Central Arrow Research Articles

Effects of working memory load on visual selective attention: behavioral and electrophysiological evidence.

Working memory and attention interact in a way that enables us to focus on relevant items and maintain current goals. The influence of working memory on attention has been noted in several studies using dual task designs. Multitasking increases the demands on working memory and reduces the amount of resources available for cognitive control functions such as resolving stimulus conflict. However, few studies have investigated the temporal activation of the cortex while multitasking. The present study addresses the extent to which working memory load influences early (P1) and late (P300) attention-sensitive event-related potential components using a dual task paradigm. Participants performed an arrow flanker task alone (single task condition) or concurrently with a Sternberg memory task (dual task condition). In the flanker task, participants responded to the direction of a central arrow surrounded by congruent or incongruent arrows. In the dual task condition, participants were presented with a Sternberg task that consisted of either four or seven consonants to remember prior to a short block of flanker trials. Participants were slower and less accurate on incongruent versus congruent trials. Furthermore, accuracy on incongruent trials was reduced in both dual task conditions. Likewise, P300 amplitude to incongruent flanker stimuli decreased when working memory load increased. These findings suggest that interference from incongruent flankers was more difficult to suppress when working memory was taxed. In addition, P1 amplitude was diminished on all flanker trials in the dual task condition. This result indicates that top-down attentional control over early visual processing is diminished by increasing demands on working memory. Both the behavioral and electrophysiological results suggest that working memory is critical in maintaining attentional focus and resolving conflict.

Testing the attention shift hypothesis as an account for the flanker sequence-based congruency modulation in spatial flanker tasks

Smaller Simon effects when stimulus locations are repeated on successive trials rather than alternated have been explained by the attention shift hypothesis, suggesting that shifts of attention result in interfering response codes. We investigated whether the attention shift hypothesis can also explain smaller flanker effects for repeated flankers than for alternated flankers, which occur only with directional information. In 3 peripheral letter identification tasks, target locations were cued by partial or complete flanker stimuli. Experiments 1 and 2 showed that directional flankers elicit shifts of attention. However, Experiment 3 revealed that directional flankers induced inverted cuing effects when reacting to the central target arrow was additionally required. These results are difficult to reconcile with the attention shift hypothesis as an explanation for the congruency reduction with repetitions of directional flankers.

Individual Differences in Subconscious Motor Control Predicted by GABA Concentration in SMA

Subliminal visual stimuli affect motor planning, but the size of such effects differs greatly between individuals. Here, we investigated whether such variation may be related to neurochemical differences between people. Cortical responsiveness is expected to be lower under the influence of more of the main inhibitory neurotransmitter, GABA. Thus, we hypothesized that, if an individual has more GABA in the supplementary motor area (SMA)--a region previously associated with automatic motor control--this would result in smaller subliminal effects. We measured the reversed masked prime--or negative compatibility--effect, and found that it correlated strongly with GABA concentration, measured with magnetic resonance spectroscopy. This occurred specifically in the SMA region, and not in other regions from which spectroscopy measurements were taken. We replicated these results in an independent cohort: more GABA in the SMA region is reliably associated with smaller effect size. These findings suggest that, across individuals, the responsiveness of subconscious motor mechanisms is related to GABA concentration in the SMA.

Bottom-up effects modulate saccadic latencies in well-known eye movement paradigm

A well-known eye movement paradigm combines saccades (fast eye movements) with a perceptual discrimination task. At a variable time after the onset of a central arrow cue indicating the target direction [the stimulus onset asynchrony (SOA)], discrimination symbols appear briefly at saccade target and non-target locations. A previous study revealed an unexpected effect of SOA on saccadic latencies: latencies were longer in trials with longer SOAs. It was suggested that this effect reflects a top-down process as observers may wait for the discrimination symbol to appear before executing saccades. However, symbol onsets may also modulate saccade latencies from the bottom-up. To clarify the origin of the SOA effect on latencies in this paradigm, we used a simplified version of the original task plus two new symbol onset conditions for comparison. The results indicate that the modulation of saccadic latencies was not due to a top-down strategy, but to a combination of two opposing bottom-up effects: the symbol onsets at the target location shortened saccade latencies, while symbol onsets at non-target locations lengthened saccade latencies.

Is object-based attention mandatory? Strategic control over mode of attention.

