Spatial attention triggered by unimodal, crossmodal, and bimodal exogenous cues: a comparison of reflexive orienting mechanisms

The aim of this study was to establish whether spatial attention triggered by bimodal exogenous cues acts differently as compared to unimodal and crossmodal exogenous cues due to crossmodal integration. In order to investigate this issue, we examined cuing effects in discrimination tasks and compared these effects in a condition wherein a visual target was preceded by both visual and auditory exogenous cues delivered simultaneously at the same side (bimodal cue), with conditions wherein the visual target was preceded by either a visual (unimodal cue) or an auditory cue (crossmodal cue). The results of two experiments revealed that cuing effects on RTs in these three conditions with an SOA of 200 ms had comparable magnitudes. Differences at a longer SOA of 600 ms (inhibition of return for bimodal cues, Experiment 1) disappeared when catch trials were included (in Experiment 2). The current data do not support an additional influence of crossmodal integration on exogenous orienting, but are well in agreement with the existence of a supramodal spatial attention module that allocates attentional resources towards stimulated locations for different sensory modalities.     

Evidence for multisensory integration in the elicitation of prior entry by bimodal cues

This study reports an experiment investigating the relative effects of intramodal, crossmodal and bimodal cues on visual and auditory temporal order judgements. Pairs of visual or auditory targets, separated by varying stimulus onset asynchronies, were presented to either side of a central fixation (±45°), and participants were asked to identify the target that had occurred first. In some of the trials, one of the targets was preceded by a short, non-predictive visual, auditory or audiovisual cue stimulus. The cue and target stimuli were presented at the exact same locations in space. The point of subjective simultaneity revealed a consistent spatiotemporal bias towards targets at the cued location. For the visual targets, the intramodal cue elicited the largest, and the crossmodal cue the smallest, bias. The bias elicited by the bimodal cue fell between the intramodal and crossmodal cue biases, with significant differences between all cue types. The pattern for the auditory targets was similar apart from a scaling factor and greater variance, so the differences between the cue conditions did not reach significance. These results provide evidence for multisensory integration in exogenous attentional cueing. The magnitude of the bimodal cueing effect was equivalent to the average of the facilitation elicited by the intramodal and crossmodal cues. Under the assumption that the visual and auditory cues were equally informative, this is consistent with the notion that exogenous attention, like perception, integrates multimodal information in an optimal way.     

Crossmodal attention between vision and touch in temporal order judgment task

Four experiments investigated the effects of cross-modal attention between vision and touch in temporal order judgment tasks combined with spatial cueing paradigm. In Experiment 1, two vibrotactile stimuli with simultaneous or successive onsets were presented bimanually to the left and right index fingers and participants were asked to judge the temporal order of the two stimuli. The tactile stimuli were preceded by a spatially uninformative visual cue. Results indicated that shift of spatial attention yielded by visual cueing resulted in the modulation of accuracy of the subsequent tactile temporal order judgment. However, this cueing effect disappeared when participants judged simultaneity of the two stimuli, instead of their temporal order (Experiment 2) or when the cue lead time between the visual cue and the stimuli was relatively long (Experiment 3). Experiment 4 replicated an effect of crossmodal attention on the direction of visual illusory line motion induced by a somatosensory cue (Shimojo, Miyauchi, & Hikosaka, 1997). These results demonstrate that substantial crossmodal links exist between vision and touch for covert exogenous orienting of attention.     

Crossmodal links in spatial attention are mediated by supramodal control processes: evidence from event-related potentials

Crossmodal links in spatial attention were studied in an experiment where participants had to detect peripheral tactile or visual targets on the attended side, while ignoring all stimuli on the unattended side and in the currently irrelevant modality. Both relevant locations and relevant modalities were specified on a trial-by-trial basis by auditory precues. Spatial orienting in the cue-target interval was reflected in anterior negativities and occipital positivities contralateral to the cued side, either when vision or touch was cued as relevant. These effects resembled previously reported ERP modulations during shifts of visual attention, implicating supramodal mechanisms in the control of spatial attention and demonstrating their independence of cue modality. Early effects of spatial attention on somatosensory and visual ERPs were of equivalent size for currently relevant and irrelevant modalities. Results support the idea that crossmodal links in spatial attention are mediated by supramodal control mechanisms.     

