Attending points in time and space

Both spatial and temporal attention improves auditory processing and these effects seem to originate at perceptual processing stages. It is not yet known if space and time are used in parallel or sequentially for stimulus selection. To directly compare when temporal and spatial attention affect stimulus processing in the auditory modality, short and long empty intervals (600 and 1,200 ms) were presented. Each interval started with a centrally presented tone (S1) and ended with a second tone (S2) presented either on the left or on the right side. Participants had to attend one point in time (offset of the short or long interval) and one position (left or right side) and had to respond to infrequent, deviant offset markers presented at the attended time point and at the attended position. The N1 of concurrently recorded event-related potentials (ERPs) to the frequent standard stimuli was enhanced by both temporal and spatial attention. The temporal and the spatial N1 attention effect had a similar scalp topography, suggesting common neural generators. By contrast, later effects of temporal and spatial attention, consisting of a posterior positivity and an anterior negativity, markedly differed.     

The influence of a vasopressin-analogue (DGAVP) on event-related potentials in a stimulus-mismatch paradigm

To assess the influence of DGAVP (des-glycinamide-arginine-8-vasopressin, a synthetic vasopressin analogue) on the processing of stimuli and stimulus deviance in humans in a double-blind cross-over experiment, 13 subjects received 60 IU DGAVP versus placebo intranasally 48, 24, and 1 h prior to the experimental session. Auditory event-related potentials (ERPs) were taken as a tool to investigate central nervous processing in an experimental task that required the subjects to count different kinds of rare tone pips deviating in pitch and probability compared to frequent standard tones. Direction of attention was manipulated by instructing the subject to count a different class of deviating tone pips in each attention condition. DGAVP enhanced the N2 of the ERP to tone pips deviating extremely in pitch from standard tones, whether or not the extremely deviating pips were to be attended to by the subjects. This influence suggests an intensified mismatch processing under DGAVP that may result from a general excitatory effect of this substance on cortical activity.     

Perceived timing of first- and second-order changes in vision and hearing

Simultaneous changes in visual stimulus attributes (such as motion or color) are often perceived to occur at different times, a fact usually attributed to differences in neural processing times of those attributes. However, other studies suggest that perceptual misalignments are not due to stimulus attributes, but to the type of change, first- or second-order. To test whether this idea generalizes across modalities, we studied perceptual synchrony of acoustic and of audiovisual cross-modal stimuli, which varied in a first- or second-order fashion. First-order changes were abrupt changes in tone intensity or frequency (auditory), or spatial position (visual), while second-order changes were an inversion of the direction of change, such as a turning point when a rising tone starts falling or a translating visual blob reverses. For both pure acoustic and cross-modal stimuli, first-order changes were systematically perceived before second-order changes. However, when both changes were first-order, or both were second-order, little or no difference in perceptual delay was found between them, regardless of attribute or modality. This shows that the type of attribute change, as well as latency differences, is a strong determinant of subjective temporal alignments. We also performed an analysis of reaction times (RTs) to the first- and second-order attribute changes used in these temporal alignment experiments. RT differences between these stimuli did not correspond with our temporal alignment data, suggesting that subjective alignments cannot be accounted for by a simple latency-based explanation.     

Intermodal attention affects the processing of the temporal alignment of audiovisual stimuli

The temporal asynchrony between inputs to different sensory modalities has been shown to be a critical factor influencing the interaction between such inputs. We used scalp-recorded event-related potentials (ERPs) to investigate the effects of attention on the processing of audiovisual multisensory stimuli as the temporal asynchrony between the auditory and visual inputs varied across the audiovisual integration window (i.e., up to 125 ms). Randomized streams of unisensory auditory stimuli, unisensory visual stimuli, and audiovisual stimuli (consisting of the temporally proximal presentation of the visual and auditory stimulus components) were presented centrally while participants attended to either the auditory or the visual modality to detect occasional target stimuli in that modality. ERPs elicited by each of the contributing sensory modalities were extracted by signal processing techniques from the combined ERP waveforms elicited by the multisensory stimuli. This was done for each of the five different 50-ms subranges of stimulus onset asynchrony (SOA: e.g., V precedes A by 125–75 ms, by 75–25 ms, etc.). The extracted ERPs for the visual inputs of the multisensory stimuli were compared among each other and with the ERPs to the unisensory visual control stimuli, separately when attention was directed to the visual or to the auditory modality. The results showed that the attention effects on the right-hemisphere visual P1 was largest when auditory and visual stimuli were temporally aligned. In contrast, the N1 attention effect was smallest at this latency, suggesting that attention may play a role in the processing of the relative temporal alignment of the constituent parts of multisensory stimuli. At longer latencies an occipital selection negativity for the attended versus unattended visual stimuli was also observed, but this effect did not vary as a function of SOA, suggesting that by that latency a stable representation of the auditory and visual stimulus components has been established.     