Is object-based attention mandatory or under strategic control? In an adapted spatial cuing paradigm, participants focused initially on a central arrow cue that was part of a perceptual group (Experiment 1) or a uniformly connected object (Experiment 2), encompassing one of the potential target locations. The cue always pointed to an opposite, different-object location. By varying cue validity, the strategic incentive to prevent the spread of attention to the entire cue object, and consequently to the same-object location, was manipulated: With invalid cuing and (consequently) equal probability of targets at same-object and different-object locations, a same-object target identification advantage was observed. With highly valid cuing and targets much more probable at the different-object location than at the same-object location, the same-object advantage disappeared. Object-based attention appears to be a default mode that may be ecologically adaptive but can be overridden by strategic control when there is a strong immediate benefit in doing so.

Response Competition in the Primary Motor Cortex: Corticospinal Excitability Reflects Response Replacement During Simple Decisions

It has been suggested that, during decisions about actions, multiple options are initially specified in parallel and then gradually eliminated in a competition for overt execution. To further test this hypothesis, we studied the modulation of human corticospinal excitability during the reaction time of the Eriksen flanker task. In the task, subjects responded with finger flexion or extension to a central arrow while ignoring congruent or incongruent flanker arrows. Single-pulse transcranial magnetic stimulation (TMS) was applied over primary motor cortex (M1) at one of five different latencies after stimulus onset, and motor-evoked potentials (MEPs) were measured in the contralateral index finger. During the control (no flankers) and congruent conditions, MEP size in the agonist increased gradually over the course of reaction time, indicating an increase in corticospinal excitability. Conversely, when the same muscle acted as an antagonist, MEP size decreased, suggesting inhibition. Critically, in the incongruent condition, MEPs briefly increased in the muscle corresponding to an initial default response to the flanker arrows and were later replaced by MEPs corresponding to the correct response to the central arrow. Finally, we found that the gradually growing MEPs for the three conditions reached a constant maximum level just before movement initiation. We propose that this dynamic modulation in corticospinal excitability reflects the competition process, leading to the selection of one response and the rejection of the other. Our results suggest that response competition influences activity in primary motor cortex and that its timing directly influences motor output latency.

'reading' dynamic facial expression in autistic spectrum disorder

Autistic Spectrum Disorder (ASD) affects about 1 in 200 individuals and is associated with deficits in social communication and reciprocity. These abilities are highly dependent upon the ability to understand others' intentions and desires by ‘reading’ facial expressions and eye gaze direction. Instruments developed to diagnose ASD find that the ability to follow the direction of other's eye gaze, is impaired in ASD and discriminates between these children and those with other disorders. However, experiments have found that children with ASD reduce their reaction times to a two-choice stimulus, by taking advantage of a precue in the form of an eye gaze stimulus. These studies may be problematic as they employed static images of eyes that were removed from the face, and which could be treated as symbolic stimuli. In this study, 10 children with ASD and an age-matched control group were presented with a realistic computer generated face (composite of 100 real faces) before the presentation of an imperative stimulus (an arrow). Participants were required to press a right key when a central arrow pointed right and a left key when it pointed left. A 10 degree eye-movement within the face seen for 1000ms provided consistently valid precue information. The imperative stimuli would appear 500–3000ms (random) after the face disappeared. In a second block of trials, the eye shift occurred concurrently with a smile or a frown. The smile meant that the cue was valid but a frown indicated an invalid cue. This design feature was explained carefully to the children. This experimental paradigm provides an objective and quantifiable measure of whether children with or without ASD can integrate facial expression and eye-gaze to derive intentional information.

Iconic Decay in Schizophrenia

Working memory impairment is considered a core deficit in schizophrenia, but the precise nature of this deficit has not been determined. Multiple lines of evidence implicate deficits at the encoding stage. During encoding, information is held in a precategorical sensory store termed iconic memory, a literal image of the stimulus with high capacity but rapid decay. Pathologically increased iconic decay could reduce the number of items that can be transferred into working memory before the information is lost and could thus contribute to the working memory deficit seen in the illness. The current study used a partial report procedure to test the hypothesis that patients with schizophrenia (n = 37) display faster iconic memory decay than matched healthy control participants (n = 28). Six letters, arranged in a circle, were presented for 50 ms. Following a variable delay of 0-1000 ms, a central arrow cue indicated the item to be reported. In both patients and control subjects, recall accuracy decreased with increasing cue delay, reflecting decay of the iconic representation of the stimulus array. Patients displayed impaired memory performance across all cue delays, consistent with an impairment in working memory, but the rate of iconic memory decay did not differ between patients and controls. This provides clear evidence against faster loss of iconic memory representations in schizophrenia, ruling out iconic decay as an underlying source of the working memory impairment in this population. Thus, iconic decay rate can be added to a growing list of unimpaired cognitive building blocks in schizophrenia.

