AIM: A network model of attention in auditory cortex

Attentional modulation of cortical networks is critical for the cognitive flexibility required to process complex scenes. Current theoretical frameworks for attention are based almost exclusively on studies in visual cortex, where attentional effects are typically modest and excitatory. In contrast, attentional effects in auditory cortex can be large and suppressive. A theoretical framework for explaining attentional effects in auditory cortex is lacking, preventing a broader understanding of cortical mechanisms underlying attention. Here, we present a cortical network model of attention in primary auditory cortex (A1). A key mechanism in our network is attentional inhibitory modulation (AIM) of cortical inhibitory neurons. In this mechanism, top-down inhibitory neurons disinhibit bottom-up cortical circuits, a prominent circuit motif observed in sensory cortex. Our results reveal that the same underlying mechanisms in the AIM network can explain diverse attentional effects on both spatial and frequency tuning in A1. We find that a dominant effect of disinhibition on cortical tuning is suppressive, consistent with experimental observations. Functionally, the AIM network may play a key role in solving the cocktail party problem. We demonstrate how attention can guide the AIM network to monitor an acoustic scene, select a specific target, or switch to a different target, providing flexible outputs for solving the cocktail party problem.     

Effects of Presentation Rate and Attention on Auditory Discrimination: A Comparison of Long-Latency Auditory Evoked Potentials in School-Aged Children and Adults

Decoding human speech requires both perception and integration of brief, successive auditory stimuli that enter the central nervous system as well as the allocation of attention to language-relevant signals. This study assesses the role of attention on processing rapid transient stimuli in adults and children. Cortical responses (EEG/ERPs), specifically mismatch negativity (MMN) responses, to paired tones (standard 100–100Hz; deviant 100–300Hz) separated by a 300, 70 or 10ms silent gap (ISI) were recorded under Ignore and Attend conditions in 21 adults and 23 children (6–11 years old). In adults, an attention-related enhancement was found for all rate conditions and laterality effects (L>R) were observed. In children, 2 auditory discrimination-related peaks were identified from the difference wave (deviant-standard): an early peak (eMMN) at about 100–300ms indexing sensory processing, and a later peak (LDN), at about 400–600ms, thought to reflect reorientation to the deviant stimuli or “second-look” processing. Results revealed differing patterns of activation and attention modulation for the eMMN in children as compared to the MMN in adults: The eMMN had a more frontal topography as compared to adults and attention played a significantly greater role in childrens’ rate processing. The pattern of findings for the LDN was consistent with hypothesized mechanisms related to further processing of complex stimuli. The differences between eMMN and LDN observed here support the premise that separate cognitive processes and mechanisms underlie these ERP peaks. These findings are the first to show that the eMMN and LDN differ under different temporal and attentional conditions, and that a more complete understanding of children’s responses to rapid successive auditory stimulation requires an examination of both peaks.     

Attentional Demands Influence Vocal Compensations to Pitch Errors Heard in Auditory Feedback

Auditory feedback is required to maintain fluent speech. At present, it is unclear how attention modulates auditory feedback processing during ongoing speech. In this event-related potential (ERP) study, participants vocalized/a/, while they heard their vocal pitch suddenly shifted downward a ½ semitone in both single and dual-task conditions. During the single-task condition participants passively viewed a visual stream for cues to start and stop vocalizing. In the dual-task condition, participants vocalized while they identified target stimuli in a visual stream of letters. The presentation rate of the visual stimuli was manipulated in the dual-task condition in order to produce a low, intermediate, and high attentional load. Visual target identification accuracy was lowest in the high attentional load condition, indicating that attentional load was successfully manipulated. Results further showed that participants who were exposed to the single-task condition, prior to the dual-task condition, produced larger vocal compensations during the single-task condition. Thus, when participants’ attention was divided, less attention was available for the monitoring of their auditory feedback, resulting in smaller compensatory vocal responses. However, P1-N1-P2 ERP responses were not affected by divided attention, suggesting that the effect of attentional load was not on the auditory processing of pitch altered feedback, but instead it interfered with the integration of auditory and motor information, or motor control itself.     

