition as well as to the use of reward-based neural signals. Based on the present results, further studies examining the influence of impulsivity on psychiatric disorders should take into account Urgency as an important modulator of behavioral adaptation.

Reward prospect rapidly speeds up response inhibition via reactive control.

Response inhibition is an important cognitive-control function that allows for already-initiated or habitual behavioral responses to be promptly withheld when needed. A typical paradigm to study this function is the stop-signal task. From this task, the stop-signal response time (SSRT) can be derived, which indexes how rapidly an already-initiated response can be canceled. Typically, SSRTs range around 200 ms, identifying response inhibition as a particularly rapid cognitive-control process. Even so, it has recently been shown that SSRTs can be further accelerated if successful response inhibition is rewarded. Since this earlier study effectively ruled out differential preparatory (proactive) control adjustments, the reward benefits likely relied on boosted reactive control. Yet, given how rapidly such control processes would need to be enhanced, alternative explanations circumventing reactive control are important to consider. We addressed this question with an fMRI study by gauging the overlap of the brain networks associated with reward-related and response-inhibition-related processes in a reward-modulated stop-signal task. In line with the view that reactive control can indeed be boosted swiftly by reward availability, we found that the activity in key brain areas related to response inhibition was enhanced for reward-related stop trials. Furthermore, we observed that this beneficial reward effect was triggered by enhanced connectivity between task-unspecific (reward-related) and task-specific (inhibition-related) areas in the medial prefrontal cortex (mPFC). The present data hence suggest that reward information can be translated very rapidly into behavioral benefits (here, within ~200 ms) through enhanced reactive control, underscoring the immediate responsiveness of such control processes to reward availability in general.

Separable mechanisms underlying global feature-based attention.

Feature-based attention is known to operate in a spatially global manner, in that the selection of attended features is not bound to the spatial focus of attention. Here we used electromagnetic recordings in human observers to characterize the spatiotemporal signature of such global selection of an orientation feature. Observers performed a simple orientation-discrimination task while ignoring task-irrelevant orientation probes outside the focus of attention. We observed that global feature-based selection, indexed by the brain response to unattended orientation probes, is composed of separable functional components. One such component reflects global selection based on the similarity of the probe with task-relevant orientation values ("template matching"), which is followed by a component reflecting selection based on the similarity of the probe with the orientation value under discrimination in the focus of attention ("discrimination matching"). Importantly, template matching occurs at ∼150 ms after stimulus onset, ∼80 ms before the onset of discrimination matching. Moreover, source activity underlying template matching and discrimination matching was found to originate from ventral extrastriate cortex, with the former being generated in more anterolateral and the latter in more posteromedial parts, suggesting template matching to occur in visual cortex higher up in the visual processing hierarchy than discrimination matching. We take these observations to indicate that the population-level signature of global feature-based selection reflects a sequence of hierarchically ordered operations in extrastriate visual cortex, in which the selection based on task relevance has temporal priority over the selection based on the sensory similarity between input representations.

Picture novelty attenuates semantic interference and modulates concomitant neural activity in the anterior cingulate cortex and the locus coeruleus.

Goal-directed behavior requires the ability to focus on information that is relevant to a given task and to ignore information that might interfere with it. In the Stroop task, for example, the influence of an irrelevant word needs to be overcome, which is believed to be difficult because it arises in a fast and automatic fashion, which effectively renders it very salient. Here we address the question of whether this can be counteracted by increasing the saliency of the task-relevant input, for example by modulating its relative novelty, which increases saliency in a fairly implicit and controlled fashion. To test the influence of novelty on interference processing, we employed a picture-word interference task in the fMRI scanner, in which we manipulated the novelty of the task-relevant picture. We found that picture novelty indeed reduced typical behavioral interference from incongruent words. Moreover, familiar incongruent trials were associated with activity increases in the anterior cingulate cortex (ACC), a prime conflict-processing region, as well as in the noradrenergic locus coeruleus (LC), which entertains connections both to and from the ACC. The lack of analogous activations in novel incongruent trials suggests that the reduction of behavioral interference was not related to enhanced conflict-resolution processes, but rather to the automatic prioritization of novel pictures which appears to avert the influence of irrelevant words at the front end. Interestingly, activity in the ACC and LC was slightly stronger in novel congruent trials compared to incongruent ones, which may reflect increased relevance of novel stimuli when encoded in a congruent context. In summary, the present data demonstrate that stimulus novelty clearly reduces semantic interference, and highlights a complex interaction of interference and novelty processing on the neural level, including an involvement of the noradrenergic system in the processing of cognitively and perceptually salient events.

