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1.  On the temporal dynamics of spatial stimulus-response transfer between spatial incompatibility and Simon tasks 
The Simon effect refers to the performance (response time and accuracy) advantage for responses that spatially correspond to the task-irrelevant location of a stimulus. It has been attributed to a natural tendency to respond toward the source of stimulation. When location is task-relevant, however, and responses are intentionally directed away (incompatible) or toward (compatible) the source of the stimulation, there is also an advantage for spatially compatible responses over spatially incompatible responses. Interestingly, a number of studies have demonstrated a reversed, or reduced, Simon effect following practice with a spatial incompatibility task. One interpretation of this finding is that practicing a spatial incompatibility task disables the natural tendency to respond toward stimuli. Here, the temporal dynamics of this stimulus-response (S-R) transfer were explored with speed-accuracy trade-offs (SATs). All experiments used the mixed-task paradigm in which Simon and spatial compatibility/incompatibility tasks were interleaved across blocks of trials. In general, bidirectional S-R transfer was observed: while the spatial incompatibility task had an influence on the Simon effect, the task-relevant S-R mapping of the Simon task also had a small impact on congruency effects within the spatial compatibility and incompatibility tasks. These effects were generally greater when the task contexts were similar. Moreover, the SAT analysis of performance in the Simon task demonstrated that the tendency to respond to the location of the stimulus was not eliminated because of the spatial incompatibility task. Rather, S-R transfer from the spatial incompatibility task appeared to partially mask the natural tendency to respond to the source of stimulation with a conflicting inclination to respond away from it. These findings support the use of SAT methodology to quantitatively describe rapid response tendencies.
doi:10.3389/fnins.2014.00243
PMCID: PMC4137233  PMID: 25191217
speed-accuracy trade-off; stimulus-response compatibility; Simon effect; spatial compatibility; S-R associations
2.  Age Differences in Response Selection for Pure and Mixed Stimulus-Response Mappings and Tasks 
Acta psychologica  2008;129(1):49-60.
Two experiments examined effects of mixed stimulus-response mappings and tasks for older and younger adults. In Experiment 1, participants performed two-choice spatial reaction tasks with blocks of pure and mixed compatible and incompatible mappings. In Experiment 2, a compatible or incompatible mapping was mixed with a Simon task for which the mapping of stimulus color to location was relevant and stimulus location irrelevant. In both experiments older adults showed larger mixing costs than younger adults and larger compatibility effects, with the differences particularly pronounced in Experiment 1 when location mappings were mixed. In mixed conditions, when stimulus location was relevant, older adults benefited more than younger adults from complete repetition of the task and stimulus from the preceding trial. When stimulus location was irrelevant, the benefit of complete repetition did not differ reliably between age groups. The results suggest that the age-related deficit associated with mixing mappings and tasks is primarily due to older adults having more difficulty separating task sets that activate conflicting response codes.
doi:10.1016/j.actpsy.2008.04.006
PMCID: PMC2608201  PMID: 18541219
Aging; Attention; Response Selection; 2860 Gerontology; 2346 Attention
3.  Endogenous orienting modulates the Simon effect: critical factors in experimental design 
Psychological Research  2007;72(3):261-272.
Responses are faster when the side of stimulus and response correspond than when they do not correspond, even if stimulus location is irrelevant to the task at hand: the correspondence, spatial compatibility effect, or Simon effect. Generally, it is assumed that an automatically generated spatial code is responsible for this effect, but the precise mechanism underlying the formation of this code is still under dispute. Two major alternatives have been proposed: the referential-coding account, which can be subdivided into a static version and an attention-centered version, and the attention-shift account. These accounts hold clear-cut predictions for attentional cuing experiments. The former would assume a Simon effect irrespective of attentional cuing in its static version, whereas the attention-centered version of the referential-coding account and the attention-shift account would predict a decreased Simon effect on validly as opposed to invalidly cued trials. However, results from previous studies are equivocal to the effects of attentional cuing on the Simon effect. We argue here that attentional cueing reliably modulates the Simon effect if some crucial experimental conditions, mostly relevant for optimizing attentional allocation, are met. Furthermore, we propose that the Simon effect may be better understood within the perspective of supra-modal spatial attention, thereby providing an explanation for observed discrepancies in the literature.
doi:10.1007/s00426-007-0110-x
PMCID: PMC2668658  PMID: 17351789
4.  A New Perceptual Bias Reveals Suboptimal Population Decoding of Sensory Responses 
PLoS Computational Biology  2012;8(4):e1002453.