Cortical activation by a salient sound modulates visual temporal order judgments: An electrophysiological study of multisensory attentional processes

Previous event-related potential (ERP) studies show that a salient lateral sound activates the visual cortex more strongly contralateral to the sound, observed as an auditory-evoked contralateral occipital positivity (ACOP). Studies showed that this activation enhances the early cortical processing of co-localized visual stimuli presented after, reflected by better detection rates, better discrimination, and sharper perceived contrast. We replicated the ACOP, using earphones, and tested whether auditory cuing can influence temporal order judgments (TOJ) for two visual stimuli (horizontal arrangement) as well as if the ACOP would predict the amplitude of this influence. A lateral salient sound was followed, after 150 or 630 ms, by the visual presentation of a pair of disks, one in left and one in right hemifield, with variable SOA. The TOJ task was to indicate which disk appeared first or which disk appeared second (controlling for response bias). We observed an ACOP at posterior electrode sites and confirmed our hypothesis that the lateral sound influenced TOJ by accelerating the perception of the disk presented on the cued side, even though the sound was irrelevant to the task. Furthermore, the ACOP amplitude was correlated to this visual perceptual change, indicating that a larger change in brain activity was associated with a faster processing of co-localized visual stimuli.     

Evaluating the feasibility of the steady‐state visual evoked potential (SSVEP) to study temporal attention

Improvements in perceptual performance can be obtained when events in the environment are temporally predictable—and temporal predictability improves attention and sensory processing. The amplitude of the steady‐state visual evoked potential (SSVEP) has been shown to correlate with attention paid to a flickering stimulus even if the flickering stimulus is irrelevant for the task. However, to our knowledge, the validity of the SSVEP to study temporal attention has not been established. Therefore, we designed an SSVEP temporal attention task to evaluate whether the SSVEP and its temporal dynamics can be used to study temporal attention. We used a forced‐choice perceptual detection task while presenting task‐irrelevant visual flicker at alpha (10 Hz) and two surrounding frequencies (6 or 15 Hz). Temporal predictability was manipulated by having the interstimulus intervals (ISI) be constant or variable. Behavioral results replicated previous studies confirming the benefits of temporal expectations on performance for trials with constant ISI. EEG analyses revealed robust SSVEP amplitudes for all flicker frequencies, although a main effect of temporal expectations on SSVEP amplitude was not significant. Additional analyses revealed temporal predictability‐related modulations of SSVEP amplitude at 10 Hz and its second harmonic (20 Hz). The effect of temporal predictability was also observed for the 6 Hz flicker, but not for 15 Hz for any ISI condition. These results provide some evidence for the feasibility of the SSVEP technique to study temporal attention for stimuli with flicker frequencies around the alpha band.     

Habituation of the Orienting Response: A Gating Mechanism Subserving Selective Attention

The purpose of this investigation was to determine whether prior habituation of the phasic orienting response (OR) to stimuli which were later to be used as distractors in an attention demanding task, would improve task performance and reduce OR frequency to distractors during the task. The task was the solution of mathematical problems from immediate memory, and the distractors were random numbers and mathematical signs. Subjects were sequentially assigned to one of four conditions: 1) 15 presentations of distractors, problems with distractors; 2) 15 presentations of a tone, problems with distractors; 3) problems with distractors; and 4) 15 presentations of distractors or tone, problems without distractors. Results clearly indicated that prior habituation of the phasic OR to the distractors improved performance relative to no prior habituation or to prior habituation to a tone. There was also evidence of decreased OR activity during problem solving in the prior habituation group relative to the tone habituation and no habituation groups. The data supported an hypothesis that selective attention consists of at least two processes: 1) sensitization of phasic and tonic ORs to salient stimuli, and 2) habituation of phasic and tonic ORs to irrelevant stimuli.     