Assessing the effect of verbal working memory load on visuo-spatial exogenous orienting

The aim of this study was to assess whether or not loading verbal working memory (WM) affects the exogenous orienting of visuo-spatial attention. Visuo-spatial exogenous orienting was measured under low/high WM load conditions by means of an orthogonal spatial cuing paradigm. Participants had to discriminate the elevation (up versus down) of a visual target preceded by a peripheral spatially-nonpredictive visual cue presented on either the left or right. This elevation discrimination task was carried out in-between a verbal WM test, in which a sequence of six digits (to be remembered for report at the end of the trial) was presented either in numerically ascending or random order (i.e., low and high load conditions, respectively). Participants made significantly more recall errors in the high load than in the low load condition, demonstrating that the verbal WM load manipulation had been effective. However, WM load did not interact with visuo-spatial exogenous orienting (i.e., comparable cuing effects were reported in both load conditions). These results are consistent with the view that peripheral visual onsets automatically capture spatial attention, regardless any concurrent increase of verbal WM load.     

Directing visual attention with spatially informative and spatially noninformative tactile cues

We investigated the tactile cuing of visual spatial attention using spatially-informative (75% valid) and spatially-noninformative (25% valid) tactile cues. The participants performed a visual change detection task following the presentation of a tactile spatial cue on their back whose location corresponded to one of the four visual quadrants on a computer monitor. The participants were explicitly instructed to use the spatially-informative tactile cues but to ignore the spatially-noninformative cues. In addition to reaction time data, participants’ eye-gaze was monitored as a measure of overt visual attention. The results showed that the spatially-informative tactile cues resulted in initial saccades toward the cued visual quadrants, and significantly reduced the visual change detection latencies. When spatially-noninformative tactile cues were used, the participants were largely successful at ignoring them as indicated by a saccade distribution that was independent of the quadrant that was cued, as well as the lack of a significant change in search time as compared to the baseline measure of no tactile cuing. The eye-gaze data revealed that the participants could not always completely ignore the spatially-noninformative tactile cues. Our results suggest that the tactile cuing of visual attention is natural but not automatic when the tactile cue and visual target are not collocated spatially, and that it takes effort to ignore the cues even when they are known to provide no useful information. In addition, our results confirm previous findings that spatially-informative tactile cues are especially effective at directing overt visual attention to locations that are not typically monitored visually, such as the bottom of a computer screen or the rearview mirror in an automobile.     

Dissociating inhibition of return from endogenous orienting of spatial attention: Evidence from detection and discrimination tasks

In the present series of experiments, peripheral informative cues were used in order to dissociate endogenous and exogenous orienting of spatial attention using the same set of stimuli. For each block of trials, the cue predicted either the same or the opposite location of target appearance. Crucially, using this manipulation, both expected and unexpected locations could be either cued or uncued. If one accepts the hypothesis that inhibition of return (IOR) is an attentional effect that inhibits the returning of attention to a previously attended location (Posner & Cohen, 1984), one would not predict an IOR effect at the expected location, since attention should not disengage from the location predicted by the cue. Detection and discrimination tasks were used to examine any potential difference in the mechanism responsible for IOR as a function of the task at hand. Two major results emerged: First, IOR was consistently observed at the expected location, where, according to the traditional "reorienting" hypothesis, IOR is not supposed to occur. Second, a different time course of cueing effects was found in detection versus discrimination tasks, even after controlling for the orienting of attention. We conclude that IOR cannot be accounted for solely by the "reorienting of attention" hypothesis. Moreover, we argue that the observed time course differences in cueing effects between detection and discrimination tasks cannot be explained by attention disengaging from cues later in discrimination than in detection tasks, as proposed by Klein (2000). The described endogenous-exogenous dissociation is consistent with models postulating that endogenous and exogenous attentional processes rely on different neural mechanisms.     

Reflexive spatial orienting of tactile attention

We investigated the covert reflexive (exogenous) orienting of tactile spatial attention. Participants made speeded discrimination responses (up vs down) to a series of tactile targets presented randomly to the index finger or thumb of either hand. These targets were preceded at a variable stimulus onset asynchrony (200, 300, or 400 ms) by a spatially nonpredictive tactile cue (presented to both finger and thumb of one or other hand) on either the same or opposite side as the target. Tactile elevation discrimination responses were more rapid and accurate when the cue and target appeared on the same side than when they appeared on opposite sides. Our results provide the first direct empirical evidence that tactile spatial attention can be reflexively directed toward peripheral tactile cues leading to the facilitation of subsequent responses to stimuli presented at that body site.     

