Peripheral Attentional Targets under Covert Attention Lead to Paradoxically Enhanced Alpha Desynchronization in Neurofibromatosis Type 1

The limited capacity of the human brain to process the full extent of visual information reaching the visual cortex requires the recruitment of mechanisms of information selection through attention. Neurofibromatosis type-1 (NF1) is a neurodevelopmental disease often exhibiting attentional deficits and learning disabilities, and is considered to model similar impairments common in other neurodevelopmental disorders such as autism. In a previous study, we found that patients with NF1 are more prone to miss targets under overt attention conditions. This finding was interpreted as a result of increased occipito-parietal alpha oscillations. In the present study, we used electroencephalography (EEG) to study alpha power modulations and the performance of patients with NF1 in a covert attention task. Covert attention was required in order to perceive changes (target offset) of a peripherally presented stimulus. Interestingly, alpha oscillations were found to undergo greater desynchronization under this task in the NF1 group compared with control subjects. A similar pattern of desynchronization was found for beta frequencies while no changes in gamma oscillations could be identified. These results are consistent with the notion that different attentional states and task demands generate different patterns of abnormal modulation of alpha oscillatory processes in NF1. Under covert attention conditions and while target offset was reported with relatively high accuracy (over 90% correct responses), excessive desynchronization was found. These findings suggest an abnormal modulation of oscillatory activity and attentional processes in NF1. Given the known role of alpha in modulating attention, we suggest that alpha patterns can show both abnormal increases and decreases that are task and performance dependent, in a way that enhanced alpha desynchronization may reflect a compensatory mechanism to keep performance at normal levels. These results suggest that dysregulation of alpha oscillations may occur in NF1 both in terms of excessive or diminished activation patterns.     

Attentional Requirements on Feature Search Are Modulated by Stimulus Properties

We report a series of dual-task experiments, in which a rapid serial visual presentation (RSVP) task was combined with a visual search task. Orientation, motion, and color were used as the defining target features in the search task. Lag between target onsets was manipulated and interference between the two tasks was quantified by measuring detection scores for the search task as a function of lag. While simultaneous performance of an orientation detection task with an RSVP letter identification task resulted in a performance decrease for lags up to 320 ms, no such decrease was detected for highly salient motion- and color-defined targets. Subsequently, detectability of the motion and color feature was matched to that of the orientation-feature resulting in the reintroduction of a (smaller) performance decrease, but only during simultaneous performance (lag 0 ms). The results suggest that there are two causes for the impaired search performance occurring when a feature search task is combined with an RSVP task. The first is short-lasting interference probably due to attentional competition; the second, which plays a role only when targets for both tasks share features, is interference that may be attributed to a central processing bottleneck.     

Distractor inhibition predicts individual differences in the attentional blink

Background

The attentional blink (AB) refers to humans'' impaired ability to detect the second of two targets (T2) in a rapid serial visual presentation (RSVP) stream of distractors if it appears within 200–600 ms of the first target (T1). Here we examined whether humans'' ability to inhibit distractors in the RSVP stream is a key determinant of individual differences in T1 performance and AB magnitude.

Methodology/Principal Findings

We presented subjects with RSVP streams (93.3 ms/item) of letters containing white distractors, a red T1 and a green T2. Subjects'' ability to suppress distractors was assessed by determining the extent to which their second target performance was primed by a preceding distractor that shared the same identity as T2. Individual subjects'' magnitude of T2 priming from this distractor was found to be negatively correlated with their T1 accuracy and positively related to their AB magnitude. In particular, subjects with attenuated ABs showed negative priming (i.e., worse T2 performance when the priming distractor appeared in the RSVP stream compared to when it was absent), whereas those with large ABs displayed positive priming (i.e., better T2 performance when the priming distractor appeared in the RSVP stream compared to when it was absent). Thus, a subject''s ability to suppress distractors, as assessed by T2 priming magnitude, predicted both their T1 performance and AB magnitude.

Conclusions/Significance

These results confirm that distractor suppression plays a key role in RSVP target selection and support the hypothesis that the AB results, at least in part, from a failure of distractor inhibition.     

