Dynamic modulation of visual detection by auditory cues in spatial neglect

One of the most constant findings of studies about selective attention is that detection of visual stimuli is enhanced when a visual cue is presented at the position of the upcoming target. In healthy participants, comparable benefits were reported when the cue was presented in a different modality than the target. The aim of this study was to examine spatial and temporal dynamics of visual attention following auditory cues in patients with spatial neglect. Twelve healthy subjects and five patients with left-sided neglect were asked to react to a small vertical line presented randomly at one of four positions. The target appeared 150 or 1000 ms after an auditory cue that was either static (continuous 380 Hz tone presented to the left or right ear) or dynamic (380 Hz tone moving from the left to the right ear or vice versa). The reaction time pattern of healthy participants was unaffected by the different tones. In contrast, reaction times of neglect patients were significantly faster to left targets following a dynamic tone moving from right to left in comparison to a tone moving from left to right. Interestingly, static unilateral tones modulated visual attention of neglect patients to a lesser degree than dynamic tones. The modulation of visual attention by dynamic auditory cues was of short duration and disappeared after 1000 ms. These results demonstrate a fast automatic shift of spatial attention in the direction of a moving tone, suggesting strong dynamic links between visual and auditory attention in patients with a severe spatial deficit.     

Activation of frontoparietal attention networks by non-predictive gaze and arrow cues

Gaze and arrow cues automatically orient visual attention, even when they have no predictive value, but the neural circuitry by which they direct attention is not clear. Recent evidence has indicated that the ventral frontoparietal attention network is primarily engaged by breaches of a viewer’s cue-related expectations. Accordingly, we hypothesized that to the extent that non-predictive gaze and arrow cues automatically engender expectations with regard to cue location, they should activate the ventral attention network when they cue attention invalidly. Using event-related fMRI, we found that invalid gaze but not arrow cues activated the ventral attention network, specifically in the area of the right temporal parietal junction (TPJ), as well as nodes along the dorsal attention network associated with a redirection of attention to the correct target location. In additional whole-brain analyses, facilitation of behavioral response time by valid gaze cues was linearly associated with the degree of activation in the right TPJ. We conclude from our findings that gaze direction elicits potent expectations in humans with regard to an actor’s intention that engage attention networks if not differently from, at least more robustly than, arrow cues.     

Executive control of spatial attention shifts in the auditory compared to the visual modality

Voluntary orienting of visual spatial attention has been shown to be associated with activation in a distributed network of frontal and parietal brain areas. Neuropsychological data suggest that at least some of these areas should be sensitive to the direction in which attention is shifted. The aim of this study was to use rapid event-related functional magnetic resonance imaging to investigate whether spatial attention in the auditory modality is subserved by the same or different brain areas as in the visual modality, and whether the auditory and visual attention networks show any degree of hemispheric lateralisation or sensitivity to the direction of attention shifts. The results suggest that auditory and visual spatial attention shifts are controlled by a supramodal network of frontal, parietal and temporal areas. Areas activated included the precuneus and superior parietal cortex, the inferior parietal cortex and temporo-parietal junction, as well as the premotor and supplementary motor areas and dorsolateral prefrontal cortex (DLPFC). In the auditory task, some of these areas, in particular the precuneus as well as the inferior parietal cortex and temporo-parietal junction, showed 'relative' asymmetry, in that they responded more strongly to attention shifts towards the contralateral than the ipsilateral hemispace. Some areas, such as the right superior parietal cortex and left DLPFC, showed 'absolute' asymmetry, in that they responded more strongly in one than in the other hemisphere.     

Time course of allocation of spatial attention by acoustic cues in non‐human primates

Spatial attention mediates the selection of information from different parts of space. When a brief cue is presented shortly before a target [cue to target onset asynchrony (CTOA)] in the same location, behavioral responses are facilitated, a process called attention capture. At longer CTOAs, responses to targets presented in the same location are inhibited; this is called inhibition of return (IOR). In the visual modality, these processes have been demonstrated in both humans and non‐human primates, the latter allowing for the study of the underlying neural mechanisms. In audition, the effects of attention have only been shown in humans when the experimental task requires sound localization. Studies in monkeys with the use of similar cues but without a sound localization requirement have produced negative results. We have studied the effects of predictive acoustic cues on the latency of gaze shifts to visual and auditory targets in monkeys experienced in localizing sound sources in the laboratory with the head unrestrained. Both attention capture and IOR were demonstrated with acoustic cues, although with a faster time course than with visual cues. Additionally, the effect was observed across sensory modalities (acoustic cue to visual target), suggesting that the underlying neural mechanisms of these effects may be mediated within the superior colliculus, a center where inputs from both vision and audition converge.     

