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.     

My eyes want to look where your eyes are looking: exploring the tendency to imitate another individual's gaze

In this study we investigated the tendency of humans to imitate the gaze direction of other individuals. Distracting gaze stimuli or non biological directional cues (arrows) were presented to observers performing an instructed saccadic eye movement task. Eye movement recordings showed that observers performed less accurately when the distracting gaze and the instructed saccade had opposite directions, with a substantial number of saccades matching the direction of the distracting gaze. Static (Experiment 1) and dynamic (Experiment 2) gaze distracters, but not pointing arrows (Experiment 3), produced the effect. Results show a strong predisposition of humans to imitate somebody else's oculomotor behaviour, even when detrimental to task performance. This is likely linked to a strong tendency to share attentional states of other individuals, known as joint attention.     

On how the motor cortices resolve an inter-hemispheric response conflict: an event-related EEG potential-guided TMS study of the flankers task

An important aspect of human motor control is the ability to resolve conflicting response tendencies. Here we used single-pulse transcranial magnetic stimulation (TMS) to track the time course of excitability changes in the primary motor hand areas (M1_HAND) while the motor system resolved response conflicts. Healthy volunteers had to respond fast with their right and left index fingers to right- and left-pointing arrows. These central target stimuli were preceded by flanking arrows, inducing premature response tendencies which competed with correct response activation. The time point of maximum premature activation was individually measured as peak latency of the lateralized readiness potential (LRP) in the EEG. In the subsequent TMS experiment, single pulses were applied to left or right M1_HAND during the same flanker task. The amplitude of the motor evoked potentials in the contralateral first dorsal interosseus muscle was taken as an index of corticospinal excitability. Guided by the previous LRP measurement, magnetic stimuli were applied 0–90 ms after the individual LRP peak, to cover the epoch of conflict resolution. When flankers were incompatible with the target, excitability of the prematurely activated M1_HAND gradually decreased during this 90 ms period. This decrease was paralleled by a mirror-symmetrical increase in excitability in the other M1_HAND. These results show that the inappropriate response tendency is cancelled in one M1_HAND simultaneously with activation of the correct response in the other. This integrated implementation of response activation and cancellation at the level of the M1_HAND presumably represents a downstream effect orchestrated by premotor brain regions.     

Lateralised cortical activity due to preparation of saccades and finger movements: a comparative study

The topography and time course of event-related asymmetries of the EEG associated with horizontal saccadic eye movements and finger movements was compared in a 4-choice response task, where the subjects had to respond to the imperative stimulus (S2) by moving the right or left index finger or by making a saccade to the right or the left. The cue stimulus (S1) contained full, partial, or no information about the direction and the effector. In case of finger movements 3 distinct lateralisations were found: (1) increased negativity over the motor cortex contralateral to the future movement direction, (2) increased contralateral negativity at temporoparietal sites beginning 200 ms after delivery of the directional information, and (3) increased ipsilateral negativity at temporo-parietal sites beginning 350–500 ms after delivered direction and effector information. The early temporo-parietal lateralisation was also visible in case of saccadic eye movements and in case of effector-unspecific directional information. Before saccadic eye movements no other distinct lateralisation could be observed at any recording site. In sum, lateralised cortical activities due to preparation processes for finger movements and due to effector-unspecific processing of directional information for motor preparation by the posterior parietal cortex could be demonstrated, whereas no distinct lateralisation due to preparation for saccadic eye movements was visible.     

Perceptual and motor contributions to performance and ERP components after incorrect motor activation in a flanker reaction task.