Neurofeedback training of gamma band oscillations improves perceptual processing

In this study, a noninvasive electroencephalography-based neurofeedback method is applied to train volunteers to deliberately increase gamma band oscillations (40 Hz) in the visual cortex. Gamma band oscillations in the visual cortex play a functional role in perceptual processing. In a previous study, we were able to demonstrate that gamma band oscillations prior to stimulus presentation have a significant influence on perceptual processing of visual stimuli. In the present study, we aimed to investigate longer lasting effects of gamma band neurofeedback training on perceptual processing. For this purpose, a feedback group was trained to modulate oscillations in the gamma band, while a control group participated in a task with an identical design setting but without gamma band feedback. Before and after training, both groups participated in a perceptual object detection task and a spatial attention task. Our results clearly revealed that only the feedback group but not the control group exhibited a visual processing advantage and an increase in oscillatory gamma band activity in the pre-stimulus period of the processing of the visual object stimuli after the neurofeedback training. Results of the spatial attention task showed no difference between the groups, which underlines the specific role of gamma band oscillations for perceptual processing. In summary, our results show that modulation of gamma band activity selectively affects perceptual processing and therefore supports the relevant role of gamma band activity for this specific process. Furthermore, our results demonstrate the eligibility of gamma band oscillations as a valuable tool for neurofeedback applications.     

Effects of nicotine on visuo-spatial selective attention as indexed by event-related potentials

Nicotine has been shown to specifically reduce reaction times to invalidly cued targets in spatial cueing paradigms. In two experiments, we used event-related potentials to test whether the facilitative effect of nicotine upon the detection of invalidly cued targets is due to a modulation of perceptual processing, as indexed by early attention-related event-related potential components. Furthermore, we assessed whether the effect of nicotine on such unattended stimuli depends upon the use of exogenous or endogenous cues. In both experiments, the electroencephalogram was recorded while non-smokers completed discrimination tasks in Posner-type paradigms after chewing a nicotine polacrilex gum (Nicorette 2 mg) in one session and a placebo gum in another session. Nicotine reduced reaction times to invalidly cued targets when cueing was endogenous. In contrast, no differential effect of nicotine on reaction times was observed when exogenous cues were used. Electrophysiologically, we found a similar attentional modulation of the P1 and N1 components under placebo and nicotine but a differential modulation of later event-related potential components at a frontocentral site. The lack of a drug-dependent modulation of P1 and N1 in the presence of a behavioral effect suggests that the effect of nicotine in endogenous visuo-spatial cueing tasks is not due to an alteration of perceptual processes. Rather, the differential modulation of frontocentral event-related potentials suggests that nicotine acts at later stages of target processing.     

Effects of divided attention on temporal processing in patients with lesions of the cerebellum or frontal lobe

Prefrontal cortex and cerebellum have both been implicated in temporal processing tasks although the exact contribution of each system remains unclear. To investigate this issue, control participants and patients with either prefrontal or cerebellar lesions were tested on temporal and nontemporal perceptual tasks under 2 levels of attentional load. Each trial involved a comparison between a standard tone and a subsequent comparison tone that varied in frequency, duration, or both. When participants had to make concurrent judgments on both dimensions, patients with frontal lobe lesions were significantly impaired on both tasks whereas the variability of cerebellar patients increased in the duration task only. This dissociation suggests that deficits on temporal processing tasks observed in frontal patients can be related to the attention demands of such tasks; cerebellar patients have a more specific problem related to timing.     

Cortisol Effects on Attentional Processes in Man as Indicated by Event-Related Potentials

Glacocorticoids which are secreted from the adrenal cortex, particularly in response to stressful stimuli, are assumed to influence brain functions related to stimulus perception and processing. To determine underlying mechanisms of this action the present study investigated the effects of physiologically enhanced plasma glucocurticoid levels on auditory event-related potentials (AERPs) reflecting different aspects of stimulus processing. Sixteen male adult subjects were tested in a dichotic listening paradigm containing four types of tone pips: standard and deviant target pips, which the subject either had to attend to, or to ignore. AERPs to the tone pips provided measures of different perceptual and attentional processes. Subjects were tested in a double-blind fashion according to a within-subject cross-over design under either 16 mg hydrocortisone, infused over the experimental period of 1 hrs, or placebo. Compared to placebo sessions, under cortisol treatment a slightly decreased amplitude of the N1 component was obtained independently of the type of tone pip presented. In addition, mismatch negativity, measured as the difference amplitude between AERP waveforms to standard and deviant target pips in the unattended ear, was substantially reduced by hydrocortisone. Results support an inhibitory action of enhanced plasma cortisol levels on stimulus processing mediated by the nonspecific sensory system.     