Modulation of auditory cortex activation by sound presentation rate and attention

We studied the effects of sound presentation rate and attention on auditory supratemporal cortex (STC) activation in 12 healthy adults using functional magnetic resonance imaging (fMRI) at 3 T. The sounds (200 ms in duration) were presented at steady rates of 0.5, 1, 1.5, 2.5, or 4 Hz while subjects either had to focus their attention to the sounds or ignore the sounds and attend to visual stimuli presented with a mean rate of 1 Hz. Consistent with previous observations, we found that both increase in stimulation rate and attention to sounds enhanced activity in STC bilaterally. Further, we observed larger attention effects with higher stimulation rates. This interaction of attention and presentation rate has not been reported previously. In conclusion, our results show both rate-dependent and attention-related modulations of STC indicating that both factors should be controlled, or at least addressed, in fMRI studies of auditory processing.     

Auditory orienting and inhibition of return in mild traumatic brain injury: A FMRI study

The semiacute phase of mild traumatic brain injury (mTBI) is associated with deficits in the cognitive domains of attention, memory, and executive function, which previous work suggests may be related to a specific deficit in disengaging attentional focus. However, to date, there have only been a few studies that have employed dynamic imaging techniques to investigate the potential neurological basis of these cognitive deficits during the semiacute stage of injury. Therefore, event‐related functional magnetic resonance imaging was used to investigate the neurological correlates of attentional dysfunction in a clinically homogeneous sample of 16 patients with mTBI during the semiacute phase of injury (<3 weeks). Behaviorally, patients with mTBI exhibited deficits in disengaging and reorienting auditory attention following invalid cues as well as a failure to inhibit attentional allocation to a cued spatial location compared to a group of matched controls. Accordingly, patients with mTBI also exhibited hypoactivation within thalamus, striatum, midbrain nuclei, and cerebellum across all trials as well as hypoactivation in the right posterior parietal cortex, presupplementary motor area, bilateral frontal eye fields, and right ventrolateral prefrontal cortex during attentional disengagement. Finally, the hemodynamic response within several regions of the attentional network predicted response times better for controls than for patients with mTBI. These objective neurological findings represent a potential biomarker for the behavioral deficits in spatial attention that characterize the initial recovery phase of mTBI. Hum Brain Mapp, 2009. © 2009 Wiley‐Liss, Inc.     

Hemispheric differences in attentional orienting by social cues

Research points to a right hemisphere bias for processing social stimuli. Hemispheric specialization for attention shifts cued by social stimuli, however, has been rarely studied. We examined the capacity of each hemisphere to orient attention in response to social and nonsocial cues using a lateralized spatial cueing paradigm. We compared the up/down orienting effects of eye gaze cues, arrow cues, and peripheral cues (change in luminance). Results revealed similar cueing effects in each visual field for nonsocial cues, but asymmetric effects for social cues. At both short (150 ms) and long (950 ms) cue-target intervals, gaze cueing was significant in the LVF, but not in the RVF. Thus, there is a right hemisphere bias for attentional orienting cued by social stimuli, but not for attentional orienting cued by nonsocial stimuli. This supports a theory of a separate neural system for socially cued orienting of attention, as well as a theory of separate parallel and simultaneous neural systems for attention in the two cerebral hemispheres.     

Temporal cues trick the visual and auditory cortices mimicking spatial cues in blind individuals

In the absence of vision, spatial representation may be altered. When asked to compare the relative distances between three sounds (i.e., auditory spatial bisection task), blind individuals demonstrate significant deficits and do not show an event‐related potential response mimicking the visual C1 reported in sighted people. However, we have recently demonstrated that the spatial deficit disappears if coherent time and space cues are presented to blind people, suggesting that they may use time information to infer spatial maps. In this study, we examined whether the modification of temporal cues during space evaluation altered the recruitment of the visual and auditory cortices in blind individuals. We demonstrated that the early (50–90 ms) occipital response, mimicking the visual C1, is not elicited by the physical position of the sound, but by its virtual position suggested by its temporal delay. Even more impressively, in the same time window, the auditory cortex also showed this pattern and responded to temporal instead of spatial coordinates.     

