Effects of attentional load on early visual processing depend on stimulus timing

A growing number of studies suggest that early visual processing is not only affected by low-level perceptual attributes but also by higher order cognitive factors such as attention or emotion. Using high-density electroencephalography, we recently demonstrated that attentional load of a task at fixation reduces the response of primary visual cortex to irrelevant peripheral stimuli, as indexed by the C1 component. In the latter study, peripheral stimuli were always presented during intervals without task-relevant stimuli. Here, we use a similar paradigm but present central task stimuli and irrelevant peripheral stimuli simultaneously while keeping all other stimulus characteristics constant. Results show that rather than to suppress responses to peripheral stimulation, high attentional load elicits higher C1 amplitudes under these conditions. These findings suggest that stimulus timing can profoundly alter the effects of attentional load on the earliest stages of processing in human visual cortex.     

Effects of attentional load on auditory scene analysis

The effects of attention on the neural processes underlying auditory scene analysis were investigated through the manipulation of auditory task load. Participants were asked to focus their attention on tuned and mistuned stimuli presented to one ear and to ignore similar stimuli presented to the other ear. For both tuned and mistuned sounds, long (standard) and shorter (deviant) duration stimuli were presented in both ears. Auditory task load was manipulated by varying task instructions. In the easier condition, participants were asked to press a button for deviant sounds (target) at the attended location, irrespective of tuning. In the harder condition, participants were further asked to identify whether the targets were tuned or mistuned. Participants were faster in detecting targets defined by duration only than by both duration and tuning. At the unattended location, deviant stimuli generated a mismatch negativity wave at frontocentral sites whose amplitude decreased with increasing task demand. In comparison, standard mistuned stimuli generated an object-related negativity at central sites whose amplitude was not affected by task difficulty. These results show that the processing of sound sequences is differentially affected by attentional load than is the processing of sounds that occur simultaneously (i.e., sequential vs. simultaneous grouping processes), and that they each recruit distinct neural networks.     

Earliest stages of visual cortical processing are not modified by attentional load

This study investigated the effects of attentional load on neural responses to attended and irrelevant visual stimuli by recording high‐density event‐related potentials (ERPs) from the scalp in normal adult subjects. Peripheral (upper and lower visual field) and central stimuli were presented in random order at a rapid rate while subjects responded to targets among the central stimuli. Color detection and color‐orientation conjunction search tasks were used as the low‐ and high‐load tasks, respectively. Behavioral results showed significant load effects on both accuracy and reaction time for target detections. ERP results revealed no significant load effect on the initial C1 component (60–100 ms) evoked by either central‐relevant or peripheral‐irrelevant stimuli. Source analysis with dipole modeling confirmed previous reports that the C1 includes the initial evoked response in primary visual cortex. Source analyses indicated that high attentional load enhanced the early (70–140 ms) neural response to central‐relevant stimuli in ventral‐lateral extrastriate cortex, whereas load effects on peripheral‐irrelevant stimulus processing started at 110 ms and were localized to more dorsal and anterior extrastriate cortical areas. These results provide evidence that the earliest stages of visual cortical processing are not modified by attentional load and show that attentional load affects the processing of task relevant and irrelevant stimuli in different ways. Hum Brain Mapp 35:3008–3024, 2014. © 2013 Wiley Periodicals, Inc .     

Covert manual response preparation triggers attentional modulations of visual but not auditory processing.

OBJECTIVE: We investigated whether covert unimanual response preparation triggers attention shifts, as postulated by the premotor theory of attention, and whether these result in spatially specific modulations of visual and auditory processing. METHODS: Visual response cues instructed participants to prepare to lift their left or right index finger in response to a subsequent target stimulus. Irrelevant visual or auditory probes were delivered to the left or right hand during the response preparation interval. ERPs were measured time-locked to cue onset, and time-locked to probe stimulus onset. RESULTS: Lateralised ERP components triggered during covert response preparation (ADAN, LDAP) were similar to components previously found during attention shifts. N1 components were enhanced to visual probes delivered adjacent to the cued response relative to those delivered to the opposite hand. Auditory probe ERPs were unaffected by manual response preparation. CONCLUSIONS: Shifts of spatial attention that are triggered during covert unimanual response preparation result in spatially specific modulations of visual but not auditory processing. SIGNIFICANCE: Results support the claim of the premotor theory that the preparation of manual responses is associated with attention shifts. However, such shifts are not based on purely supramodal processes, as they result in a modality-specific pattern of sensory modulations.     