Several studies have reported optimal population decoding of sensory responses in two-alternative visual discrimination tasks. Such decoding involves integrating noisy neural responses into a more reliable representation of the likelihood that the stimuli under consideration evoked the observed responses. Importantly, an ideal observer must be able to evaluate likelihood with high precision and only consider the likelihood of the two relevant stimuli involved in the discrimination task. We report a new perceptual bias suggesting that observers read out the likelihood representation with remarkably low precision when discriminating grating spatial frequencies. Using spectrally filtered noise, we induced an asymmetry in the likelihood function of spatial frequency. This manipulation mainly affects the likelihood of spatial frequencies that are irrelevant to the task at hand. Nevertheless, we find a significant shift in perceived grating frequency, indicating that observers evaluate likelihoods of a broad range of irrelevant frequencies and discard prior knowledge of stimulus alternatives when performing two-alternative discrimination.
Author Summary
An attractive view on human information processing proposes that inference problems are dealt with in a statistically optimal fashion. This hypothesis can explain aspects of perception, movement planning, cognition and decision making. In the present study, I use a new psychophysical paradigm that reveals surprisingly suboptimal perceptual decision making. Observers discriminate between two sinusoidal gratings of a different spatial frequency. Making use of visual noise, I induce an asymmetry in neural population responses to the gratings and find this asymmetry to effectively bias perceptual decision making. A simple ideal observer model, uninformed about the presence of visual noise but only considering the two grating spatial frequencies relevant to the task at hand, manages to avoid such a bias. I conclude that observers are limited in their ability to make use of prior knowledge of relevant visual features when performing this task. These results are in line with a growing number of findings suggesting that near-optimal decoders, although straightforward to implement and achieving near-maximal performance, consistently overestimate empirical performance in simple perceptual tasks.
doi:10.1371/journal.pcbi.1002453
PMCID: PMC3325184  PMID: 22511853
5.  The Task-Relevant Attribute Representation Can Mediate the Simon Effect 
PLoS ONE  2014;9(3):e90954.
Researchers have previously suggested a working memory (WM) account of spatial codes, and based on this suggestion, the present study carries out three experiments to investigate how the task-relevant attribute representation (verbal or visual) in the typical Simon task affects the Simon effect. Experiment 1 compared the Simon effect between the between- and within-category color conditions, which required subjects to discriminate between red and blue stimuli (presumed to be represented by verbal WM codes because it was easy and fast to name the colors verbally) and to discriminate between two similar green stimuli (presumed to be represented by visual WM codes because it was hard and time-consuming to name the colors verbally), respectively. The results revealed a reliable Simon effect that only occurs in the between-category condition. Experiment 2 assessed the Simon effect by requiring subjects to discriminate between two different isosceles trapezoids (within-category shapes) and to discriminate isosceles trapezoid from rectangle (between-category shapes), and the results replicated and expanded the findings of Experiment 1. In Experiment 3, subjects were required to perform both tasks from Experiment 1. Wherein, in Experiment 3A, the between-category task preceded the within-category task; in Experiment 3B, the task order was opposite. The results showed the reliable Simon effect when subjects represented the task-relevant stimulus attributes by verbal WM encoding. In addition, the response times (RTs) distribution analysis for both the between- and within-category conditions of Experiments 3A and 3B showed decreased Simon effect with the RTs lengthened. Altogether, although the present results are consistent with the temporal coding account, we put forth that the Simon effect also depends on the verbal WM representation of task-relevant stimulus attribute.