ERD as an index of anticipatory attention? Effects of stimulus degradation

Previous research has suggested that the stimulus-preceding negativity (SPN) is largely independent of stimulus modality. In contrast, the scalp topography of the event-related desynchronization (ERD) related to the anticipation of stimuli providing knowledge of results (KR) is modality dependent. These findings, combined with functional SPN research, lead to the hypothesis that anticipatory ERD reflects anticipatory attention, whereas the SPN mainly depends on the affective-motivational properties of the anticipated stimulus. To further investigate the prestimulus ERD, and compare this measure with the SPN, 12 participants performed a time-estimation task, and were informed about the quality of their time estimation by an auditory or a visual stimulus providing KR. The KR stimuli could be either intact or degraded. Auditory degraded KR stimuli were less effective than other KR stimuli in guiding subsequent behavior, and were preceded by a larger SPN. There were no effects of degradation on the SPN in the visual modality. Preceding auditory KR stimuli no ERD was present, whereas preceding visual stimuli an occipital ERD was found. However, contrary to expectation, the latter was larger preceding intact than preceding degraded stimuli. It is concluded that the data largely agree with an interpretation of the pre-KR SPN as a reflection of the anticipation of the affective-motivational value of KR stimuli, and of the prestimulus ERD as a perceptual anticipatory attention process.     

Does temporal preparation speed up visual processing? Evidence from the N2pc

A growing number of studies show that temporal preparation, which denotes processes of anticipation and preparation for an upcoming stimulus, facilitates perceptual processing. Recently, it has been hypothesized that this perceptual benefit arises due to an acceleration of early perceptual processing. Whereas this idea receives some direct support from a recent study showing that temporal preparation reduces the latency of early auditory ERPs, supportive evidence regarding the visual modality lacks so far. To further investigate this acceleration account, we measured the latency of early visual ERPs in a visual search task. We observed that temporal preparation, manipulated via constant foreperiods, reduced the latency of early visual ERPs, specifically the N2pc as an index of attentional target processing. This finding supports a modality‐independent acceleration of perceptual processing by temporal preparation.     

Toward the influence of temporal attention on the selection of targets in a visual search task: An ERP study

We used ERPs to investigate whether temporal attention interacts with spatial attention and feature‐based attention to enhance visual processing. We presented a visual search display containing one singleton stimulus among a set of homogenous distractors. Participants were asked to respond only to target singletons of a particular color and shape that were presented in an attended spatial position. We manipulated temporal attention by presenting a warning signal before each search display and varying the foreperiod (FP) between the warning signal and the search display in a blocked manner. We observed distinctive ERP effects of both spatial and temporal attention. The amplitudes for the N2pc, SPCN, and P3 were enhanced by spatial attention indicating a processing benefit of relevant stimulus features at the attended side. Temporal attention accelerated stimulus processing; this was indexed by an earlier onset of the N2pc component and a reduction in reaction times to targets. Most importantly, temporal attention did not interact with spatial attention or stimulus features to influence visual processing. Taken together, the results suggest that temporal attention fosters visual perceptual processing in a visual search task independently from spatial attention and feature‐based attention; this provides support for the nonspecific enhancement hypothesis of temporal attention.     

Attention modulates auditory pattern memory as indexed by event-related brain potentials

The role of selective attention on auditory pattern processing was investigated using the mismatch negativity, an event-related brain potential component associated with sensory memory. Participants responded to changes in an alternating tone pattern in a designated ear while a similar auditory pattern was presented in the opposite ear. Participants were also presented with the same sequences while reading a book (no response required). In all conditions, changes in the pattern elicited a mismatch negativity (MMN) that peaked at 160–220 ms poststimulus. MMN amplitude varied with attention: the amplitude was higher in response to deviant stimuli presented in the attended ear than to the deviant stimuli presented in the unattended ear or during reading. The results show that selective attention modulates auditory pattern memory.     

Share or compete? Load‐dependent recruitment of prefrontal cortex during dual‐task performance

Dual-task performance requires flexible attention allocation to two or more streams of information. Dorsolateral prefrontal cortex (DLPFC) is considered important for executive function, and recent modeling work proposes that attention control may arise from selective activation and inhibition of different processing units within this region. Here, we used a tone discrimination task and a visual letter memory task to examine whether this type of competition could be measurable using a neuroimaging technique, the event-related optical signal, with high spatial and temporal resolution. Left and right DLPFC structures were differentially affected by task priority and load, with the left middle frontal gyrus (MFG) being preferentially recruited by the visual memory task, whereas the two tasks competed for recruitment, in a spatially segregated manner, in right MFG. The data provide support for a competition view of dual-task processing.     