Neuronal effects of nicotine during auditory selective attention in schizophrenia

Although nicotine has been shown to improve attention deficits in schizophrenia, the neurobiological mechanisms underlying this effect are poorly understood. We hypothesized that nicotine would modulate attention‐associated neuronal response in schizophrenia patients in the ventral parietal cortex (VPC), hippocampus, and anterior cingulate based on previous findings in control subjects. To test this hypothesis, the present study examined response in these regions in a cohort of nonsmoking patients and healthy control subjects using an auditory selective attention task with environmental noise distractors during placebo and nicotine administration. In agreement with our hypothesis, significant diagnosis (Control vs. Patient) X drug (Placebo vs. Nicotine) interactions were observed in the VPC and hippocampus. The interaction was driven by task‐associated hyperactivity in patients (relative to healthy controls) during placebo administration, and decreased hyperactivity in patients after nicotine administration (relative to placebo). No significant interaction was observed in the anterior cingulate. Task‐associated hyperactivity of the VPC predicted poor task performance in patients during placebo. Poor task performance also predicted symptoms in patients as measured by the Brief Psychiatric Rating Scale. These results are the first to suggest that nicotine may modulate brain activity in a selective attention‐dependent manner in schizophrenia. Hum Brain Mapp 37:410–421, 2016. © 2015 Wiley Periodicals, Inc.     

Staring in the eye of auditory neglect: comments on 'gaze direction modulates auditory spatial deficits in stroke patients with neglect'

In the current issue of Cortex, Pavani and colleagues show that directing gaze toward the contralesional hemifield significantly diminishes the auditory deficits associated with unilateral neglect. The authors suggest that this beneficial effect of gaze direction may arise due to the recruitment of crossmodal attentional links between audition and vision. A complementary interpretation of these findings is that directing gaze toward neglected sound sources encourages the recoding of auditory spatial location in a relatively preserved frame of reference, and that it is through this coordinate transformation process that awareness of auditory stimuli may be regained. Rehabilitation strategies aimed at recoding stimuli within relatively preserved reference frames may be a useful step forward in managing unilateral neglect.     

Interaction of spatial and non‐spatial cues in auditory stream segregation in the European starling

Integrating sounds from the same source and segregating sounds from different sources in an acoustic scene are an essential function of the auditory system. Naturally, the auditory system simultaneously makes use of multiple cues. Here, we investigate the interaction between spatial cues and frequency cues in stream segregation of European starlings (Sturnus vulgaris) using an objective measure of perception. Neural responses to streaming sounds were recorded, while the bird was performing a behavioural task that results in a higher sensitivity during a one‐stream than a two‐stream percept. Birds were trained to detect an onset time shift of a B tone in an ABA‐ triplet sequence in which A and B could differ in frequency and/or spatial location. If the frequency difference or spatial separation between the signal sources or both were increased, the behavioural time shift detection performance deteriorated. Spatial separation had a smaller effect on the performance compared to the frequency difference and both cues additively affected the performance. Neural responses in the primary auditory forebrain were affected by the frequency and spatial cues. However, frequency and spatial cue differences being sufficiently large to elicit behavioural effects did not reveal correlated neural response differences. The difference between the neuronal response pattern and behavioural response is discussed with relation to the task given to the bird. Perceptual effects of combining different cues in auditory scene analysis indicate that these cues are analysed independently and given different weights suggesting that the streaming percept arises consecutively to initial cue analysis.     

Voluntary and involuntary attention affect face discrimination differently

Do voluntary (endogenous) and involuntary (exogenous) attention have the same perceptual consequences? Here we used fMRI to examine activity in the fusiform face area (FFA--a region in ventral visual cortex responsive to faces) and frontal-parietal areas (dorsal regions involved in spatial attention) under voluntary and involuntary spatial cueing conditions. The trial and stimulus parameters were identical for both cueing conditions. However, the cue predicted the location of an upcoming target face in the voluntary condition but was nonpredictive in the involuntary condition. The predictable cue condition led to increased activity in the FFA compared to the nonpredictable cue condition. These results show that voluntary attention leads to more activity in areas of the brain associated with face processing than involuntary attention, and they are consistent with differential behavioral effects of attention on recognition-related processes.     