doi:10.1371/journal.pone.0090954
PMCID: PMC3949726  PMID: 24618692
6.  Anticipatory Regulation of Action Control in a Simon Task: Behavioral, Electrophysiological, and fMRI Correlates 
With the present study we investigated cue-induced preparation in a Simon task and measured electroencephalogram and functional magnetic resonance imaging (fMRI) data in two within-subjects sessions. Cues informed either about the upcoming (1) spatial stimulus-response compatibility (rule cues), or (2) the stimulus location (position cues), or (3) were non-informative. Only rule cues allowed anticipating the upcoming compatibility condition. Position cues allowed anticipation of the upcoming location of the Simon stimulus but not its compatibility condition. Rule cues elicited fastest and most accurate performance for both compatible and incompatible trials. The contingent negative variation (CNV) in the event-related potential (ERP) of the cue-target interval is an index of anticipatory preparation and was magnified after rule cues. The N2 in the post-target ERP as a measure of online action control was reduced in Simon trials after rule cues. Although compatible trials were faster than incompatible trials in all cue conditions only non-informative cues revealed a compatibility effect in additional indicators of Simon task conflict like accuracy and the N2. We thus conclude that rule cues induced anticipatory re-coding of the Simon task that did not involve cognitive conflict anymore. fMRI revealed that rule cues yielded more activation of the left rostral, dorsal, and ventral prefrontal cortex as well as the pre-SMA as compared to POS and NON-cues. Pre-SMA and ventrolateral prefrontal activation after rule cues correlated with the effective use of rule cues in behavioral performance. Position cues induced a smaller CNV effect and exhibited less prefrontal and pre-SMA contributions in fMRI. Our data point to the importance to disentangle different anticipatory adjustments that might also include the prevention of upcoming conflict via task re-coding.
doi:10.3389/fpsyg.2013.00047
PMCID: PMC3569607  PMID: 23408377
cognitive conflict; cueing; EEG; fMRI; pre-SMA; Simon task; anticipation; cognitive control
7.  The role of social cues in the deployment of spatial attention: head-body relationships automatically activate directional spatial codes in a Simon task 
The role of body orientation in the orienting and allocation of social attention was examined using an adapted Simon paradigm. Participants categorized the facial expression of forward facing, computer-generated human figures by pressing one of two response keys, each located left or right of the observers' body midline, while the orientation of the stimulus figure's body (trunk, arms, and legs), which was the task-irrelevant feature of interest, was manipulated (oriented toward the left or right visual hemifield) with respect to the spatial location of the required response. We found that when the orientation of the body was compatible with the required response location, responses were slower relative to when body orientation was incompatible with the response location. In line with a model put forward by Hietanen (1999), this reverse compatibility effect suggests that body orientation is automatically processed into a directional spatial code, but that this code is based on an integration of head and body orientation within an allocentric-based frame of reference. Moreover, we argue that this code may be derived from the motion information implied in the image of a figure when head and body orientation are incongruent. Our results have implications for understanding the nature of the information that affects the allocation of attention for social orienting.
doi:10.3389/fnint.2012.00004
PMCID: PMC3269793  PMID: 22347172
social attention; spatial attention; Simon task; head-body orientation; implied motion
8.  The Carry-Over Effect of Competition in Task-Sharing: Evidence from the Joint Simon Task 
PLoS ONE  2014;9(6):e97991.
The Simon effect, that is the advantage of the spatial correspondence between stimulus and response locations when stimulus location is a task-irrelevant dimension, occurs even when the task is performed together by two participants, each performing a go/no-go task. Previous studies showed that this joint Simon effect, considered by some authors as a measure of self-other integration, does not emerge when during task performance co-actors are required to compete. The present study investigated whether and for how long competition experienced during joint performance of one task can affect performance in a following joint Simon task. In two experiments, we required pairs of participants to perform together a joint Simon task, before and after jointly performing together an unrelated non-spatial task (the Eriksen flanker task). In Experiment 1, participants always performed the joint Simon task under neutral instructions, before and after performing the joint flanker task in which they were explicitly required either to cooperate with (i.e., cooperative condition) or to compete against a co-actor (i.e., competitive condition). In Experiment 2, they were required to compete during the joint flanker task and to cooperate during the subsequent joint Simon task. Competition experienced in one task affected the way the subsequent joint task was performed, as revealed by the lack of the joint Simon effect, even though, during the Simon task participants were not required to compete (Experiment 1). However, prior competition no longer affected subsequent performance if a new goal that created positive interdependence between the two agents was introduced (Experiment 2). These results suggest that the emergence of the joint Simon effect is significantly influenced by how the goals of the co-acting individuals are related, with the effect of competition extending beyond the specific competitive setting and affecting subsequent interactions.