Emotion modulation of visual attention: categorical and temporal characteristics

Background

Experimental research has shown that emotional stimuli can either enhance or impair attentional performance. However, the relative effects of specific emotional stimuli and the specific time course of these differential effects are unclear.

Methodology/Principal Findings

In the present study, participants (n = 50) searched for a single target within a rapid serial visual presentation of images. Irrelevant fear, disgust, erotic or neutral images preceded the target by two, four, six, or eight items. At lag 2, erotic images induced the greatest deficits in subsequent target processing compared to other images, consistent with a large emotional attentional blink. Fear and disgust images also produced a larger attentional blinks at lag 2 than neutral images. Erotic, fear, and disgust images continued to induce greater deficits than neutral images at lag 4 and 6. However, target processing deficits induced by erotic, fear, and disgust images at intermediate lags (lag 4 and 6) did not consistently differ from each other. In contrast to performance at lag 2, 4, and 6, enhancement in target processing for emotional stimuli was observed in comparison to neutral stimuli at lag 8.

Conclusions/Significance

These findings suggest that task-irrelevant emotion information, particularly erotica, impairs intentional allocation of attention at early temporal stages, but at later temporal stages, emotional stimuli can have an enhancing effect on directed attention. These data suggest that the effects of emotional stimuli on attention can be both positive and negative depending upon temporal factors.     

Negative priming under rapid serial visual presentation

Negative priming (NP) was examined under a new paradigm wherein a target and distractors were temporally separated using rapid serial visual presentation (RSVP). The results from the two experiments revealed that (a) NP was robust under RSVP, such that the responses to a target were slower when the target served as a distractor in a previous trial than when it did not; (b) NP was found regardless of whether the distractors appeared before or after the targets; and (c) NP was stronger when the distractor was more distinctive. These findings are generally similar to those on NP in the spatial search task. The implications for the processes causing NP under RSVP are discussed in the current paper.     

Converging levels of analysis in the cognitive neuroscience of visual attention.

Experiments using behavioural, lesion, functional imaging and single neuron methods are considered in the context of a neuropsychological model of visual attention. According to this model, inputs compete for representation in multiple visually responsive brain systems, sensory and motor, cortical and subcortical. Competition is biased by advance priming of neurons responsive to current behavioural targets. Across systems competition is integrated such that the same, selected object tends to become dominant throughout. The behavioural studies reviewed concern divided attention within and between modalities. They implicate within-modality competition as one main restriction on concurrent stimulus identification. In contrast to the conventional association of lateral attentional focus with parietal lobe function, the lesion studies show attentional bias to be a widespread consequence of unilateral cortical damage. Although the clinical syndrome of unilateral neglect may indeed be associated with parietal lesions, this probably reflects an assortment of further deficits accompanying a simple attentional imbalance. The functional imaging studies show joint involvement of lateral prefrontal and occipital cortex in lateral attentional focus and competition. The single unit studies suggest how competition in several regions of extrastriate cortex is biased by advance priming of neurons responsive to current behavioural targets. Together, the concepts of competition, priming and integration allow a unified theoretical approach to findings from behavioural to single neuron levels.     

Intellectual functioning and multi-dimensional attentional processes in long-term survivors of a central nervous system related pediatric malignancy

Aims

Central nervous system (CNS) malignancies and/or their treatment in pediatric cancer survivors are known to be associated with deficits in neuropsychological functions. We report findings from a nation-wide study of childhood cancer survivors to investigate intelligence and attention/concentration from a multi-dimensional perspective in a diverse sample from this population.

Main methods

Four hundred forty-four pediatric cancer survivors between 6 and 17 years of age, who had suffered CNS involvement associated with their malignancy, were evaluated. All patients completed a measure of general intelligence. Attention was measured by a continuous performance test (CPT) and by parental report using a standardized psychological inventory.

Key findings

Social economic status (SES) was a significant predictor of intellectual functioning and scores on independent measures of attention. After controlling for SES, cranial radiation therapy (CRT) was strongly predictive of impairments in intellectual functioning. Patients who had completed a transplant procedure did not have significant impairments in intellectual functioning when compared to other participants. CPT performance was most clearly influenced by a younger age at diagnosis and the presence of a supratentorial brain tumor. Reaction time was lower in patients who had received CRT. Gender did not correlate with CPT performance, but caregiver reports of deficits in attentional functioning were more prevalent in girls compared to boys.