Selective auditory attention effects in tonotopically organized cortical areas: A topographic ERP study

This study reinvestigates some aspects of the neurophysiological mechanisms of auditory selective attention, by testing the hypothesis that auditory attention can exert a selective control over the sensory processing of acoustic stimuli in tonotopic auditory cortex. Event-related potentials (ERPs) were recorded from human subjects while they listened selectively to a tone sequence in one ear and ignored a concurrent sequence of a different frequency in the opposite ear. The tone frequencies were 500, 1,000, 2,000, or 4,000 Hz in separate sequences, and were delivered at a constant rate of one every 800 ms. The effects of attention were analyzed in the difference waves obtained by subtracting ERPs to ignore tones from those of the same tones when attended. The earliest effect of attention (70–80 ms post-stimulus) was found to present the same spatio-temporal organization as the obligatory, sensory-evoked N1 wave, with similar tonotopic changes in scalp distribution with the tone frequencies. At longer latencies, two other attentional effects were observed, of probable endogenous origin: one around 175 ms post-stimulus, possibly originating from non-tonotropic auditory cortex, and the latest one (after 300 ms) from non-specific areas. The results support the hypothesis of a genuine sensory gating mechanism for auditory attention, taking place in tonotopic auditory cortex at an early stage of sensory processing, even in low attentional load conditions. © 1995 Wiley-Liss, Inc.     

Selective attention differentially affects brainstem auditory evoked potentials of electrodermal responders and nonresponders

Brainstem auditory evoked potentials (BAEPs) were recorded in adult male chronic schizophrenics on maintenance neuroleptic therapy, drug-free alcoholics, and normal subjects. Subjects were subdivided into electrodermal responders (Rs) and nonresponders (NRs). Attention was directed toward auditory or visual stimuli. Independent of diagnosis, latencies of BAEP wave V were longer in NRs when visual rather than auditory stimuli were attended to, while there was no task effect for Rs. This finding is interpreted as a sign of excessive selective filtering of auditory stimuli in NRs. In the alcoholic NRs, wave I-V conduction times were longer than those in any other subgroup, possibly indicating retarded neural transmission.     

Mapping feature-sensitivity and attentional modulation in human auditory cortex with functional magnetic resonance imaging

Feature-specific enhancement refers to the process by which selectively attending to a particular stimulus feature specifically increases the response in the same region of the brain that codes that stimulus property. Whereas there are many demonstrations of this mechanism in the visual system, the evidence is less clear in the auditory system. The present functional magnetic resonance imaging (fMRI) study examined this process for two complex sound features, namely frequency modulation (FM) and spatial motion. The experimental design enabled us to investigate whether selectively attending to FM and spatial motion enhanced activity in those auditory cortical areas that were sensitive to the two features. To control for attentional effort, the difficulty of the target-detection tasks was matched as closely as possible within listeners. Locations of FM-related and motion-related activation were broadly compatible with previous research. The results also confirmed a general enhancement across the auditory cortex when either feature was being attended to, as compared with passive listening. The feature-specific effects of selective attention revealed the novel finding of enhancement for the nonspatial (FM) feature, but not for the spatial (motion) feature. However, attention to spatial features also recruited several areas outside the auditory cortex. Further analyses led us to conclude that feature-specific effects of selective attention are not statistically robust, and appear to be sensitive to the choice of fMRI experimental design and localizer contrast.     

Modulating auditory selective attention by non‐invasive brain stimulation: Differential effects of transcutaneous vagal nerve stimulation and transcranial random noise stimulation

Selective attention is a basic process required to maintain goal‐directed behavior by appropriately responding to target stimuli and suppressing reactions to non‐target stimuli. It has been proposed that auditory selective attention is linked to the activity of the locus coeruleus‐norepinergic (LC‐NE) system and a large‐scale fronto‐parietal cortical network, but there is still sparse causal evidence for these assumptions. By applying transcutaneous vagal nerve stimulation (tVNS) and transcranial random noise stimulation (tRNS) over the frontal cortex, we systematically assessed the involvement of these subcortical and cortical components in the regulation of auditory selective attention. Using a single‐blinded, sham‐controlled, within‐subject design we analyzed online effects of tVNS and tRNS in 20 healthy participants during an auditory oddball paradigm. We show significant stimulation‐dependent modulations of auditory selective attention on the behavioral and electrophysiological level. Compared to sham, tVNS increased the P3 amplitude, while tRNS reduced the reaction time to target stimuli. Moreover, both techniques reduced the P3 latency. Our data provide evidence for the functional relevance of the subcortical NE system in the regulation of neural resources that allows a phasic response to incoming target stimuli. They indicate that frontal cortex structures are crucially involved in the successful evaluation of the respective information. Moreover, our results are in favor of the LC‐P3 hypothesis claiming the vital role of the NE system in auditory selective attention and in the generation of the P3. Of note, the effects of tVNS on auditory selective attention are comparable with those evoked by pharmacological interventions and invasive vagal nerve stimulation.     