Sensory gain control (amplification) as a mechanism of selective attention: electrophysiological and neuroimaging evidence.

Both physiological and behavioral studies have suggested that stimulus-driven neural activity in the sensory pathways can be modulated in amplitude during selective attention. Recordings of event-related brain potentials indicate that such sensory gain control or amplification processes play an important role in visual-spatial attention. Combined event-related brain potential and neuroimaging experiments provide strong evidence that attentional gain control operates at an early stage of visual processing in extrastriate cortical areas. These data support early selection theories of attention and provide a basis for distinguishing between separate mechanisms of attentional suppression (of unattended inputs) and attentional facilitation (of attended inputs).     

How do we select perceptions and actions? Human brain imaging studies.

The selective nature of human perception and action implies a modulatory interaction between sensorimotor processes and attentional processes. This paper explores the use of functional imaging in humans to explore the mechanisms of perceptual selection and the fate of irrelevant stimuli that are not selected. Experiments with positron emission tomography show that two qualitatively different patterns of modulation of cerebral blood flow can be observed in experiments where non-spatial visual attention and auditory attention are manipulated. These patterns of modulation of cerebral blood flow modulation can be described as gain control and bias signal mechanisms. In visual and auditory cortex, the dominant change in cerebral blood flow associated with attention to either modality is related to a bias signal. The relation of these patterns of modulation to attentional effects that have been observed in single neurons is discussed. The existence of mechanisms for selective perception raises the more general question of whether irrelevant ignored stimuli are nevertheless perceived. Lavie''s theory of attention proposes that the degree to which ignored stimuli are processed varies depending on the perceptual load of the current task. Evidence from behavioural and functional magnetic resonance imaging studies of ignored visual motion processing is presented in support of this proposal.     

Precognitive and cognitive elements in sound localization

Sound localization behavior is of great importance for an animal's survival. To localize a sound, animals have to detect a sound source and assign a location to it. In this review we discuss recent results on the underlying mechanisms and on modulatory influences in the barn owl, an auditory specialist with very well developed capabilities to localize sound. Information processing in the barn owl auditory pathway underlying the computations of detection and localization is well understood. This analysis of the sensory information primarily determines the following orienting behavior towards the sound source. However, orienting behavior may be modulated by cognitive (top-down) influences such as attention. We show how advanced stimulation techniques can be used to determine the importance of different cues for sound localization in quasi-realistic stimulation situations, how attentional influences can improve the response to behaviorally relevant stimuli, and how attention can modulate related neural responses. Taken together, these data indicate how sound localization might function in the usually complex natural environment.     

Multisensory integration and attention in autism spectrum disorder: evidence from event-related potentials

Successful integration of various simultaneously perceived perceptual signals is crucial for social behavior. Recent findings indicate that this multisensory integration (MSI) can be modulated by attention. Theories of Autism Spectrum Disorders (ASDs) suggest that MSI is affected in this population while it remains unclear to what extent this is related to impairments in attentional capacity. In the present study Event-related potentials (ERPs) following emotionally congruent and incongruent face-voice pairs were measured in 23 high-functioning, adult ASD individuals and 24 age- and IQ-matched controls. MSI was studied while the attention of the participants was manipulated. ERPs were measured at typical auditory and visual processing peaks, namely, P2 and N170. While controls showed MSI during divided attention and easy selective attention tasks, individuals with ASD showed MSI during easy selective attention tasks only. It was concluded that individuals with ASD are able to process multisensory emotional stimuli, but this is differently modulated by attention mechanisms in these participants, especially those associated with divided attention. This atypical interaction between attention and MSI is also relevant to treatment strategies, with training of multisensory attentional control possibly being more beneficial than conventional sensory integration therapy.     