Effects of prismatic adaptation on spatial gradients in unilateral neglect: A comparison of visual and auditory target detection with central attentional load

Prismatic adaptation is increasingly recognised as an effective procedure for rehabilitating symptoms of unilateral spatial neglect - producing relatively long-lasting improvements on a variety of spatial attention tasks. The mechanisms by which the aftereffects of adaptation change neglect patients’ performance on these tasks remain controversial. It is not clear, for example, whether adaptation directly influences the pathological ipsilesional attention bias that underlies neglect, or whether it simply changes exploratory motor behaviour. Here we used visual and auditory versions of a target detection task with a secondary task at fixation. Under these conditions, patients with neglect demonstrated a spatial gradient in their ability to orient to the brief, peripheral visual or auditory targets. Following prism adaptation, we found that overall performance on both the auditory and visual task improved, however, most patients in our sample did not show changes in their visual or auditory spatial gradient of attention, despite adequate aftereffects of adaptation and significant improvement in neglect on visual cancellation. Although there were individual cases that suggested prism-induced changes in visual target detection, and even reversal of the visual spatial gradient, such cases were not evident for the auditory modality. The findings indicate that spatial gradients in stimulus-driven attention may be less responsive to the effects of prism adaptation than neglect symptoms in voluntary orienting and exploratory behaviour. Individual factors such as lesion site and symptom severity may also determine the expression of prism effects on spatial neglect.     

Top-down attentional processing enhances auditory evoked gamma band activity

In contrast to animal studies, relatively little is known about the functional significance of the early evoked gamma band activity in humans. We investigated whether evoked and induced 40 Hz activity differentiate automatic, bottom-up aspects of attention from voluntary, top-down related attentional demands. An auditory novelty-oddball task was applied to 14 healthy subjects. As predicted, more evoked gamma was found for the target condition than in the two task-irrelevant conditions. Since gamma band activity was not enhanced for novel stimuli, the evoked gamma response cannot be explained with a simple concept of stimulus arousal. Neither induced gamma nor the degree of 40 Hz phase-locking were different between the experimental conditions. Taken together, our data emphasize the role of evoked gamma band activity for top-down attentional processing.     

Hemispheric asymmetries of visual and auditory information processing

The lateralization of visual and auditory functions, interhemispheric interactions and their relationship to speech dominance were studied. Spectral analysis of visual and auditory evoked potentials (EPs) recorded from left and right speech dominant subjects was employed. The results suggested a complementary processing of the temporal and spatial properties of the afferent information in the speech dominant and non-dominant hemispheres, respectively.     

Reward reduces conflict by enhancing attentional control and biasing visual cortical processing

How does motivation interact with cognitive control during challenging behavioral conditions? Here, we investigated the interactions between motivation and cognition during a response conflict task and tested a specific model of the effect of reward on cognitive processing. Behaviorally, participants exhibited reduced conflict during the reward versus no-reward condition. Brain imaging results revealed that a group of subcortical and fronto-parietal regions was robustly influenced by reward at cue processing and, importantly, that cue-related responses in fronto-parietal attentional regions were predictive of reduced conflict-related signals in the medial pFC (MPFC)/ACC during the upcoming target phase. Path analysis revealed that the relationship between cue responses in the right intraparietal sulcus (IPS) and interference-related responses in the MPFC during the subsequent target phase was mediated via signals in the left fusiform gyrus, which we linked to distractor-related processing. Finally, reward increased functional connectivity between the right IPS and both bilateral putamen and bilateral nucleus accumbens during the cue phase, a relationship that covaried with across-individual sensitivity to reward in the case of the right nucleus accumbens. Taken together, our findings are consistent with a model in which motivationally salient cues are employed to upregulate top-down control processes that bias the selection of visual information, thereby leading to more efficient stimulus processing during conflict conditions.     

Effects of auditory and visual deprivation on human brain development

Brain imaging studies of congenitally deaf and congenitally blind adults are summarized. The results suggest that different subsystems within vision and audition display different degrees of experience-dependent modification of cortical organization. Within vision, the organization of systems important in processing peripheral space and motion information is most altered following auditory deprivation. Within audition, fast auditory processing and attention to and localization within peripheral space is enhanced following visual deprivation. Hypotheses concerning the origins of these differential effects of early experience are discussed.     