OBJECTIVES: The aims of this study are to evaluate contributions of response and perceptual processes to reaction performance in a flanker reaction task, and to investigate whether event-related potential (ERP) component N2 and error negativity N(e) represent similar or functionally distinct cortical mechanisms. METHODS: ERPs, lateralized readiness potentials (LRPs) and reaction performance were measured in a flanker task with arrows as targets and congruent or incongruent flankers. Squares were used as neutral flankers. Target color signaled a response of the hand indicated by (PRO) or against (ANTI) the target arrow's pointing direction. RESULTS: On both PRO and ANTI conditions, performance was facilitated by congruent and impaired by incongruent flankers. In the ERPs on trials with late response errors an N2 was evident before an N(e). In addition, ERPs on correct trials showed an N2 particularly after incongruent flankers on PRO but for each flanker type on ANTI conditions. On incongruent ANTI trials, two successive response conflicts occurred but only a single N2 appeared. CONCLUSIONS: The results indicate that differences in perceptual processing contribute significantly to the flanker effects on task performance and provide further evidence that N2 and N(e) represent different cortical mechanisms. The data also suggest that N2 is not a real-time correlate of incorrect response suppression.     

Modulation of hemispatial neglect by directional and numerical cues in the line bisection task

We investigated the effects of arrows, eye gaze, and digits presented as irrelevant flankers in a line bisection task that was administered to 17 right brain damaged patients with or without left neglect. The rightward bias of neglect patients was selectively modulated by the direction of eye gaze and by the magnitude of two identical digits. The bisection error was shifted contralesionally by leftward-gazing eyes and "small" digits, whereas it was shifted ipsilesionally by rightward-gazing eyes and "large" digits. Therefore, the performance of neglect patients was influenced by task-irrelevant cues whose directional meaning was either explicitly represented (eye gaze) or related to the activation of a spatially oriented mental representation (digits). Regression analyses of the overall performance revealed that size of the rightward bias and error variability were predicted by neglect assessment scores across the entire sample of right brain damaged patients. The increased variability in line bisection performance is consistent with the "indifference zone" theory and it appears to be a subtle but stable marker of neglect.     

Vertical scanning biases and their possible influence on reading direction: celtic wisdom or folly?

The reason people read from top to bottom is unknown, but could be related to brain-mediated directional biases or environmental factors. To learn if there is a brain-mediated directional bias responsible for top-down reading direction, we evaluated the directional scanning in the vertical dimension by using directional letter and face cancellation tasks. Twenty participants were instructed to cancel either target letters or faces using either an up-down or down-up direction, with the stimuli located in left, right, and center hemispace. The results indicated significant differences in completion time between the search direction (up vs. down) and spatial position for the letter cancellation task, with a faster completion time for the bottom-up scan in right space and top-down in left space. Because the left hemisphere primarily attends to contralateral right hemispace our results suggest that, when attending to letter stimuli, the left hemisphere is biased to scan in a proximal to distal (upward) direction. Although the reasons why this is reversed in left hemispace and why we did not see directional biases in the face condition remains unclear, these results do suggest that the direction in which we learn to read is inconsistent with the brain's intrinsic directional bias.     

Influence of perceptual factors on line bisection

In the present study we investigated whether masking/perceptual factors may influence line bisection performance in normal subjects. We carried out two experiments. In Experiment 1 the stimuli consisted of a line flanked at each end by: (1) parallel arrows, (2) oblique arrows, (3) or oblique geometrical figures. Of the two pairs of labels, one pair converged on the line and appeared to mask the flanked portion of the line. The results showed the presence of a bisection bias in the direction opposite to location of masking labels. In Experiment 2 we examined if orientation factors might have contributed to the observed bisection bias. The stimuli consisted of a line flanked by only one pair of oblique geometrical figures. The results showed that bisection bias was always in the direction opposite to label location, independently of the direction pointed out by oblique figures. These findings suggest that masking/perceptual factors may influence bisection performance in normal subjects.     