Endogenous cueing attenuates object substitution masking

Object substitution masking (OSM) is a form of visual masking in which a briefly presented target surrounded by four small dots is masked by the continuing presence of the four dots after target offset. A major parameter in the prediction of OSM is the time required for attention to be directed to the target following its onset. Object substitution theory (Di Lollo et al. in J Exp Psychol Gen 129:481-507, 2000) predicts that the sooner attention can be focused at the target's location, the less masking will ensue. However, recently Luiga and Bachmann (Psychol Res 71:634-640, 2007) presented evidence that precueing of attention to the target location prior to target-plus-mask onset by means of a central (endogenous) arrow cue does not reduce OSM. When attention was cued exogenously, OSM was attenuated. Based on these results, Luiga and Bachmann argued that object substitution theory should be adapted by differentiating the ways of directing attention to the target location. The goal of the present study was to further examine the dissociation between the effects of endogenous and exogenous precueing on OSM. Contrary to Luiga and Bachmann, our results show that prior shifts of attention to the target location initiated by both exogenous and endogenous cues reduce OSM as predicted by object substitution theory and its computational model CMOS.

Involuntary but not voluntary orienting contributes to a disengage deficit in visual neglect

Patients with left neglect are particularly slow to respond to visual targets on their left when attention is first engaged to their right. This deficit is known as the disengage deficit (DD). Studies investigating the DD typically employ nonpredictive peripheral onset cues to measure involuntary orienting and predictive central arrow cues to measure voluntary orienting. A DD has been observed with both cues, suggesting that a DD occurs for involuntary and for voluntary orienting. Recent evidence questions this conclusion because nonpredictive central arrow cues trigger involuntary orienting. This implies that predictive central arrows also involve involuntary orienting and do not measure only voluntary attention. This new knowledge suggests a new conceptualization of the DD. While it is undisputed that a DD occurs when attention is shifted involuntarily, it is uncertain whether a DD is produced by voluntary orienting because most previous cuing studies of the DD have involved shifts of involuntary attention. To address this critical question, we tested neglect and control patients with nonpredictive and predictive peripheral onset cues ( Experiment 1), nonpredictive and predictive central arrow cues ( Experiment 2), and predictive central number cues ( Experiment 3). The experiments provide three lines of converging evidence that voluntary orienting does not contribute to a DD. First, the DD was the same whether attention was engaged involuntarily by nonpredictive peripheral cues or engaged involuntarily and voluntarily by predictive peripheral cues ( Experiment 1), indicating that voluntary orienting does not modulate the DD. Second, the DD was the same whether attention was engaged involuntarily by nonpredictive central arrow cues or engaged involuntarily and voluntarily by predictive central arrow cues ( Experiment 2), replicating the finding of Experiment 1 with very different cues. Third, the DD was not present when attention was only engaged voluntarily by central predictive number cues ( Experiment 3).

The neural correlates of social attention: automatic orienting to social and nonsocial cues

Previous evidence suggests that directional social cues (e.g., eye gaze) cause automatic shifts in attention toward gaze direction. It has been proposed that automatic attentional orienting driven by social cues (social orienting) involves a different neural network from automatic orienting driven by nonsocial cues. However, previous neuroimaging studies on social orienting have only compared gaze cues to symbolic cues, which typically engage top-down mechanisms. Therefore, we directly compared the neural activity involved in social orienting to that involved in purely automatic nonsocial orienting. Twenty participants performed a spatial cueing task consisting of social (gaze) cues and automatic nonsocial (peripheral squares) cues presented at short and long stimulus (cue-to-target) onset asynchronies (SOA), while undergoing fMRI. Behaviorally, a facilitation effect was found for both cue types at the short SOA, while an inhibitory effect (inhibition of return: IOR) was found only for nonsocial cues at the long SOA. Imaging results demonstrated that social and nonsocial cues recruited a largely overlapping fronto-parietal network. In addition, social cueing evoked greater activity in occipito-temporal regions at both SOAs, while nonsocial cueing recruited greater subcortical activity, but only for the long SOA (when IOR was found). A control experiment, including central arrow cues, confirmed that the occipito-temporal activity was at least in part due to the social nature of the cue and not simply to the location of presentation (central vs. peripheral). These results suggest an evolutionary trajectory for automatic orienting, from predominantly subcortical mechanisms for nonsocial orienting to predominantly cortical mechanisms for social orienting.