Working memory load improves early stages of independent visual processing

Increasing evidence suggests that working memory and perceptual processes are dynamically interrelated due to modulating activity in overlapping brain networks. However, the direct influence of working memory on the spatio-temporal brain dynamics of behaviorally relevant intervening information remains unclear. To investigate this issue, subjects performed a visual proximity grid perception task under three different visual-spatial working memory (VSWM) load conditions. VSWM load was manipulated by asking subjects to memorize the spatial locations of 6 or 3 disks. The grid was always presented between the encoding and recognition of the disk pattern. As a baseline condition, grid stimuli were presented without a VSWM context. VSWM load altered both perceptual performance and neural networks active during intervening grid encoding. Participants performed faster and more accurately on a challenging perceptual task under high VSWM load as compared to the low load and the baseline condition. Visual evoked potential (VEP) analyses identified changes in the configuration of the underlying sources in one particular period occurring 160-190 ms post-stimulus onset. Source analyses further showed an occipito-parietal down-regulation concurrent to the increased involvement of temporal and frontal resources in the high VSWM context. Together, these data suggest that cognitive control mechanisms supporting working memory may selectively enhance concurrent visual processing related to an independent goal. More broadly, our findings are in line with theoretical models implicating the engagement of frontal regions in synchronizing and optimizing mnemonic and perceptual resources towards multiple goals.     

The effect of non-visual working memory load on top-down modulation of visual processing

While a core function of the working memory (WM) system is the active maintenance of behaviorally relevant sensory representations, it is also critical that distracting stimuli are appropriately ignored. We used functional magnetic resonance imaging to examine the role of domain-general WM resources in the top-down attentional modulation of task-relevant and irrelevant visual representations. In our dual-task paradigm, each trial began with the auditory presentation of six random (high load) or sequentially ordered (low load) digits. Next, two relevant visual stimuli (e.g., faces), presented amongst two temporally interspersed visual distractors (e.g., scenes), were to be encoded and maintained across a 7-s delay interval, after which memory for the relevant images and digits was probed. When taxed by high load digit maintenance, participants exhibited impaired performance on the visual WM task and a selective failure to attenuate the neural processing of task-irrelevant scene stimuli. The over-processing of distractor scenes under high load was indexed by elevated encoding activity in a scene-selective region-of-interest relative to low load and passive viewing control conditions, as well as by improved long-term recognition memory for these items. In contrast, the load manipulation did not affect participants’ ability to upregulate activity in this region when scenes were task-relevant. These results highlight the critical role of domain-general WM resources in the goal-directed regulation of distractor processing. Moreover, the consequences of increased WM load in young adults closely resemble the effects of cognitive aging on distractor filtering [Gazzaley, A., Cooney, J. W., Rissman, J., & D’Esposito, M. (2005). Top-down suppression deficit underlies working memory impairment in normal aging. Nature Neuroscience 8, 1298-1300], suggesting the possibility of a common underlying mechanism.     

Effects of lesions of temporal-parietal junction on perceptual and attentional processing in humans

When stimuli with larger forms (global) containing smaller forms (local) are presented to subjects with large lesions in the right hemisphere, they are more likely to miss the global form than the local form, whereas subjects with large lesions in the left are more likely to miss the local than the global form. The present study tested whether the global/local impairment in subjects with posterior lesions was due to deficits in controlled attentional processes, passive perceptual processes, or both. Attentional control was examined by measuring reaction time changes when the probability of a target appearing at either the global or local level was varied. Patients with unilateral right or left lesions centered in temporal-parietal regions and age-matched controls served as subjects. Because neurophysiological and neuropsychological evidence have implicated temporal regions in visual discrimination and inferior parietal regions in the allocation of attention to locations in the visual field, patients with left hemisphere lesions were further subdivided into those with lesions centered in the superior temporal gyrus (LSTG) or rostral inferior parietal lobule (LIPL). Patients with right hemisphere injury could not be analogously subdivided. The results revealed that the LSTG group was able to control the allocation of attention to global and local levels normally, while the LIPL group was not. In contrast, the LSTG group showed a strong baseline reaction time advantage toward global targets, while normals and the LIPL group showed no advantage toward one level or the other. Finally, the perceptual component was affected differentially by lesions in the right hemisphere and LSTG, with lesions in the left favoring global targets and lesions in the right favoring local targets. These findings indicate that the hemispheric global/local asymmetry is due to a perceptual mechanism with a critical anatomical locus centered in the STG.     