Common neural substrates for response selection across modalities and mapping paradigms

In many situations, people can only compute one stimulus-to-response mapping at a time, suggesting that response selection constitutes a "central processing bottleneck" in human information processing. Using fMRI, we tested whether common or distinct brain regions were involved in response selection across visual and auditory inputs, and across spatial and nonspatial mapping rules. We isolated brain regions involved in response selection by comparing two conditions that were identical in perceptual input and motor output, but differed in the complexity of the mapping rule. In the visual-manual task of Experiment 1, four vertical lines were positioned from left to right, and subjects pressed one of four keys to report which line was unique in length. In the auditory-manual task of Experiment 2, four tones were presented in succession, and subjects pressed one of four keys to report which tone was unique in duration. For both visual and auditory tasks, the mapping between target position and key position was either spatially compatible or incompatible. In the verbal task of Experiment 3, subjects used nonspatial mappings that were either compatible ("same" if colors matched; "different" if they mismatched) or incompatible (the opposite). Extensive activation overlap was observed across all three experiments for incompatible versus compatible mapping in bilateral parietal and frontal regions. Our results indicate that common neural substrates are involved in response selection across input modalities and across spatial and nonspatial domains of stimulus-to-response mapping, consistent with behavioral evidence that response selection is a central process.     

Can expectancy influence hemispheric asymmetries?

We carried out three experiments with the aim of verifying a critical assumption of Kinsbourne's (Acta Psychol., 33 (1970), 193-201; Attention and Performance V, London: Academic press, (1975), pp. 81-96) 'dynamic' attentional hypothesis of hemispheric asymmetries, namely, that asymmetries arise only when subjects know in advance what type of stimulus and/or cognitive mode they are about to be engaged with. We used a paradigm modified from Posner (J. Exp. Psychol., 109 (1980), 160-174) to study the effects of non-spatial 'cognitive' cueing on hemispheric asymmetries using a lexical decision and a visuo-spatial discrimination task (acute vs. obtuse angles). While we did not find significant overall hemispheric asymmetries with the spatial material, we found a consistent advantage of the left hemisphere in the lexical decision task. In Experiment 2 where the cue was presented in central vision and only the stimuli were lateralised and in Experiment 3 where both cue and stimuli were lateralised to the same hemisphere, the left hemisphere advantage did not interact with the effect of cueing. In contrast, in Experiment 4, where only the cue was lateralised and the stimuli were centrally presented, the left hemisphere advantage in the lexical decision task emerged only following invalid cueing. While the results of Experiments 2 and 3 are not in keeping with Kinsbourne's hypothesis, the result of Experiment 4 shows that some pre-exposural mechanisms may indeed affect the emergence of hemispheric asymmetries. A differential susceptibility in 'disengaging' from the processing mode induced by an invalid cue might represent another interesting example of hemispheric difference.     

The effects of reading-writing direction on the asymmetry of space perception and directional tendencies: a comparison between French and Tunisian children

We compared the influence of reading and writing habits on the asymmetry of space perception and the directional tendencies of French and Tunisian right-handers, aged 5, 7, and 9 years. By comparing two groups of children who use the opposite direction for writing (from left to right for French, from right to left for Arabic), before and after being taught to read in school, we evaluated the impact of writing direction on these asymmetries. A bisection task, a circle-drawing task, and a dot-filling task were used to assess spatial asymmetries and directional tendencies. On the bisection task, a group difference emerged at 9 years, with the French children bisecting the line to the left of the true centre, and the Tunisian children showing no bias. On the circle-drawing task, there was a group difference from 7 years on, as the French children, but not the Tunisian children, used increasing counterclockwise movements. Finally, on the dot-filling task performed with the right hand, the French children filled in significantly more dots when going from left to right from 7 years on, whereas Tunisian children filled in more dots when going from right to left. These results show the impact of basic tendencies in younger children (ipsilateral bias in line bisection, clockwise direction in circle drawing, outward tendency for horizontal displacement in dot filling), as well as the impact of writing direction on spatial asymmetries after learning to read. The results are also discussed in reference to the differences between the two languages, the closeness of the French direction of writing to spontaneous neural-based tendencies, and the influence of learning French at age 8 for the Tunisian children.     