Increased effect of target eccentricity on covert shifts of visual attention in patients with neglect

Debate continues regarding the mechanisms underlying covert shifts of visual attention. We examined the relationship between target eccentricity and the speed of covert shifts of attention in normal subjects and patients with brain lesions using a cued-response task in which cues and targets were presented at 2° or 8° lateral to the fixation point. Normal subjects were slower on invalid trials in the 8° as compared to 2° condition. Patients with right-hemisphere stroke with neglect were slower in their responses to left-sided invalid targets compared to valid targets, and demonstrated a significant increase in the effect of target validity as a function of target eccentricity. Additional data from one neglect patient (JM) demonstrated an exaggerated validity × eccentricity × side interaction for contralesional targets on a cued reaction time task with a central (arrow) cue. We frame these results in the context of a continuous ‘moving spotlight’ model of attention, and also consider the potential role of spatial saliency maps. By either account, we argue that neglect is characterized by an eccentricity-dependent deficit in the allocation of attention.

Visuospatial attention and redundancy gain

Two experiments investigated the effect of visuospatial attention on redundancy gain in simple reaction time tasks. In each trial participants were given a central arrow cue indicating where a stimulus would most likely be presented (i.e., upper or lower half of the display in Experiment 1; left or right half of the display in Experiment 2). Then, a single stimulus or two redundant stimuli could be presented in either expected or unexpected locations. Replicating previous findings, responses were faster when stimuli appeared in expected rather than unexpected locations, and they were also faster when two redundant stimuli were presented than when only one was. Critically, redundancy gain was statistically equivalent for stimuli in expected and unexpected locations, suggesting that the effect of redundancy gain arises after the perceptual processes influenced by the allocation of visuospatial attention.

Attending to objects: Endogenous cues can produce inhibition of return

Inhibition of Return (IOR) is effective in a wide range of experimental settings but has proven elusive under conditions of volitional (endogenous) attentional control. This result may be due to a continuing attentional bias towards the cued location. Here we ask whether IOR can be unmasked under endogenous cueing conditions when a predictive cue prevents such a bias. In Experiment 1, a central arrow directed attention endogenously to one end of a rectangular object. Attention initially radiated to an unpredicted location in the object but IOR later impaired target detection at that location. In Experiment 2, attention was directed endogenously to a dynamic object. While target detection was facilitated at a subsequent (and predicted) location of the rotating object, detection-latencies at its original location were slowed. Together the results show that it is possible to produce IOR using endogenous cues.

Attentional modulation of motion-induced blindness

When a global moving pattern is superimposed on high-contrast stationary or slowly moving stimuli, the stimuli can be perceived as disappearing and reappearing alternately for periods of several seconds. This visual illusory phenomenon was named “motion-induced blindness” (MIB) in recent literature. So far there is no consensus on the mechanism of MIB, especially on the role of attention in this phenomenon. To examine the effect of spatial attention on MIB, the present study manipulated the participants’ spatial attention by asking them to respond to two targets simultaneously presented in bilateral visual fields (the divided-attention condition) or only respond to one of them (the focused-attention condition). A central arrow was presented as an endogenous cue to index the target visual field in the focused-attention condition, while a point was presented instead in the divided-attention condition. The results show that the percentage of accumulated invisibility period was larger for the targets in the focused-attention condition than for those in the divided-attention condition. This effect of attention is significant in upper visual field (UVF) and left lower visual field (left LVF); that is, this effect shows a hemispheric asymmetry in LVF but not in UVF. Furthermore, the percentage of accumulated invisibility period was larger for targets in left LVF than for those in right LVF in the focused-attention condition, but no hemispheric asymmetry was found in the divided-attention condition. In addition, the increased percentage of accumulated invisibility period in the focused-attention condition originated merely in the enhancement of the mean phase duration of disappearance in LVF, while the disappearance occurred more frequently and lasted longer for each occurrence, which led to an increase in the total invisibility period, in the focused-than divided-attention condition in UVF. These results suggest that the modulation of spatial attention on MIB has different patterns in UVF and LVF.

Unilateral Amygdala Lesions Hamper Attentional Orienting Triggered by Gaze Direction

The newly discovered deficit in a bilateral amygdala-damaged case, of not being able to allocate attention to the critical feature of a face (Adolphs R, Gosselin F, Buchanan TW, Tranel D, Schyns P, Damasio AR. 2005. A mechanism for impaired fear recognition after amygdala damage. Nature. 433:68--72.), has opened a new window into the function of the amygdala. This case implies that the amygdala might be essential in detecting potentially relevant social stimuli, and directing attention accordingly. In this study, we have sought to test this implication by investigating the behavioral performance of 5 unilateral amygdala-damaged subjects on spatial cueing tasks. The tasks employed central gaze and arrow direction as cues to trigger attentional orienting in peripheral target detection. Although age-matched normal controls demonstrated a significant congruency effect such that targets presented congruently to cue direction elicited faster detection, amygdala subjects demonstrated no such congruency effect for gaze cues in the face of a significant congruency effect for arrow cues. The results suggest that the social valence of a stimulus is critical for amygdala involvement in visual processing. The results also support the implicated role of the amygdala in detecting and analyzing relevant social stimuli, and orienting attention accordingly.