The N2pc component and its links to attention shifts and spatially selective visual processing

The N2pc component has recently become a popular tool in attention research. To investigate whether this component exclusively reflects attentional target selection or also prior stages in attentional processing (covert orienting, target-unspecific spatial attention), a spatial cuing procedure was combined with a visual search task. In some blocks, informative cues indicated the side of upcoming singleton targets that were present on most trials among uniform distractors. In other blocks, cues were spatially uninformative, and no preparatory shifts of attention were possible. The N2pc in response to targets was unaffected by this manipulation, showing that this component is not associated with attention shifts. Following informative cues, an attenuated N2pc was elicited by uniform nontarget arrays, suggesting that the N2pc may also reflect spatially specific processing of stimulus features at task-relevant locations prior to target selection.     

Temporal dynamics of lateralized ERP components elicited during endogenous attentional shifts to relevant tactile events

To investigate the temporal dynamics of lateralized event-related brain potential (ERP) components elicited during covert shifts of spatial attention, ERPs were recorded in a task where central visual symbolic cues instructed participants to direct attention to their left or right hand in order to detect infrequent tactile targets presented to that hand, and to ignore tactile stimuli presented to the other hand, as well as all randomly intermingled peripheral visual stimuli. In different blocks, the stimulus onset asynchrony (SOA) between cue and target was 300 ms, 700 ms, or 1,100 ms. Anterior and posterior ERP modulations sensitive to the direction of an attentional shift were time-locked to the attentional cue, rather than to the anticipated arrival of a task-relevant stimulus. These components thus appear to reflect central attentional control rather than the anticipatory preparation of sensory areas. In addition, attentional modulations of ERPs to task-irrelevant visual stimuli were found, providing further evidence for crossmodal links in spatial attention between touch and vision.     

Crossmodal exogenous orienting improves the accuracy of temporal order judgments

Although many studies have demonstrated that crossmodal exogenous orienting can lead to a facilitation of reaction times, the issue of whether exogenous spatial orienting also affects the accuracy of perceptual judgments has proved to be much more controversial. Here, we examined whether or not exogenous spatial attentional orienting would affect sensitivity in a temporal discrimination task. Participants judged which of the two target letters, presented on either the same or opposite sides, had been presented first. A spatially non-predictive tone was presented 200 ms prior to the onset of the first visual stimulus. In two experiments, we observed improved performance (i.e., a decrease in the just noticeable difference) when the target letters were presented on opposite sides and the auditory cue was presented on the side of the first visual stimulus, even when central fixation was monitored ("Experiment 2"). A shift in the point of subjective simultaneity was also observed in both experiments, indicating ‘prior entry’ for cued as compared to uncued first target trials. No such JND or PSS effects were observed when the auditory tone was presented after the second visual stimulus ("Experiment 3"), thus confirming the attentional nature of the effects observed. These findings clearly show that the crossmodal exogenous orienting of spatial attention can affect the accuracy of temporal judgments.

Preparatory states in crossmodal spatial attention: spatial specificity and possible control mechanisms

We used event-related functional magnetic resonance imaging to study the neural correlates of endogenous spatial attention for vision and touch. We examined activity associated with attention-directing cues (central auditory pure tones), symbolically instructing subjects to attend to one hemifield or the other prior to upcoming stimuli, for a visual or tactile task. In different sessions, subjects discriminated either visual or tactile stimuli at the covertly attended side, during bilateral visuotactile stimulation. To distinguish cue-related preparatory activity from any modulation of stimulus processing, unpredictably on some trials only the auditory cue was presented. The use of attend-vision and attend-touch blocks revealed whether preparatory attentional effects were modality-specific or multimodal. Unimodal effects of spatial attention were found in somatosensory cortex for attention to touch, and in occipital areas for attention to vision, both contralateral to the attended side. Multimodal spatial effects (i.e. effects of attended side irrespective of task-relevant modality) were detected in contralateral intraparietal sulcus, traditionally considered a multimodal brain region; and also in the middle occipital gyrus, an area traditionally considered purely visual. Critically, all these activations were observed even on cue-only trials, when no visual or tactile stimuli were subsequently presented. Endogenous shifts of spatial attention result in changes of brain activity prior to the presentation of target stimulation (baseline shifts). Here, we show for the first time the separable multimodal and unimodal components of such preparatory activations. Additionally, irrespective of the attended side and modality, attention-directing auditory cues activated a network of superior frontal and parietal association areas that may play a role in voluntary control of spatial attention for both vision and touch. Electronic Publication     