Early processing stages are modulated when auditory stimuli are presented at an attended moment in time: an event-related potential study

The present study investigated with event-related potentials whether attending to a moment in time modulates the processing of auditory stimuli at a similar early, perceptual level as attending to a location in space. The participants listened to short (600 ms) and long (1,200 ms) intervals marked by white noise bursts. The task was to attend in alternating runs either to the short or to the long intervals and to respond to rare offset markers that differed in intensity from the frequent standard offset markers. Prior to the to-be-attended moment, a slow negative potential developed over the frontal scalp. Stimuli presented at the attended compared to the unattended moments in time elicited an enhanced N1 and an enhanced posteriorly distributed positivity (300-370 ms). The results show that attention can be flexibly controlled in time and that not only late but also early perceptual processing stages are modulated by attending to a moment in time.     

Altered auditory-tactile interactions in congenitally blind humans: an event-related potential study

It has been shown that stimuli of a task-irrelevant modality receive enhanced processing when they are presented at an attended location in space (crossmodal attention). The present study investigated the effects of visual deprivation on the interaction of the intact sensory systems. Random streams of tactile and auditory stimuli were presented at the left or right index finger of congenitally blind participants. They had to attend to one modality (auditory or tactile) of one side (left or right) and had to respond to deviant stimuli of the attended modality and side. While in a group of sighted participants, early event-related potentials (ERPs) were negatively displaced to stimuli presented at the attended position, compared to the unattended, for both the task-relevant and the task-irrelevant modality, starting as early as 80 ms after stimulus onset (unimodal and crossmodal spatial attention effects, respectively), corresponding crossmodal effects could not be detected in the blind. In the sighted, spatial attention effects after 200 ms were only significant for the task-relevant modality, whereas a crossmodal effect for this late time window was observed in the blind. This positive rather than negative effect possibly indicates an active suppression of task-irrelevant stimuli at an attended location in space. The present data suggest that developmental visual input is essential for the use of space to integrate input of the non-visual modalities, possibly because of its high spatial resolution. Alternatively, enhanced perceptual skills of the blind within the intact modalities may result in reduced multisensory interactions ("inverse effectiveness of multisensory integration").     

Examining task-dependencies of different attentional processes as reflected in the P3a and reorienting negativity components of the human event-related brain potential

Unexpected changes in task-irrelevant auditory stimuli are capable to distract processing of task-relevant visual information. This effect is accompanied by the elicitation of event-related potential (ERP) components associated with attentional orientation, i.e. P3a and reorienting negativity (RON). In the present study we varied the demands of a visual task in order to test whether the RON component -- as an index of attentional reorientation after distraction -- is confined to a semantic task requiring working memory. In two ERP experiments we applied an auditory-visual distraction paradigm in which subjects were instructed to discriminate visual stimuli preceded by a task-irrelevant sound, this being either a standard tone (600 Hz, 88%) or a deviant tone (660 Hz, 12%). The visual stimuli were numbers which had to be judged on basis of a semantic (odd or even) or physical feature (either size or colour). As expected, deviance related ERP components namely the mismatch negativity (MMN), P3a, and RON were elicited. Importantly, the RON was affected by the variation of the task: within the semantic task an early RON and within the physical task a late RON was obtained. These results suggest that the RON component reflects two functionally distinct processes of attentional allocation after distraction: refocusing on task-relevant information on the working memory level, and general reorientation of attention, e.g. preparation for the upcoming task.     

Configural processing of other-race faces is delayed but not decreased

The processing of other-race faces has been suggested to differ from own-race face processing with regard to the extent to which configural and/or holistic information is taken into account. We aimed at investigating the underlying mechanisms more precisely by applying event-related potentials (ERP). We presented upright and inverted own-race, other-race, and other-species faces (apes), as well as non-facial control stimuli (houses), and analysed the effects of stimulus type and inversion on the N170. Peak latencies for same-race, other-race, other-species, and non-face stimuli gradually increased, and inversion of all types of face stimuli led to an additional delay. Importantly, the two factors did not interact for face stimuli. Inversion increased N170 amplitudes for own- and other-race faces only. We conclude that early perceptual mechanisms of face processing are not qualitatively different for own- and other-race faces. Instead, the same underlying processes appear to work less efficiently for other-race faces.