Significance

These findings are important given the large, representative sample and multi-dimensional assessment of attentional functioning. The presence of a very strong SES effect on all dependent variables must be addressed in studies of this nature.     

Event-Related Potential Evidence for Two Functionally Dissociable Sources of Semantic Effects in the Attentional Blink

Three target words (T1, T2, and T3) were embedded in a rapid serial visual presentation (RSVP) stream of non-word distractors, and participants were required to report the targets at the end of each RSVP stream. T2 and T3 were semantically related words in half of the RSVP streams, and semantically unrelated words in the other half of the RSVP streams. Using an identical design, a recent study reported distinct reflections of the T2–T3 semantic relationship on the P2 and N400 components of event-related potentials (ERPs) time-locked to T3, suggesting an early, automatic, source of P2 semantic effects and a late, controlled, source of N400 semantic effects. Here, P2 and N400 semantic effects were examined by manipulating list-wide context. Relative to participants performing in a semantically unbiased context, participants over-exposed to filler RSVP streams always including semantically related T2/T3 words reported a dilution of T3-locked P2 semantic effects and a magnification of T3-locked N400 semantic effects. Opposite effects on P2 and N400 ERP components of list-wide semantic context are discussed in relation to recent proposals on the representational status of RSVP targets at processing stages prior to consolidation in visual short-term memory.     

Impaired Contingent Attentional Capture Predicts Reduced Working Memory Capacity in Schizophrenia

Although impairments in working memory (WM) are well documented in schizophrenia, the specific factors that cause these deficits are poorly understood. In this study, we hypothesized that a heightened susceptibility to attentional capture at an early stage of visual processing would result in working memory encoding problems. 30 patients with schizophrenia and 28 demographically matched healthy participants were presented with a search array and asked to report the orientation of the target stimulus. In some of the trials, a flanker stimulus preceded the search array that either matched the color of the target (relevant-flanker capture) or appeared in a different color (irrelevant-flanker capture). Working memory capacity was determined in each individual using the visual change detection paradigm. Patients needed considerably more time to find the target in the no-flanker condition. After adjusting the individual exposure time, both groups showed equivalent capture costs in the irrelevant-flanker condition. However, in the relevant-flanker condition, capture costs were increased in patients compared to controls when the stimulus onset asynchrony between the flanker and the search array was high. Moreover, the increase in relevant capture costs correlated negatively with working memory capacity. This study demonstrates preserved stimulus-driven attentional capture but impaired contingent attentional capture associated with low working memory capacity in schizophrenia. These findings suggest a selective impairment of top-down attentional control in schizophrenia, which may impair working memory encoding.     

Focusing on attention: the effects of working memory capacity and load on selective attention

Background

Working memory (WM) is imperative for effective selective attention. Distractibility is greater under conditions of high (vs. low) concurrent working memory load (WML), and in individuals with low (vs. high) working memory capacity (WMC). In the current experiments, we recorded the flanker task performance of individuals with high and low WMC during low and high WML, to investigate the combined effect of WML and WMC on selective attention.

Methodology/Principal Findings

In Experiment 1, distractibility from a distractor at a fixed distance from the target was greater when either WML was high or WMC was low, but surprisingly smaller when both WML was high and WMC low. Thus we observed an inverted-U relationship between reductions in WM resources and distractibility. In Experiment 2, we mapped the distribution of spatial attention as a function of WMC and WML, by recording distractibility across several target-to-distractor distances. The pattern of distractor effects across the target-to-distractor distances demonstrated that the distribution of the attentional window becomes dispersed as WM resources are limited. The attentional window was more spread out under high compared to low WML, and for low compared to high WMC individuals, and even more so when the two factors co-occurred (i.e., under high WML in low WMC individuals). The inverted-U pattern of distractibility effects in Experiment 1, replicated in Experiment 2, can thus be explained by differences in the spread of the attentional window as a function of WM resource availability.