Orienting of visual attention in dyslexia: evidence for asymmetric hemispheric control of attention

The purpose of this study was to determine whether the transport of small hydrophilic molecules across the blood-brain barrier (BBB) during focal cerebral ischemia could be altered by a topical application of endothelin-1 (ET-1) in the ischemic cortex (IC). Forty minutes after middle cerebral artery (MCA) occlusion, patches of 10 nM ET-1 (low-endothelin group), 100 nM ET-1 (high-endothelin group), or normal saline (control group) were placed on the IC of rats for a 20-min period. One hour after MCA occlusion, transfer coefficient (K _i) of [¹⁴C-α-]aminoisobutyric acid (¹⁴C-AIB) or regional cerebral blood flow (rCBF) was determined. Vital signs were not significantly different among the experimental groups. In the control group (n=8), the K _i of the IC was significantly higher than that of the contralateral cortex (CC; 11.9±5.8 vs 5.0±1.9 μl/g per minute). In the low-endothelin group (n=8), the K _i of the IC was still significantly higher than that of the CC (9.4±5.2 vs 5.3±2.5 μl/g per minute). However, in the High-endothelin group (n=8), the K _i of the IC was not different from that of the CC (6.9±2.1 vs 5.6±2.3 μl/g per minute) and 42% lower than that of the control group. The rCBF was not affected by 100 nM of ET-1 [control (n=6): IC 53±18 ml/100 g per minute, CC 94±23 ml/100 g per minute; high-endothelin (n=6): IC 49±15 ml/100 g per minute, CC 98±24 ml/100 g per minute]. Our data suggest that the application of endothelin-1 in the IC could reduce the transfer coefficient of small hydrophilic molecules across the BBB during focal ischemia. Electronic Publication     

The time course of spatial orienting elicited by central and peripheral cues: evidence from event-related brain potentials

To study differences in the time course of attentional orienting triggered by salient peripheral events and by central symbolic precues, event-related brain potentials (ERPs) were recorded in response to letter stimuli following spatially informative symbolic or peripheral precues after a cue-target interval (CTI) of either 200 or 700 ms. Stimuli at cued (attended) locations elicited an enhanced negativity relative to stimuli at uncued locations. With short CTIs, these effects started around 150 ms post-stimulus for peripheral cues. They were delayed by about 100 ms for central cues. This latency difference is assumed to reflect fast exogenous orienting elicited by peripheral, but not by central cues. Beyond 200 ms post-stimulus, attentional negativities were larger with long CTIs than with short CTIs for both cue types, presumably related to the gradual build-up of endogenous orienting triggered by spatially predictive events.     

Role of audiovisual synchrony in driving head orienting responses

Many studies now suggest that optimal multisensory integration sometimes occurs under conditions where auditory and visual stimuli are presented asynchronously (i.e. at asynchronies of 100 ms or more). Such observations lead to the suggestion that participants’ speeded orienting responses might be enhanced following the presentation of asynchronous (as compared to synchronous) peripheral audiovisual spatial cues. Here, we report a series of three experiments designed to investigate this issue. Upon establishing the effectiveness of bimodal cuing over the best of its unimodal components (Experiment 1), participants had to make speeded head-turning or steering (wheel-turning) responses toward the cued direction (Experiment 2), or an incompatible response away from the cue (Experiment 3), in response to random peripheral audiovisual stimuli presented at stimulus onset asynchronies ranging from ?100 to 100 ms. Race model inequality analysis of the results (Experiment 1) revealed different mechanisms underlying the observed multisensory facilitation of participants’ head-turning versus steering responses. In Experiments 2 and 3, the synchronous presentation of the component auditory and visual cues gave rise to the largest facilitation of participants’ response latencies. Intriguingly, when the participants had to subjectively judge the simultaneity of the audiovisual stimuli, the point of subjective simultaneity occurred when the auditory stimulus lagged behind the visual stimulus by 22 ms. Taken together, these results appear to suggest that the maximally beneficial behavioural (head and manual) orienting responses resulting from peripherally presented audiovisual stimuli occur when the component signals are presented in synchrony. These findings suggest that while the brain uses precise temporal synchrony in order to control its orienting responses, the system that the human brain uses to consciously judge synchrony appears to be less fine tuned.     