Normal and impaired reflexive orienting of attention after central nonpredictive cues

Recent studies suggest that stimuli with directional meaning can trigger lateral shifts of visuospatial attention when centrally presented as noninformative cues. We investigated covert orienting in healthy participants and in a group of 17 right brain-damaged patients (9 with hemispatial neglect) comparing arrows, eye gaze, and digits as central nonpredictive cues in a detection task. Orienting effects elicited by arrows and eye gaze were overall consistent in healthy participants and in right brain-damaged patients, whereas digit cues were ineffective. Moreover, patients with neglect showed, at the shortest delay between cue and target, a disengage deficit for arrow cueing whose magnitude was predicted by neglect severity. We conclude that the peculiar form of attentional orienting triggered by the directional meaning of arrow cues presents some features previously thought to characterize only the stimulus-driven (exogenous) orienting to noninformative peripheral cues.     

Egocentric frame of reference: its role in spatial bias after right hemisphere lesions.

The reference shift hypothesis of unilateral neglect holds that spatial bias in left neglect stems from a rightward deviation of patients' egocentric frame of reference (ER). Twenty five unselected right brain-damaged patients participated in a straight-ahead pointing task to assess the position of their ER (Experiment 1). A rightward ER shift emerged only in the subgroup of patients with extensive parietal lesions. In Experiment 2, we found that the position of the ER did not predict the outcome of various visuospatial neglect tests (r = 0.07 to 0.27). In Experiment 3, no significant positive correlation emerged between the ER position and visual (r = 0.26) or tactile (r = -0.48) extinction. Two further experiments examined the relationships between the ER position and patients' performance on a reaction time test of directional motor bias (Experiment 4), and on a test of response times to lateralised visual stimuli (Experiment 5). Results showed that the ER position did not predict the distribution of accuracy scores or response times in either task (Experiment 3: accuracy: r = 0.06; response times: r = 0.16; Experiment 4: accuracy: r = 0.09; response times: r = 0.04). We concluded that the position of the ER plays no crucial role in the behavioural consequences of spatial bias induced by right hemisphere lesions.     

Utilitarian Attention by Children with Autism Spectrum Disorder on a Filtering Task

The findings are evidence that persons with ASD benefit more than typically developing (TD) persons from spatial framing cues in focusing their attention on a visual target. Participants were administered a forced-choice task to assess visual filtering. A target stimulus was presented on a screen and flanker stimuli were presented simultaneously with or after the target, with varying stimuli onset asynchronies (SOAs). Regardless of SOA, TD children showed the expected distracting effects with slower reaction times (RTs) when flankers were at closer distances from the target. However, children with ASD displayed shorter RTs in the conditions in which the stimuli were presented simultaneously or with a short SOA. These findings are interpreted as reflecting utilitarian attention among children with ASD.     

Responses of hippocampal formation neurons in the monkey related to delayed spatial response and object-place memory tasks

The memory for where in the environment a particular visual stimulus has been seen is one of the types of memory relatively specifically impaired by hippocampal damage in primates including man. In order to investigate what processing might be performed by the hippocampus related to this type of memory, the activity of hippocampal neurons was recorded while monkeys performed an object-place memory task. In this task, the monkey was shown a sample stimulus in one position on a video screen, there was a delay of 2 s, and then the same or a different stimulus was shown in the same or in a different position. The monkey remembered the sample and its position, and if both matched the delayed stimulus, he licked to obtain fruit juice. Of the 600 neurons analysed in this task, 3.8% responded differently for the different spatial positions, with some of these responding differentially during the sample presentation, some in the delay period, and some in the match period. Thus some hippocampal neurons respond differently for stimuli shown in different positions in space, and some respond differently when the monkey is remembering different positions in space. In addition some of the neurons responded to a combination of object and place information, in that they responded only to a novel object in a particular place. These neuronal responses were not due to any response being made or prepared by the monkey, for information about which behavioral response was required was not available until the match stimulus was shown. This is the first demonstration that some hippocampal neurons in the primate have activity related to the spatial position of stimuli. The activity of these neurons was also measured in a delayed spatial response task, in which the monkey was shown a stimulus in one position, and, after a 2 s delay when two identical stimuli were shown, had to reach to touch the stimulus which was in the position in which it had previously been seen. It was found that the majority of the neurons which responded in the object-place memory task did not respond in the delayed response task. Instead, a different population of neurons (5.7% of the total) responded in the delayed spatial response task, with differential left-right responses in the sample, delay, or match periods.(ABSTRACT TRUNCATED AT 400 WORDS)     