Effects of aerobic exercise and gender on visual and auditory P300, reaction time, and accuracy

Visual and auditory reaction times (RTs) have been reported to decrease during moderate aerobic exercise, and this has been interpreted as reflecting an exercise-induced activation (EIA) of cognitive information processing. In the present study we examined changes in several independent measures of information processing (RT, accuracy, P300 latency and amplitude) during exercise, and their relationship to visual or auditory modalities and to gender. P300 latencies offer independent measures of cognitive speed that are unrelated to motor output, and P300 amplitudes have been used as measures of attentional allocation. Twenty-four healthy college students [mean (SD) age 20 (2) years] performed auditory and visual "oddball" tasks during resting baseline, aerobic exercise, and recovery periods. Consistent with previous studies, both visual and auditory RTs during exercise were significantly shortened compared to control and recovery periods (which did not differ from each other). We now report that, paralleling the RT changes, auditory and visual P300 latencies decreased during exercise, indicating the occurrence of faster cognitive information processing in both sensory modalities. However, both auditory and visual P300 amplitudes decreased during exercise, suggesting diminished attentional resource allocation. In addition, error rates increased during exercise. Taken together, these results suggest that the enhancement of cognitive information processing speed during moderate aerobic exercise, although operating across genders and sensory modalities, is not a global facilitation of cognition, but is accompanied by decreased attention and increased errors.     

Effects of Attention to Auditory Motion on Cortical Activations during Smooth Pursuit Eye Tracking

Background

In contrast to traditional views that consider smooth pursuit as a relatively automatic process, evidence has been reported for the importance of attention for accurate pursuit performance. However, the exact role that attention might play in the maintenance of pursuit remains unclear.

Methodology/Principal Findings

We analysed the neuronal activity associated with healthy subjects executing smooth pursuit eye movements (SPEM) during concurrent attentive tracking of a moving sound source, which was either in-phase or in antiphase to the executed eye movements. Assuming that attentional resources must be allocated to the moving sound source, the simultaneous execution of SPEM and auditory tracking in diverging directions should result in increased load on common attentional resources. By using an auditory stimulus as a distractor rather then a visual stimulus we guaranteed that cortical activity cannot be caused by conflicts between two simultaneous visual motion stimuli. Our results revealed that the smooth pursuit task with divided attention led to significantly higher activations bilaterally in the posterior parietal cortex and lateral and medial frontal cortex, presumably containing the parietal, frontal and supplementary eye fields respectively.

Conclusions

The additional cortical activation in these areas is apparently due to the process of dividing attention between the execution of SPEM and the covert tracking of the auditory target. On the other hand, even though attention had to be divided the attentional resources did not seem to be exhausted, since the identification of the direction of the auditory target and the quality of SPEM were unaffected by the congruence between visual and auditory motion stimuli. Finally, we found that this form of task-related attention modulated not only the cortical pursuit network in general but also affected modality specific and supramodal attention regions.     

P50 Sensory Gating in Infants

Attentional deficits are common in a variety of neuropsychiatric disorders including attention deficit-hyperactivity disorder, autism, bipolar mood disorder, and schizophrenia. There has been increasing interest in the neurodevelopmental components of these attentional deficits; neurodevelopmental meaning that while the deficits become clinically prominent in childhood or adulthood, the deficits are the results of problems in brain development that begin in infancy or even prenatally. Despite this interest, there are few methods for assessing attention very early in infancy. This report focuses on one method, infant auditory P50 sensory gating.Attention has several components. One of the earliest components of attention, termed sensory gating, allows the brain to tune out repetitive, noninformative sensory information. Auditory P50 sensory gating refers to one task designed to measure sensory gating using changes in EEG. When identical auditory stimuli are presented 500 ms apart, the evoked response (change in the EEG associated with the processing of the click) to the second stimulus is generally reduced relative to the response to the first stimulus (i.e. the response is "gated"). When response to the second stimulus is not reduced, this is considered a poor sensory gating, is reflective of impaired cerebral inhibition, and is correlated with attentional deficits.Because the auditory P50 sensory gating task is passive, it is of potential utility in the study of young infants and may provide a window into the developmental time course of attentional deficits in a variety of neuropsychiatric disorders. The goal of this presentation is to describe the methodology for assessing infant auditory P50 sensory gating, a methodology adapted from those used in studies of adult populations.     

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.     