Interruption from irrelevant auditory and visual onsets even when attention is in a focused state

The common view on the interplay between exogenous and endogenous orienting holds that abrupt onsets are not capable of attracting attention when they occur outside the current focus of attention. Does this also apply to sudden irrelevant auditory onsets and when irrelevant visual onsets occur far in the periphery? In addition, does focused attention also reduce the alerting effect of auditory onsets, or vice versa, do highly alerting stimuli distort the attentional state? Crossmodal and unimodal variants of the Posner paradigm were examined in two experiments with targets and irrelevant onsets occurring at 28.3 and 19.3° from fixation. Either centrally presented arrows indicated the forthcoming position of visual targets to be discriminated, or warning cues signaled the likely moment of target occurrence. The targets could be preceded by peripheral auditory or visual onsets that were to be ignored. Crossmodal and unimodal exogenous orienting effects of these irrelevant onsets were observed while participants focused at the relevant side. In addition, no evidence was found that the alerting effect of auditory onsets was dependent on focused attention. Our findings indicate that, at least under the current conditions, neither crossmodal nor unimodal orienting effects of peripheral events dissipate when attention is in a focused state.     

Response interference in touch,vision, and crossmodally: beyond the spatial dimension

To date, tactile distractor processing has primarily been investigated by focusing on the spatial characteristics of distractors and the impact of their presentation on the orienting of attention. In two experiments, we examined the influence of tactile distractors when the location of stimulus presentation was kept constant, thus controlling for the effects of spatial attention. A response priming paradigm was used in which two stimuli were sequentially presented from the same (fixated) direction. Typically, target responses are facilitated when the previously presented distractor (i.e., the prime) happens to map on to the same response as compared to the distractor maps on to the opposite response. Similar response priming effects were observed for tactile and visual distractors within a unimodal experimental setting (Experiments 1a and 1b). Interestingly, however, when the targets and distractors were presented in different sensory modalities, only the visual distractors exerted a crossmodal effect on the subsequent processing of vibrotactile targets (Experiment 2). These results therefore indicate that visual stimuli automatically trigger their corresponding response even when the task at hand is not visual, whereas tactile stimuli are only processed up to the level of response generation when the participants’ task is tactile.     

The modulation of crossmodal integration by unimodal perceptual grouping: a visuotactile apparent motion study

We adapted the crossmodal dynamic capture task to investigate the modulation of visuotactile crossmodal integration by unimodal visual perceptual grouping. The influence of finger posture on this interaction was also explored. Participants were required to judge the direction of a tactile apparent motion stream (moving either to the left or to the right) presented to their crossed or uncrossed index fingers. The participants were instructed to ignore a distracting visual apparent motion stream, comprised of either 2 or 6 lights presented concurrently with the tactile stimuli. More crossmodal dynamic capture of the direction of the tactile apparent motion stream by the visual apparent motion stream was observed in the 2-lights condition than in the 6-lights condition. This interaction was not modulated by finger posture. These results suggest that visual intramodal perceptual grouping constrains the crossmodal binding of visual and tactile apparent motion information, irrespective of finger posture.     

Location and features of instructive spatial cues do not influence the time course of covert shifts of visual spatial attention

The time course of shifting visual spatial attention to flickering stimuli in the left and right visual hemifield was investigated. The goal was to test whether an instructive peripheral salient cue located close to the newly to-be-attended location triggers faster shifts per se compared to a central cue. Besides behavioural data an objective electrophysiological measure, the steady-state visual evoked potential (SSVEP) was used to measure the time course of visual pathway facilitation in the human brain for centrally and peripherally cued shifts of spatial attention. Results revealed that both spatial cues resulted in identical time courses of shifts of covert spatial attention. This was true with respect to behavioural data and SSVEP amplitude. Results support the notion that a salient peripheral spatial cue does not automatically produce faster shifts of spatial attention to the to-be-attended location when this cue is informative and embedded in an ongoing stimulation.     

Competition between auditory and visual spatial cues during visual task performance

There is debate in the crossmodal cueing literature as to whether capture of visual attention by means of sound is a fully automatic process. Recent studies show that when visual attention is endogenously focused sound still captures attention. The current study investigated whether there is interaction between exogenous auditory and visual capture. Participants preformed an orthogonal cueing task, in which, the visual target was preceded by both a peripheral visual and auditory cue. When both cues were presented at chance level, visual and auditory capture was observed. However, when the validity of the visual cue was increased to 80% only visual capture and no auditory capture was observed. Furthermore, a highly predictive (80% valid) auditory cue was not able to prevent visual capture. These results demonstrate that crossmodal auditory capture does not occur when a competing predictive visual event is presented and is therefore not a fully automatic process.