Conclusions/Significance

The current findings show that limitations in WM resources, due to either WML or individual differences in WMC, affect the spatial distribution of attention. The difference in attentional constraining between high and low WMC individuals demonstrated in the current experiments helps characterise the nature of previously established associations between WMC and controlled attention.     

Neuronal coherence during selective attentional processing and sensory-motor integration.

Groups of neurons synchronize their activities during a variety of conditions, but whether this synchronization is functionally relevant has remained a matter of debate. Here, we survey recent findings showing that synchronization is dynamically modulated during cognitive processes. Based on this evidence, synchronization appears to reflect a general mechanism that renders interactions among selective subsets of neurons effective. We show that neuronal synchronization predicts which sensory input is processed and how efficient it is transmitted to postsynaptic target neurons during sensory-motor integration. Four lines of evidence are presented supporting the hypothesis that rhythmic neuronal synchronization, also called neuronal coherence, underlies effective and selective neuronal communication. (1) Findings from intracellular recordings strongly suggest that postsynaptic neurons are particularly sensitive to synaptic input that is synchronized in the gamma-frequency (30-90 Hz) range. (2) Neurophysiological studies in awake animals revealed enhanced rhythmic synchronization among neurons encoding task-relevant information. (3) The trial-by-trial variation in the precision of neuronal synchronization predicts part of the trial-by-trial variation in the speed of visuo-motor integration. (4) The planning and selection of specific movements can be predicted by the strength of coherent oscillations among local neuronal groups in frontal and parietal cortex. Thus, neuronal coherence appears as a neuronal substrate of an effective neuronal communication structure that dynamically links neurons into functional groups processing task-relevant information and selecting appropriate actions during attention and effective sensory-motor integration.     

Dissociated deficits in attentional networks in social anxiety and depression

A critical cognitive symptom that is commonly involved in social anxiety and depression is attentional deficit. However, the functional relationship between attentional deficit and these two disorders remains poorly understood. Here, we behaviorally disentangled the three key attentional components(alerting, orienting, and executive control) using the established attentional network task(ANT) to investigate how social anxiety and depression are related to deficits in these attention components. We identified a double dissociation between the symptoms of social anxiety and depression and the attentional component deficits when processing non-emotional stimuli. While individuals vulnerable to social anxiety exhibited deficits in the orienting component, individuals vulnerable to depression were impaired in the executive control component. Our findings showed that social anxiety and depression were associated with deficits in different attentional components, which are not specific to emotional information.     

Quick minds slowed down: effects of rotation and stimulus category on the attentional blink

Background

Most people show a remarkable deficit to report the second of two targets when presented in close temporal succession, reflecting an attentional restriction known as the ‘attentional blink’ (AB). However, there are large individual differences in the magnitude of the effect, with some people showing no such attentional restrictions.

Methodology/Principal Findings

Here we present behavioral and electrophysiological evidence suggesting that these ‘non-blinkers’ can use alphanumeric category information to select targets at an early processing stage. When such information was unavailable and target selection could only be based on information that is processed relatively late (rotation), even non-blinkers show a substantial AB. Electrophysiologically, in non-blinkers this resulted in enhanced distractor-related prefrontal brain activity, as well as delayed target-related occipito-parietal activity (P3).

Conclusion/Significance

These findings shed new light on possible strategic mechanisms that may underlie individual differences in AB magnitude and provide intriguing clues as to how temporal restrictions as reflected in the AB can be overcome.     

The BTBR Mouse Model of Autism Spectrum Disorders Has Learning and Attentional Impairments and Alterations in Acetylcholine and Kynurenic Acid in Prefrontal Cortex