Interaction between Attention and Bottom-Up Saliency Mediates the Representation of Foreground and Background in an Auditory Scene

The mechanism by which a complex auditory scene is parsed into coherent objects depends on poorly understood interactions between task-driven and stimulus-driven attentional processes. We illuminate these interactions in a simultaneous behavioral–neurophysiological study in which we manipulate participants' attention to different features of an auditory scene (with a regular target embedded in an irregular background). Our experimental results reveal that attention to the target, rather than to the background, correlates with a sustained (steady-state) increase in the measured neural target representation over the entire stimulus sequence, beyond auditory attention's well-known transient effects on onset responses. This enhancement, in both power and phase coherence, occurs exclusively at the frequency of the target rhythm, and is only revealed when contrasting two attentional states that direct participants' focus to different features of the acoustic stimulus. The enhancement originates in auditory cortex and covaries with both behavioral task and the bottom-up saliency of the target. Furthermore, the target's perceptual detectability improves over time, correlating strongly, within participants, with the target representation's neural buildup. These results have substantial implications for models of foreground/background organization, supporting a role of neuronal temporal synchrony in mediating auditory object formation.     

Modality-specific control of strategic spatial attention in parietal cortex

The neural basis of selective spatial attention presents a significant challenge to cognitive neuroscience. Recent neuroimaging studies have suggested that regions of the parietal and temporal cortex constitute a "supramodal" network that mediates goal-directed attention in multiple sensory modalities. Here we used transcranial magnetic stimulation (TMS) to determine which cortical subregions control strategic attention in vision and touch. Healthy observers undertook an orienting task in which a central arrow cue predicted the location of a subsequent visual or somatosensory target. To determine the attentional role of cortical subregions at different stages of processing, TMS was delivered to the right hemisphere during cue or target events. Results indicated a critical role of the inferior parietal cortex in strategic orienting to visual events, but not to somatosensory events. These findings are inconsistent with the existence of a supramodal attentional network and instead provide direct evidence for modality-specific attentional processing in parietal cortex.     

Attention modulates earliest responses in the primary auditory and visual cortices

A fundamental question about the neural correlates of attention concerns the earliest sensory processing stage that it can affect. We addressed this issue by recording magnetoencephalography (MEG) signals while subjects performed detection tasks, which required employment of spatial or nonspatial attention, in auditory or visual modality. Using distributed source analysis of MEG signals, we found that, contrary to previous studies that used equivalent current dipole (ECD) analysis, spatial attention enhanced the initial feedforward response in the primary visual cortex (V1) at 55-90 ms. We also found attentional modulation of the putative primary auditory cortex (A1) activity at 30-50 ms. Furthermore, we reproduced our findings using ECD modeling guided by the results of distributed source analysis and suggest a reason why earlier studies using ECD analysis failed to identify the modulation of earliest V1 activity.     

Acutely Elevated Cortisol in Response to Stressor Is Associated with Attentional Bias Toward Depression-Related Stimuli but Is Not Associated with Attentional Function

Cortisol induces attentional bias toward a negative stimulus and impaired attentional function. Depressed individuals have high levels of cortisol, and exhibit an attentional bias toward a depression-related stimulus and impaired processing speed and executive attention, which are components of attentional function. Therefore, the study tested the hypotheses that an acute increase in cortisol in response to a stressor is associated with attentional bias toward a depression-related stimulus and impaired processing speed and executive attention. Thirty-six participants were administered the dot-probe task for the measurement of attentional bias toward a depression-related stimulus and the Trail Making Test A and B for the measurement of processing speed and executive attention before and after a mental arithmetic task. It was revealed that attentional bias toward a depression-related stimulus following the stressor was observed only among the responders (i.e., participants with cortisol elevation in response to a stressor). On the other hand, no differences in the performance of processing speed and executive attention were noted between the responders and non-responders. The results indicate that acutely elevated cortisol is related to attentional bias, but is not related to processing speed and executive attention. The results have an implication for the etiology of depression.     