The role of the pulvinar in distractor processing and visual search.

The pulvinar nuclei of the thalamus are hypothesized to coordinate attentional selection in the visual cortex. Different models have, however, been proposed for the precise role of the pulvinar in attention. One proposal is that the pulvinar mediates shifts of spatial attention; a different proposal is that it serves the filtering of distractor information. At present, the relation between these possible operations and their relative importance in the pulvinar remains unresolved. We address this issue by contrasting these proposals in two fMRI experiments. We used a visual search paradigm that permitted us to dissociate neural activity reflecting shifts of attention from activity underlying distractor filtering. We find that distractor filtering, but not the operation of shifting attention, is associated with strong activity enhancements in dorsal and ventral regions of the pulvinar as well as in early visual cortex areas including the primary visual cortex. Our observations indicate that distractor filtering is the preponderant attentional operation subserved by the pulvinar, presumably mediated by a modulation of processing in visual areas where spatial resolution is sufficiently high to separate target from distractor input.

The involvement of the dopaminergic midbrain and cortico-striatal-thalamic circuits in the integration of reward prospect and attentional task demands.

Reward has been shown to promote human performance in multiple task domains. However, an important debate has developed about the uniqueness of reward-related neural signatures associated with such facilitation, as similar neural patterns can be triggered by increased attentional focus independent of reward. Here, we used functional magnetic resonance imaging to directly investigate the neural commonalities and interactions between the anticipation of both reward and task difficulty, by independently manipulating these factors in a cued-attention paradigm. In preparation for the target stimulus, both factors increased activity within the midbrain, dorsal striatum, and fronto-parietal areas, while inducing deactivations in default-mode regions. Additionally, reward engaged the ventral striatum, posterior cingulate, and occipital cortex, while difficulty engaged medial and dorsolateral frontal regions. Importantly, a network comprising the midbrain, caudate nucleus, thalamus, and anterior midcingulate cortex exhibited an interaction between reward and difficulty, presumably reflecting additional resource recruitment for demanding tasks with profitable outcome. This notion was consistent with a negative correlation between cue-related midbrain activity and difficulty-induced performance detriments in reward-predictive trials. Together, the data demonstrate that expected value and attentional demands are integrated in cortico-striatal-thalamic circuits in coordination with the dopaminergic midbrain to flexibly modulate resource allocation for an effective pursuit of behavioral goals.

The neural underpinnings of how reward associations can both guide and misguide attention.

It is commonly accepted that reward is an effective motivator of behavior, but little is known about potential costs resulting from reward associations. Here, we used functional magnetic resonance imaging (fMRI) to investigate the neural underpinnings of such reward-related performance-disrupting effects in a reward-modulated Stroop task in humans. While reward associations in the task-relevant dimension (i.e., ink color) facilitated performance, behavioral detriments were found when the task-irrelevant dimension (i.e., word meaning) implicitly referred to reward-predictive ink colors. Neurally, only relevant reward associations invoked a typical reward-anticipation response in the nucleus accumbens (NAcc), which was in turn predictive of behavioral facilitation. In contrast, irrelevant reward associations increased activity in a medial prefrontal motor-control-related region, namely the presupplementary motor area (pre-SMA), which likely reflects the preemption and inhibition of automatic response tendencies that are amplified by irrelevant reward-related words. This view was further supported by a positive relationship between pre-SMA activity and pronounced response slowing in trials containing reward-related as compared with reward-unrelated incongruent words. Importantly, the distinct neural processes related to the beneficial and detrimental behavioral effects of reward associations appeared to arise from preferential-coding mechanisms in visual-processing areas that were shared by the two stimulus dimensions, suggesting a transfer of reward-related saliency to the irrelevant dimension, but with highly differential behavioral and neural ramifications. More generally, the data demonstrate that even entirely irrelevant reward associations can influence stimulus-processing and response-selection pathways relatively automatically, thereby representing an important flipside of reward-driven performance enhancements.

Neural mechanisms of surround attenuation and distractor competition in visual search.