doi:10.1371/journal.pone.0097991
PMCID: PMC4045759  PMID: 24897120
9.  Priming and Backward Influences in the Human Brain: Processing Interactions during the Stroop Interference Effect 
Cerebral Cortex (New York, NY)  2009;19(11):2508-2521.
This study investigated neural processing interactions during Stroop interference by varying the temporal separation of relevant and irrelevant features of congruent, neutral, and incongruent colored-bar/color-word stimulus components. High-density event-related potentials (ERPs) and behavioral performance were measured as participants reported the bar color as quickly as possible, while ignoring the color words. The task-irrelevant color words could appear at 1 of 5 stimulus onset asynchronies (SOAs) relative to the task-relevant bar-color occurrence: −200 or −100 ms before, +100 or +200 ms after, or simultaneously. Incongruent relative to congruent presentations elicited slower reaction times and higher error rates (with neutral in between), and ERP difference waves containing both an early, negative-polarity, central-parietal deflection, and a later, more left-sided, positive-polarity component. These congruency-related differences interacted with SOA, showing the greatest behavioral and electrophysiological effects when irrelevant stimulus information preceded the task-relevant target and reduced effects when the irrelevant information followed the relevant target. We interpret these data as reflecting 2 separate processes: 1) a ‘priming influence’ that enhances the magnitude of conflict-related facilitation and conflict-related interference when a task-relevant target is preceded by an irrelevant distractor; and 2) a reduced ‘backward influence’ of stimulus conflict when the irrelevant distractor information follows the task-relevant target.
doi:10.1093/cercor/bhp036
PMCID: PMC2764508  PMID: 19321654
conflict processing; event-related potentials (ERPs); incongruency; stimulus onset asynchrony (SOA); Stroop task
10.  Functional Difference between Sustained and Transient Modulations of Cognitive Control in the Simon Task: Evidence from False Alarm Responses on No-Go Trials 
PLoS ONE  2013;8(11):e81804.
Cognitive control in response compatibility tasks is modulated by the task context. Two types of contextual modulations have been demonstrated; sustained (block-wise) and transient (trial-by-trial). Recent research suggests that these modulations have different underlying mechanisms. This study presents new evidence supporting this claim by comparing false alarm (FA) responses on no-go trials of the Simon task between the sustained and transient contexts. In Experiment 1, the sustained context was manipulated so that a block included a larger number of incongruent trials. Results showed that participants made more FA responses by the hand opposite to the stimulus location. This suggests a generation of response bias in which the task-irrelevant location information is utilized in a reversed manner (i.e., to respond with the right hand to a stimulus presented on the left side and vice versa). Next, Experiment 2 examined the effect of the transient context and found that overall FA rate was lower when a no-go trial was preceded by an incongruent trial than by a congruent trial, whereas such response bias as that shown in Experiment 1 was not demonstrated. This suggests that the transient conflict context enhances inhibition of the task-irrelevant process but does not make the task-irrelevant information actively usable. Based on these results, we propound two types of cognitive control modulations as adaptive behaviors: response biasing based on utilization of the task-irrelevant information under the sustained conflict context and transient enhancement of inhibition of the task-irrelevant process based on the online conflict monitoring.
doi:10.1371/journal.pone.0081804
PMCID: PMC3841129  PMID: 24303072
11.  Diagnostic Features of Emotional Expressions Are Processed Preferentially 
PLoS ONE  2012;7(7):e41792.