Autism is a complex spectrum of disorders characterized by core behavioral deficits in social interaction, communication, repetitive stereotyped behaviors and restricted interests. Autism frequently presents with additional cognitive symptoms, including attentional deficits and intellectual disability. Preclinical models are important tools for studying the behavioral domains and biological underpinnings of autism, and potential treatment targets. The inbred BTBR T+tf/J (BTBR) mouse strain has been used as an animal model of core behavioral deficits in autism. BTBR mice exhibit repetitive behaviors and deficits in sociability and communication, but other aspects of their cognitive phenotype, including attentional performance, are not well characterized. We examined the attentional abilities of BTBR mice in the 5-choice serial reaction time task (5-CSRTT) using an automated touchscreen testing apparatus. The 5-CSRTT is an analogue of the human continuous performance task of attention, and so both the task and apparatus have translational relevance to human touchscreen cognitive testing. We also measured basal extracellular levels of a panel of neurotransmitters within the medial prefrontal cortex, a brain region critically important for performing the 5-CSRTT. We found that BTBR mice have increased impulsivity, defined as an inability to withhold responding, and decreased motivation, as compared to C57Bl/6J mice. Both of these features characterize attentional deficit disorders in humans. BTBR mice also display decreased accuracy in detecting short stimuli, lower basal levels of extracellular acetylcholine and higher levels of kynurenic acid within the prefrontal cortex. Intact cholinergic transmission in prefrontal cortex is required for accurate performance of the 5-CSRTT, consequently this cholinergic deficit may underlie less accurate performance in BTBR mice. Based on our findings that BTBR mice have attentional impairments and alterations in a key neural substrate of attention, we propose that they may be valuable for studying mechanisms for treatment of cognitive dysfunction in individuals with attention deficits and autism.     

Attention Increases the Temporal Precision of Conscious Perception: Verifying the Neural-ST2 Model

What role does attention play in ensuring the temporal precision of visual perception? Behavioural studies have investigated feature selection and binding in time using fleeting sequences of stimuli in the Rapid Serial Visual Presentation (RSVP) paradigm, and found that temporal accuracy is reduced when attentional control is diminished. To reduce the efficacy of attentional deployment, these studies have employed the Attentional Blink (AB) phenomenon. In this article, we use electroencephalography (EEG) to directly investigate the temporal dynamics of conscious perception. Specifically, employing a combination of experimental analysis and neural network modelling, we test the hypothesis that the availability of attention reduces temporal jitter in the latency between a target''s visual onset and its consolidation into working memory. We perform time-frequency analysis on data from an AB study to compare the EEG trials underlying the P3 ERPs (Event-related Potential) evoked by targets seen outside vs. inside the AB time window. We find visual differences in phase-sorted ERPimages and statistical differences in the variance of the P3 phase distributions. These results argue for increased variation in the latency of conscious perception during the AB. This experimental analysis is complemented by a theoretical exploration of temporal attention and target processing. Using activation traces from the Neural-ST² model, we generate virtual ERPs and virtual ERPimages. These are compared to their human counterparts to propose an explanation of how target consolidation in the context of the AB influences the temporal variability of selective attention. The AB provides us with a suitable phenomenon with which to investigate the interplay between attention and perception. The combination of experimental and theoretical elucidation in this article contributes to converging evidence for the notion that the AB reflects a reduction in the temporal acuity of selective attention and the timeliness of perception.     

Distractor Suppression When Attention Fails: Behavioral Evidence for a Flexible Selective Attention Mechanism

Despite consistent evidence showing that attention is a multifaceted mechanism that can operate at multiple levels of processing depending on the structure and demands of the task, investigations of the attentional blink phenomenon have consistently shown that the impairment in reporting the second of two targets typically occurs at a late, or post-perceptual, stage of processing. This suggests that the attentional blink phenomenon may represent the operation of a unique attentional mechanism that is not as flexible as other attentional mechanisms. To test whether the attentional blink is a fixed or flexible phenomenon, we manipulated first target task demands (i.e., difficulty) and measured the influence this had on processing a subsequently presented distractor and the second target. If the attentional blink represents a mechanism that is fixed and consistently fails at a single stage of processing, then manipulations of task difficulty should not affect distractor processing. However, if the attentional blink represents a more multifaceted and flexible mechanism, then task difficulty should modulate distractor processing. The results revealed that distractor processing during the AB was attenuated under high task difficulty. In addition, unlike previous studies, we failed to find a correlation between distractor processing and the severity of the attentional blink. Using a simulation, we demonstrate that the previously reported correlations may have been spurious and due to using variables that were not independent. Overall, the present results support the conclusion that the selectivity of attention during the AB is flexible and depends on the structure and demands of the task.     