Grapheme-color synesthetes show enhanced crossmodal processing between auditory and visual modalities

Synesthesia is an involuntary experience in which stimulation of one sensory modality triggers additional, atypical sensory experiences. Strong multisensory processes are present in the general population, but the relationship between these ‘normal’ sensory interactions and synesthesia is currently unknown. Neuroimaging research suggests that some forms of synesthesia are caused by enhanced cross-activation between brain areas specialized for the processing of different sensory attributes, and finds evidence of increased white matter connections among regions known to be involved in typical crossmodal processes. Using two classic crossmodal integration tasks we show that grapheme-color synesthetes exhibit enhanced crossmodal interactions between auditory and visual modalities, suggesting that the experience of synesthesia in one modality generalizes to enhanced crossmodal processes with other modalities. This finding supports our conjecture that the atypical sensory experiences of synesthetes represent a selective expression of a more diffuse propensity toward ‘typical’ crossmodality interactions.     

Effects of expectation on negative potentials during visual processing

ERPs were recorded during several RT tasks: simple RT; oddball choice RT; a LIE condition in which subjects were told stimuli would infrequently change, but did not; differential responding to two equiprobable stimuli that were randomized in one condition and alternated in another condition. Subtracting ERPs elicited during simple RT from those elicited during the other conditions, it was found that a negative component, NA, was enhanced, relative to simple RT, in all the other RT tasks. The data of the LIE condition indicated that NA was enhanced by the expectation that unpredictable stimulus changes would occur, even when they did not. The data of the 50/50 alternating RT condition indicated that stimulus changes by themselves enhance NA, even when they are predictable. There appear to be several deflections that comprise NA. NA was obtained with a variety of subtractions that balanced stimulus probability, the structure of the stimulus sequence and task instructions. Similar results were obtained whether subjects made a finger lift response or counted stimuli.     

Effects of passive tactile and auditory stimuli on left visual neglect

Patients with left-sided visual neglect fail to copy the left part of drawings or the drawings on the left side of a sheet of paper. Our aim was to study the variations in copying drawings induced by passive stimulation in patients with left-sided visual neglect. No stimulation at all, tactile unilateral and bilateral, binaural auditory verbal, and nonverbal stimuli were randomly applied to 14 patients with right-hemisphere strokes. Only nonverbal stimuli decreased the neglect. As nonverbal stimuli mainly activate the right hemisphere, the decrease in neglect suggests right-hemispheric hypoactivity at rest in these patients. The absence of modification of neglect during verbal stimulation suggests a bilateral hemispheric activation and the persistence of interhemispheric imbalance. Our results showed that auditory pathways take part in the network involved with neglect. Passive nonverbal auditory stimuli may be of interest in the rehabilitation of patients with left visual neglect.     

Top-down modulation of auditory processing: effects of sound context, musical expertise and attentional focus

By recording auditory electrical brain potentials, we investigated whether the basic sound parameters (frequency, duration and intensity) are differentially encoded among speech vs. music sounds by musicians and non-musicians during different attentional demands. To this end, a pseudoword and an instrumental sound of comparable frequency and duration were presented. The accuracy of neural discrimination was tested by manipulations of frequency, duration and intensity. Additionally, the subjects' attentional focus was manipulated by instructions to ignore the sounds while watching a silent movie or to attentively discriminate the different sounds. In both musicians and non-musicians, the pre-attentively evoked mismatch negativity (MMN) component was larger to slight changes in music than in speech sounds. The MMN was also larger to intensity changes in music sounds and to duration changes in speech sounds. During attentional listening, all subjects more readily discriminated changes among speech sounds than among music sounds as indexed by the N2b response strength. Furthermore, during attentional listening, musicians displayed larger MMN and N2b than non-musicians for both music and speech sounds. Taken together, the data indicate that the discriminative abilities in human audition differ between music and speech sounds as a function of the sound-change context and the subjective familiarity of the sound parameters. These findings provide clear evidence for top-down modulatory effects in audition. In other words, the processing of sounds is realized by a dynamically adapting network considering type of sound, expertise and attentional demands, rather than by a strictly modularly organized stimulus-driven system.     