Prior information of stimulus location: effects on ERP measures of visual selection and response selection

This paper examines the effects of prior information of the location of an upcoming stimulus on event-related EEG potentials associated with the focusing of attention. Results of two tasks, reported in a previous publication (Praamstra, P., Boutsen, L., Humphreys, G.W., 2005. Frontoparietal control of spatial attention and motor intention in human EEG. J. Neurophysiol. 94, 764-774), were compared: one in which spatial attention was cued to the stimulus location and one in which the cue was non-informative. Only informative directional cues elicited directing-attention EEG potentials in the delay period between cue and target. Notwithstanding these electrophysiological signs of an attentional orientation prior to the occurrence of the target, there were no reaction time effects related to the presence of advance spatial information. By contrast, the advance information did have effects on EEG potentials following the target stimulus. The N2pc, reflecting an attentional selection mechanism in extrastriate cortex, was reduced in amplitude with advance spatial information. The N2cc, coinciding in time with the N2pc but measured over the motor cortex, was preempted by the advance spatial information. These results support that the N2cc is not due to overlap of the N2pc with movement execution-related activity. It is proposed that the neural activity underlying this EEG potential arises from the dorsal premotor cortex and serves an executive-attentional function that helps to ensure that the selection of a manual response is not biased by the direction of spatial attention.     

Mu rhythm,visual processing and motor control

ObjectiveThe Mu-rhythm has been proposed as both an inhibitor (“idling hypothesis”) and as a promoter (“processing hypothesis”) of information processing in the primary sensory-motor cortex (SM-C). We tested these possibilities by analyzing the phase-locked and non-phase-locked Mu response during the execution of a visual–motor task.MethodsEEG was recorded in 13 subjects during the visual presentation of an arrow which indicated the direction of the finger motion to be executed after the presentation of a second stimulus. The EEG activity in the α-range (Mu-α) and β-range was evaluated by a method which segregated the phase-locked and the non-phase-locked response. The event-related Mu-response observed during this task was compared with that computed when the subjects saw the same arrow-stimuli but did not perform any task (passive test).ResultsVisual stimuli induced a phase-locked α-oscillation which began ≈50 ms after the stimulus onset and persisted for about 150–200 ms. This response was much higher when stimuli were used for motion planning than when they were passively observed, and was more marked in the α-range than in the β-range. The phase-locked response was followed by a persistent decrease of the non-phase-locked Mu-activity similar to that previously reported with the event-related desynchronization/synchronization method.ConclusionsThe Mu-wave is not a single phenomenon. It was segregated here into two components, one with an early and short-lasting phase locked-response to visual stimuli, which increased during the task execution, and the other without phase-locked responses which persistently decreased during the task execution.SignificancePresent data suggest that Mu-activity performs a double action, increasing the information processing of one task (according to the “processing hypothesis”) and decreasing the computation of other potentially interfering tasks (according to the “idling hypothesis”), with task selection being achieved by choosing their phase-association to the Mu-wave.     