Monarch butterflies reared under autumn-like conditions have more efficient flight and lower post-flight metabolism

1. Many migratory animals undergo physiological and behavioural changes to prepare for and sustain long-distance movements. Because insect migrations are common and diverse, studies that examine how migratory insects meet the energetic demands of long-distance movements are badly needed. 2. Monarch butterflies (Danaus plexippus) migrate up to 4000 km annually from eastern North America to wintering sites in central Mexico. Autumn generation monarchs undergo physiological and behavioural changes in response to environmental cues to initiate migration. In particular, exposure to cooler temperatures and shorter day lengths in early autumn causes monarchs to enter the hormonally induced state of reproductive diapause. 3. This study examined differences in flight-associated metabolic rate (MR) and flight performance metrics for monarchs experimentally reared under autumn-like conditions (typically experienced before the southward migration) relative to monarchs reared under summer-like conditions. 4. Adult monarchs reared under autumn-like conditions showed lower post-flight MRs, greater flight efficiency, and lower measures of reproductive activity relative to monarchs reared under summer-like conditions. Increases in post-flight metabolism were associated with monarch body weight, age, and flight velocity. 5. These findings suggest that a trans-generational shift in flight energetics is an important component of the monarch's complex migratory syndrome, and that physiological changes that accompany reproductive diapause facilitate energy conservation during flight.     

Keep Calm and Carry On: Improved Frustration Tolerance and Processing Speed by Transcranial Direct Current Stimulation (tDCS)

Cognitive control (CC) of attention is a major prerequisite for effective information processing. Emotional distractors can bias and impair goal-directed deployment of attentional resources. Frustration-induced negative affect and cognition can act as internal distractors with negative impact on task performance. Consolidation of CC may thus support task-oriented behavior under challenging conditions. Recently, transcranial direct current stimulation (tDCS) has been put forward as an effective tool to modulate CC. Particularly, anodal, activity enhancing tDCS to the left dorsolateral prefrontal cortex (dlPFC) can increase insufficient CC in depression as indicated by a reduction of attentional biases induced by emotionally salient stimuli. With this study, we provide first evidence that, compared to sham stimulation, tDCS to the left dlPFC enhances processing speed measured by an adaptive version of the Paced Auditory Serial Addition Task (PASAT) that is typically thwarted by frustration. Notably, despite an even larger amount of error-related negative feedback, the task-induced upset was suppressed in the group receiving anodal tDCS. Moreover, inhibition of task-related negative affect was correlated with performance gains, suggesting a close link between enhanced processing speed and consolidation of CC by tDCS. Together, these data provide first evidence that activity enhancing anodal tDCS to the left dlPFC can support focused cognitive processing particularly when challenged by frustration-induced negative affect.     

How Does Information Processing Speed Relate to the Attentional Blink?

Background

When observers are asked to identify two targets in rapid sequence, they often suffer profound performance deficits for the second target, even when the spatial location of the targets is known. This attentional blink (AB) is usually attributed to the time required to process a previous target, implying that a link should exist between individual differences in information processing speed and the AB.

Methodology/Principal Findings

The present work investigated this question by examining the relationship between a rapid automatized naming task typically used to assess information-processing speed and the magnitude of the AB. The results indicated that faster processing actually resulted in a greater AB, but only when targets were presented amongst high similarity distractors. When target-distractor similarity was minimal, processing speed was unrelated to the AB.

Conclusions/Significance

Our findings indicate that information-processing speed is unrelated to target processing efficiency per se, but rather to individual differences in observers'' ability to suppress distractors. This is consistent with evidence that individuals who are able to avoid distraction are more efficient at deploying temporal attention, but argues against a direct link between general processing speed and efficient information selection.     

The role of attention in visual processing

Attention to a visual stimulus typically increases the responses of cortical neurons to that stimulus. Because many studies have shown a close relationship between the performance of individual neurons and behavioural performance of animal subjects, it is important to consider how attention affects this relationship. Measurements of behavioural and neuronal performance taken from rhesus monkeys while they performed a motion detection task with two attentional states show that attention alters the relationship between behaviour and neuronal response. Notably, attention affects the relationship differently in different cortical visual areas. This indicates that a close relationship between neuronal and behavioural performance on a given task persists over changes in attentional state only within limited regions of visual cortex.