Visual attention biases relevant processing in the visual system by amplifying relevant or attenuating irrelevant sensory input. A potential signature of the latter operation, referred to as surround attenuation, has recently been identified in the electromagnetic brain response of human observers performing visual search. It was found that a zone of attenuated cortical excitability surrounds the target when the search required increased spatial resolution for item discrimination. Here we address the obvious hypothesis that surround attenuation serves distractor suppression in the vicinity of the target where interference from irrelevant search items is maximal. To test this hypothesis, surround attenuation was assessed under conditions when the target was presented in isolation versus when it was surrounded by distractors. Surprisingly, substantial and indistinguishable surround attenuation was seen under both conditions, indicating that it reflects an attentional operation independent of the presence of distractors. Adding distractors in the target's surround, however, increased the amplitude of the N2pc--an evoked response known to index distractor competition in visual search. Moreover, adding distractors led to a topographical change of source activity underlying the N2pc toward earlier extrastriate areas. In contrast, the topography of reduced source activity due to surround attenuation remained unaltered with and without distractors in the target's surround. We conclude that surround attenuation is not a direct consequence of the attenuation of distractors in visual search and that it dissociates from attentional operations reflected by the N2pc. A theoretical framework is proposed that links both operations in a common model of top-down attentional selection in visual cortex.

Task-load-dependent activation of dopaminergic midbrain areas in the absence of reward.

Dopamine release in cortical and subcortical structures plays a central role in reward-related neural processes. Within this context, dopaminergic inputs are commonly assumed to play an activating role, facilitating behavioral and cognitive operations necessary to obtain a prospective reward. Here, we provide evidence from human fMRI that this activating role can also be mediated by task-demand-related processes and thus extends beyond situations that only entail extrinsic motivating factors. Using a visual discrimination task in which varying levels of task demands were precued, we found enhanced hemodynamic activity in the substantia nigra (SN) for high task demands in the absence of reward or similar extrinsic motivating factors. This observation thus indicates that the SN can also be activated in an endogenous fashion. In parallel to its role in reward-related processes, reward-independent activation likely serves to recruit the processing resources needed to meet enhanced task demands. Simultaneously, activity in a wide network of cortical and subcortical control regions was enhanced in response to high task demands, whereas areas of the default-mode network were deactivated more strongly. The present observations suggest that the SN represents a core node within a broader neural network that adjusts the amount of available neural and behavioral resources to changing situational opportunities and task requirements, which is often driven by extrinsic factors but can also be controlled endogenously.

Strategic allocation of attention reduces temporally predictable stimulus conflict.

Humans are able to continuously monitor environmental situations and adjust their behavioral strategies to optimize performance. Here we investigate the behavioral and brain adjustments that occur when conflicting stimulus elements are, or are not, temporally predictable. ERPs were collected while manual response variants of the Stroop task were performed in which the SOAs between the relevant color and irrelevant word stimulus components were either randomly intermixed or held constant within each experimental run. Results indicated that the size of both the neural and behavioral effects of stimulus incongruency varied with the temporal arrangement of the stimulus components, such that the random-SOA arrangements produced the greatest incongruency effects at the earliest irrelevant first SOA (-200 msec) and the constant-SOA arrangements produced the greatest effects with simultaneous presentation. These differences in conflict processing were accompanied by rapid (∼150 msec) modulations of the sensory ERPs to the irrelevant distractor components when they occurred consistently first. These effects suggest that individuals are able to strategically allocate attention in time to mitigate the influence of a temporally predictable distractor. As these adjustments are instantiated by the participants without instruction, they reveal a form of rapid strategic learning for dealing with temporally predictable stimulus incongruency.

Rapid modulation of sensory processing induced by stimulus conflict.

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 conflicting stimulus inputs reflects performance-degrading attentional distraction because of their incompatibility rather than any rapid task-enhancing cognitive control mechanisms. The present findings thus provide neural evidence for a model in which attentional distraction is the key initial trigger for the temporal cascade of processes by which the human brain responds to conflicting stimulus input in the environment.

Object-based selection of irrelevant features is not confined to the attended object.

Attention to one feature of an object can bias the processing of unattended features of that object. Here we demonstrate with ERPs in visual search that this object-based bias for an irrelevant feature also appears in an unattended object when it shares that feature with the target object. Specifically, we show that the ERP response elicited by a distractor object in one visual field is modulated as a function of whether a task-irrelevant color of that distractor is also present in the target object that is presented in the opposite visual field. Importantly, we find this modulation to arise with a delay of approximately 80 msec relative to the N2pc--a component of the ERP response that reflects the focusing of attention onto the target. In a second experiment, we demonstrate that this modulation reflects enhanced neural processing in the unattended object. These observations together facilitate the surprising conclusion that the object-based selection of irrelevant features is spatially global even after attention has selected the target object.