The scope and limits of top-down attention in unconscious visual processing

Attentional selection plays a critical role in conscious perception. When attention is diverted, even salient stimuli fail to reach visual awareness. Attention can be voluntarily directed to a spatial location or a visual feature for facilitating the processing of information relevant to current goals. In everyday situations, attention and awareness are tightly coupled. This has led some to suggest that attention and awareness might be based on a common neural foundation, whereas others argue that they are mediated by distinct mechanisms. A body of evidence shows that visual stimuli can be processed at multiple stages of the visual-processing streams without evoking visual awareness. To illuminate the relationship between visual attention and conscious perception, we investigated whether top-down attention can target and modulate the neural representations of unconsciously processed visual stimuli. Our experiments show that spatial attention can target only consciously perceived stimuli, whereas feature-based attention can modulate the processing of invisible stimuli. The attentional modulation of unconscious signals implies that attention and awareness can be dissociated, challenging a simplistic view of the boundary between conscious and unconscious visual processing.     

Influences of attention on auditory aftereffects following purely visual adaptation

Recently, Kitagawa and Ichihara (2002) demonstrated that visual adaptation to an expanding or contracting disk produces a cross-modal visually-induced auditory loudness aftereffect (VALAE), which they attributed to cross-correlations of motion in three-dimensional space. Our experiments extend their results by providing evidence that attending selectively to one of two competing visual stimuli of the same saliency produces a cross-modal VALAE that favors the attended stimulus. These cross-modal attentional effects suggest the existence of integrative spatial mechanisms between vision and audition that are affected by attention.     

Separate attentional resources for vision and audition

Current models of attention, typically claim that vision and audition are limited by a common attentional resource which means that visual performance should be adversely affected by a concurrent auditory task and vice versa. Here, we test this implication by measuring auditory (pitch) and visual (contrast) thresholds in conjunction with cross-modal secondary tasks and find that no such interference occurs. Visual contrast discrimination thresholds were unaffected by a concurrent chord or pitch discrimination, and pitch-discrimination thresholds were virtually unaffected by a concurrent visual search or contrast discrimination task. However, if the dual tasks were presented within the same modality, thresholds were raised by a factor of between two (for visual discrimination) and four (for auditory discrimination). These results suggest that at least for low-level tasks such as discriminations of pitch and contrast, each sensory modality is under separate attentional control, rather than being limited by a supramodal attentional resource. This has implications for current theories of attention as well as for the use of multi-sensory media for efficient informational transmission.     

The dark side of visual attention

The limited capacity of neural processing restricts the number of objects and locations that can be attended to. Selected events are readily enhanced: the bright side of attention. However, such focal processing comes at a cost, namely, functional blindness for unattended events: the dark side of visual attention. Recent work has advanced our understanding of the neural mechanisms that facilitate visual processing, as well as the neural correlates of unattended, unconscious visual events. Also, new results have revealed how attentional deployment is optimized by non-visual factors such as behavioral set, past experience, and emotional salience.     

视觉注意与神经振荡

人脑每时每刻都要接收大量视觉信息，由于人脑加工信息的能力有限，所以在较大视野内将注意分配给相关信息，同时抑制引起注意分散的不相关信息，对执行目标导向的行为至关重要。这种对视觉信息的选择性和主动性加工以适应当前目标的过程被称作视觉注意（visual attention），且视觉注意可分为自上而下的注意与自下而上的注意两种不同功能。由于来自大脑电信号的神经振荡活动在认知加工中发挥重要作用，已有研究综述了视觉注意与神经振荡（neural oscillation）的密切关系，但并未涉及不同的注意功能与神经振荡的关系。本文系统性调查了不同注意功能与神经振荡的关系，发现额-顶区域的theta频带振荡活动反映了自上而下的认知控制，而后部脑区的theta振荡与自下而上的注意相关。顶-枕区域alpha振荡的偏侧化有助于注意分配，而alpha频带的大规模同步促成了注意对视皮层自上而下的影响。Beta振荡介导了自上而下的信息与自下而上的信息之间的互动，作为信息载体促进了视觉信息处理。Gamma振荡则可能与自上而下和自下而上的注意间整合相关。本文就视觉注意功能与神经振荡关系的研究现状展开综述，旨在揭示不同的神经振荡活动在特定的视觉注意功能中的作用。