Attention to Arrows: Pointing to a New Direction

It was long believed that central arrows needed to be spatially predictive to produce a shift in spatial attention. Recent evidence indicates, however, that central spatially nonpredictive directional cues, like arrows, will trigger reflexive shifts in attention. We asked what this recent discovery means for past studies that used predictive directional cues such as arrows. Our findings indicate that predictive arrows produce attention effects that greatly exceed the individual or summed effects of reflexive orienting to nonpredictive arrows and volitional orienting to predictive numbers. This suggests that the especially large effect produced by predictive arrows reflects an interaction between reflexive and volitional orienting. Given the broad application of the predictive arrow cueing paradigm in both past and current research, the present data shed new light on a wide range of investigations, from psychophysical studies of basic attention to behavioural and neuroimaging studies of cognition and social development.

Capturing focused attention

When attention is diffuse, as in a visual search task, an abrupt onset almost invariably succeeds in capturing attention. But if attention had been cued in advance to a different location, the same onset may then fail to capture attention (Theeuwes, 1991; Yantis and Jonides, 1990). In previous demonstrations, the onset appeared frequently. This may have diminished the onset's novelty, which in turn could have affected its potency in capturing attention. The question we asked was whether preserving the onset's novelty could enhance its capacity in capturing attention, even when attention had been prioritized elsewhere. As in Theeuwes's (1991) study, observers were cued to the target location with a 100%-predictive central arrow cue. The frequency with which visual transients were introduced was varied across experiments. When the onset stimulus appeared frequently (Experiment 1), it indeed failed to capture attention. But when its appearance was relatively infrequent (Experiment 2), capture effects were clearly observed. In Experiment 3, the target appeared in one location throughout the experiment. This would have improved target localization, but an infrequently appearing onset stimulus still successfully captured attention.

The number line effect reflects top-down control

Recent evidence indicates that central directional stimuli, such as eyes and arrows, trigger rapid, reflexive shifts of spatial attention. A study by Fischer, Castel, Dodd, and Pratt (2003) suggested that a similar effect might also apply to central numbers, as if a digit's meaning causes attention to be oriented to its relative position on a left-to-right mental number line. However, unlike central eyes and arrows, the orienting effect for central digits emerges slowly, suggesting that top-down endogenous processes may be mediating this effect. Here, we report a series of three experiments that strongly support this hypothesis. Experiment 1 replicated Fischer et al.'s left-to-right number line effect. Experiment 2 showed that this effect could be completely reversed by merely asking participants to imagine a number line running from right to left. Experiment 3 showed that a left-to-right number line effect could be abolished by presenting targets above and below central fixation, as well as to the left and right of center. Experiment 3 also showed that other mental sets, such as imagining a clock, result in attention's being oriented in accordance with where the central digits are represented on a clock face. Together, these data indicate that the spatial representations and attentional orienting related to the perception of digits are both fragile and flexible and depend critically on the top-down spatial mental sets adopted by individuals.

Context Modulates Early Stimulus Processing when Resolving Stimulus-response Conflict

When responding to stimuli in our environment, the presence of multiple items associated with task-relevant responses affects both ongoing response selection and subsequent behavior. Computational modeling of conflict monitoring and neuroimaging data predict that the recent context of response competition will bias the selection of certain stimuli over others very early in the processing stream through increased focal spatial attention. We used high-density EEG to test this hypothesis and to investigate the contextual effects on nonspatial, early stimulus processing in a modified flanker task. Subjects were required to respond to a central arrow and to ignore potentially conflicting information from flanking arrows in trials preceded by a series of either compatible or incompatible trials. On some trials, we presented the flanking arrows in the absence of the central target. The visual P1 component was selectively enhanced only for incompatible trials when preceded by incompatible ones, suggesting that contextual effects depend on feature-based processing, and not only simple enhancement of the target location. Context effects also occurred on no-target trials as evidenced by an enhanced early-evoked response when they followed compatible compared to incompatible trials, suggesting that spatial attention was also modulated by recent context. These results support a multi-componential account of spatial and nonspatial attention and they suggest that contextually driven cognitive control mechanisms can operate on specific stimulus features at extremely early stages of processing within stimulus-response conflict tasks.