The role of automatic orienting of attention towards ipsilesional stimuli in non-visual (tactile and auditory) neglect: A critical review

The aim of the present survey was to review scientific articles dealing with the non-visual (auditory and tactile) forms of neglect to determine: (a) whether behavioural patterns similar to those observed in the visual modality can also be observed in the non-visual modalities; (b) whether a different severity of neglect can be found in the visual and in the auditory and tactile modalities; (c) the reasons for the possible differences between the visual and non-visual modalities.Data pointing to a contralesional orienting of attention in the auditory and the tactile modalities in visual neglect patients were separately reviewed.Results showed: (a) that in patients with right brain damage manifestations of neglect for the contralesional side of space can be found not only in the visual but also in the auditory and tactile modalities; (b) that the severity of neglect is greater in the visual than in the non-visual modalities.This asymmetry in the severity of neglect across modalities seems due to the greater role that the automatic capture of attention by irrelevant ipsilesional stimuli seems to play in the visual modality.     

Watching where you look: modulation of visual processing of foveal stimuli by spatial attention

Two experiments investigated the effect of sustained selective spatial attention upon the perceptual analysis of stimuli within the center of gaze. Spatial attention has typically been studied in relation to peripheral stimuli, and its relevance to the processing of central stimuli has remained relatively ignored. Here, behavioral measures in normal human volunteer participants showed that focused spatial attention can also influence responses to central stimuli, over and beyond the advantage conferred by their foveation. Event-related potentials (ERPs) showed that the action of attention upon foveal stimuli included the modulation of perceptual processing in extrastriate visual areas. Surprisingly, the visual modulation revealed an intriguing and consistent pattern of hemispheric asymmetry, in both experiments. These findings show that in addition to the established right hemisphere dominance of the brain areas that direct attention, the consequences of directed attention may also be asymmetrical.     

No auditory conduction abnormality in children with attention deficit hyperactivity disorder

Attention deficit hyperactivity disorder (ADHD) is a neurobehavioral developmental disorder characterized by lack of sustained attention and hyperactivity. It has been suggested that asymmetrical conduction of the auditory stimulus in the brainstem plays a role in the pathophysiological process of ADHD. In the present study, the functional integrity of the central auditory pathway was assessed using the auditory brainstem response (ABR), mid-latency response (MLR) and slow vertex response (SVR). Twenty ADHD children and twenty controls were recruited for the study and recordings were done on a computerized evoked potential recorder using the 10-20 system of electrode placement. There emerged no significant difference in absolute peak latencies, interpeak latencies and amplitude of ABR or latency of MLR in the ADHD children as compared with the controls. Prolongation of the SVR latency was found in the children with ADHD versus the controls, but the difference was statistically insignificant. The present study does not suggest any auditory conduction abnormality as a contributory factor in ADHD.     

The functional consequences of social attention on memory precision and on memory-guided orienting in development

Adults are slower at locating targets in naturalistic scenes containing a social distractor compared to an equally salient non-social distractor, and their subsequent memory for targets in social scenes is poorer. Therefore, adults’ social biases affect not only attention, but also their memory. Six-to-ten year-old children and young adults took part in the current study, employing a combination of behavioural and eye-tracking measures. Social stimuli in naturalistic scenes distracted both children and adults during visual search, as demonstrated by their gaze behavior and search times. In addition, eye-tracking revealed even greater attentional capture by social distractors for children. Memory for targets was worse in social compared to non-social scenes. Intriguingly, children demonstrated overall better memory precision than adults. Finally, when participants detected previously learnt targets within visual scenes, adults were slower for targets appearing at unexpected (invalid) locations within social scenes compared to non-social scenes, but this was not the case for children. In their entirety, these findings suggest that the interplay between social attentional biases, memory and memory-guided attention is complex and modulated by age-related differences. Complementary methodologies in developmental cognitive neuroscience shed light on the mechanisms through which social attention and memory interact over development.