The anterior N1 component as an index of modality shifting

Processing of a given target is facilitated when it is defined within the same (e.g., visual-visual), compared to a different (e.g., tactile-visual), perceptual modality as on the previous trial [Spence, C., Nicholls, M., & Driver, J. The cost of expecting events in the wrong sensory modality. Perception & Psychophysics, 63, 330-336, 2001]. The present study was designed to identify electrocortical (EEG) correlates underlying this "modality shift effect." Participants had to discriminate (via foot pedal responses) the modality of the target stimulus, visual versus tactile (Experiment 1), or respond based on the target-defining features (Experiment 2). Thus, modality changes were associated with response changes in Experiment 1, but dissociated in Experiment 2. Both experiments confirmed previous behavioral findings with slower discrimination times for modality change, relative to repetition, trials. Independently of the target-defining modality, spatial stimulus characteristics, and the motor response, this effect was mirrored by enhanced amplitudes of the anterior N1 component. These findings are explained in terms of a generalized "modality-weighting" account, which extends the "dimension-weighting" account proposed by Found and Müller [Searching for unknown feature targets on more than one dimension: Investigating a "dimension-weighting" account. Perception & Psychophysics, 58, 88-101, 1996] for the visual modality. On this account, the anterior N1 enhancement is assumed to reflect the detection of a modality change and initiation of the readjustment of attentional weight-setting from the old to the new target-defining modality in order to optimize target detection.     

Auditory-somatosensory multisensory interactions in humans: Dissociating detection and spatial discrimination

Simple reaction times (RTs) to auditory-somatosensory (AS) multisensory stimuli are facilitated over their unisensory counterparts both when stimuli are delivered to the same location and when separated. In two experiments we addressed the possibility that top-down and/or task-related influences can dynamically impact the spatial representations mediating these effects and the extent to which multisensory facilitation will be observed. Participants performed a simple detection task in response to auditory, somatosensory, or simultaneous AS stimuli that in turn were either spatially aligned or misaligned by lateralizing the stimuli. Additionally, we also informed the participants that they would be retrogradely queried (one-third of trials) regarding the side where a given stimulus in a given sensory modality was presented. In this way, we sought to have participants attending to all possible spatial locations and sensory modalities, while nonetheless having them perform a simple detection task. Experiment 1 provided no cues prior to stimulus delivery. Experiment 2 included spatially uninformative cues (50% of trials). In both experiments, multisensory conditions significantly facilitated detection RTs with no evidence for differences according to spatial alignment (though general benefits of cuing were observed in Experiment 2). Facilitated detection occurs even when attending to spatial information. Performance with probes, quantified using sensitivity (d′), was impaired following multisensory trials in general and significantly more so following misaligned multisensory trials. This indicates that spatial information is not available, despite being task-relevant. The collective results support a model wherein early AS interactions may result in a loss of spatial acuity for unisensory information.     

The effect of Parkinson's disease on interference control during action selection

Basal ganglia structures comprise a portion of the neural circuitry that is hypothesized to coordinate the selection and suppression of competing responses. Parkinson's disease (PD) may produce a dysfunction in these structures that alters this capacity, making it difficult for patients with PD to suppress interference arising from the automatic activation of salient or overlearned responses. Empirical observations thus far have confirmed this assumption in some studies, but not in others, due presumably to considerable inter-individual variability among PD patients. In an attempt to help resolve this controversy, we measured the performance of 50 PD patients and 25 healthy controls on an arrow version of the Eriksen flanker task in which participants were required to select a response based on the direction of a target arrow that was flanked by arrows pointing in the same (congruent) or opposite (incongruent) direction. Consistent with previous findings, reaction time (RT) increased with incongruent flankers compared to congruent or neutral flankers, and this cost of incongruence was greater among PD patients. Two novel findings are reported. First, distributional analyses, guided by dual-process models of conflict effects and the activation-suppression hypothesis, revealed that PD patients are less efficient at suppressing the activation of conflicting responses, even when matched to healthy controls on RT in a neutral condition. Second, this reduced efficiency was apparent in half of the PD patients, whereas the remaining patients were as efficient as healthy controls. These findings suggest that although poor suppression of conflicting responses is an important feature of PD, it is not evident in all medicated patients.