Efferent gating of human auditory attentional processes

Brainstem evoked potentials were measured while subjects reduced either frontalis or lips/throat activity in response to visual information regarding EMG activity while attending to dichotically presented click stimuli. An inhibition of left ear activity occurred at the level of the cochlear nucleus (Wave 1) during articulatory muscle activity, suggesting that the right ear advantage is related to active inhibition of ipsilateral auditory pathways. Contralateral central inhibition of disattended information at the level of the brainstem (Wave 5) was also implicated. A neurophysiological model accounting for both attenuation of irrelevant stimuli and vigilance for high priority information is proposed.     

Clinical neurophysiology of visual and auditory processing in dyslexia: A review

Neurophysiological studies on children and adults with dyslexia provide a deeper understanding of how visual and auditory processing in dyslexia might relate to reading deficits. The goal of this review is to provide an overview of research findings in the last two decades on motion related and contrast sensitivity visual evoked potentials and on auditory event related potentials to basic tone and speech sound processing in dyslexia. These results are particularly relevant for three important theories about causality in dyslexia: the magnocellular deficit hypothesis, the temporal processing deficit hypothesis and the phonological deficit hypothesis. Support for magnocellular deficits in dyslexia are primarily provided from evidence for altered visual evoked potentials to rapidly moving stimuli presented at low contrasts. Consistently ERP findings revealed altered neurophysiological processes in individuals with dyslexia to speech stimuli, but evidence for deficits processing certain general acoustic information relevant for speech perception, such as frequency changes and temporal patterns, are also apparent.     

Effects of aging on peripheral and central auditory processing in rats

Hearing loss is a hallmark sign in the elderly population. Decline in auditory perception provokes deficits in the ability to localize sound sources and reduces speech perception, particularly in noise. In addition to a loss of peripheral hearing sensitivity, changes in more complex central structures have also been demonstrated. Related to these, this study examines the auditory directional maps in the deep layers of the superior colliculus of the rat. Hence, anesthetized Sprague–Dawley adult (10 months) and aged (22 months) rats underwent distortion product of otoacoustic emissions (DPOAEs) to assess cochlear function. Then, auditory brainstem responses (ABRs) were assessed, followed by extracellular single‐unit recordings to determine age‐related effects on central auditory functions. DPOAE amplitude levels were decreased in aged rats although they were still present between 3.0 and 24.0 kHz. ABR level thresholds in aged rats were significantly elevated at an early (cochlear nucleus – wave II) stage in the auditory brainstem. In the superior colliculus, thresholds were increased and the tuning widths of the directional receptive fields were significantly wider. Moreover, no systematic directional spatial arrangement was present among the neurons of the aged rats, implying that the topographical organization of the auditory directional map was abolished. These results suggest that the deterioration of the auditory directional spatial map can, to some extent, be attributable to age‐related dysfunction at more central, perceptual stages of auditory processing.     

Effects of DBS on auditory and somatosensory processing in Parkinson's disease

Motor symptoms of Parkinson's disease (PD) can be relieved by deep brain stimulation (DBS). The mechanism of action of DBS is largely unclear. Magnetoencephalography (MEG) studies on DBS patients have been unfeasible because of strong magnetic artifacts. An artifact suppression method known as spatiotemporal signal space separation (tSSS) has mainly overcome these difficulties. We wanted to clarify whether tSSS enables noninvasive measurement of the modulation of cortical activity caused by DBS. We have studied auditory and somatosensory‐evoked fields (AEFs and SEFs) of advanced PD patients with bilateral subthalamic nucleus (STN) DBS using MEG. AEFs were elicited by 1‐kHz tones and SEFs by electrical pulses to the median nerve with DBS on and off. Data could be successfully acquired and analyzed from 12 out of 16 measured patients. The motor symptoms were significantly relieved by DBS, which clearly enhanced the ipsilateral auditory N100m responses in the right hemisphere. Contralateral N100m responses and somatosensory P60m responses also had a tendency to increase when bilateral DBS was on. MEG with tSSS offers a novel and powerful tool to investigate DBS modulation of the evoked cortical activity in PD with high temporal and spatial resolution. The results suggest that STN‐DBS modulates auditory processing in advanced PD. Hum Brain Mapp, 2011. © 2010 Wiley‐Liss, Inc.