Diagnostic features of emotional expressions are differentially distributed across the face. The current study examined whether these diagnostic features are preferentially attended to even when they are irrelevant for the task at hand or when faces appear at different locations in the visual field. To this aim, fearful, happy and neutral faces were presented to healthy individuals in two experiments while measuring eye movements. In Experiment 1, participants had to accomplish an emotion classification, a gender discrimination or a passive viewing task. To differentiate fast, potentially reflexive, eye movements from a more elaborate scanning of faces, stimuli were either presented for 150 or 2000 ms. In Experiment 2, similar faces were presented at different spatial positions to rule out the possibility that eye movements only reflect a general bias for certain visual field locations. In both experiments, participants fixated the eye region much longer than any other region in the face. Furthermore, the eye region was attended to more pronouncedly when fearful or neutral faces were shown whereas more attention was directed toward the mouth of happy facial expressions. Since these results were similar across the other experimental manipulations, they indicate that diagnostic features of emotional expressions are preferentially processed irrespective of task demands and spatial locations. Saliency analyses revealed that a computational model of bottom-up visual attention could not explain these results. Furthermore, as these gaze preferences were evident very early after stimulus onset and occurred even when saccades did not allow for extracting further information from these stimuli, they may reflect a preattentive mechanism that automatically detects relevant facial features in the visual field and facilitates the orientation of attention towards them. This mechanism might crucially depend on amygdala functioning and it is potentially impaired in a number of clinical conditions such as autism or social anxiety disorders.
doi:10.1371/journal.pone.0041792
PMCID: PMC3405011  PMID: 22848607
12.  Irrelevant Features of a Stimulus Can Either Facilitate or Disrupt Performance in a Working Memory Task: The Role of Fluid Intelligence 
PLoS ONE  2011;6(10):e26249.
It has been shown that fluid intelligence (gf) is fundamental to overcome interference due to information of a previously encoded item along a task-relevant domain. However, the biasing effect of task-irrelevant dimensions is still unclear as well as its relation with gf. The present study aimed at clarifying these issues. Gf was assessed in 60 healthy subjects. In a different session, the same subjects performed two versions (letter-detection and spatial) of a three-back working memory task with a set of physically identical stimuli (letters) presented at different locations on the screen. In the letter-detection task, volunteers were asked to match stimuli on the basis of their identity whereas, in the spatial task, they were required to match items on their locations. Cross-domain bias was manipulated by pseudorandomly inserting a match between the current and the three back items on the irrelevant domain. Our findings showed that a task-irrelevant feature of a salient stimulus can actually bias the ongoing performance. We revealed that, at trials in which the current and the three-back items matched on the irrelevant domain, group accuracy was lower (interference). On the other hand, at trials in which the two items matched on both the relevant and irrelevant domains, the group showed an enhancement of the performance (facilitation). Furthermore, we demonstrated that individual differences in fluid intelligence covaries with the ability to override cross-domain interference in that higher gf subjects showed better performance at interference trials than low gf subjects. Altogether, our findings suggest that stimulus features irrelevant to the task can affect cognitive performance along the relevant domain and that gf plays an important role in protecting relevant memory contents from the hampering effect of such a bias.
doi:10.1371/journal.pone.0026249
PMCID: PMC3194836  PMID: 22022580
13.  Attention improves encoding of task-relevant features in the human visual cortex 
When spatial attention is directed towards a particular stimulus, increased activity is commonly observed in corresponding locations of the visual cortex. Does this attentional increase in activity indicate improved processing of all features contained within the attended stimulus, or might spatial attention selectively enhance the features relevant to the observer’s task? We used fMRI decoding methods to measure the strength of orientation-selective activity patterns in the human visual cortex while subjects performed either an orientation or contrast discrimination task, involving one of two laterally presented gratings. Greater overall BOLD activation with spatial attention was observed in areas V1-V4 for both tasks. However, multivariate pattern analysis revealed that orientation-selective responses were enhanced by attention only when orientation was the task-relevant feature, and not when the grating’s contrast had to be attended. In a second experiment, observers discriminated the orientation or color of a specific lateral grating. Here, orientation-selective responses were enhanced in both tasks but color-selective responses were enhanced only when color was task-relevant. In both experiments, task-specific enhancement of feature-selective activity was not confined to the attended stimulus location, but instead spread to other locations in the visual field, suggesting the concurrent involvement of a global feature-based attentional mechanism. These results suggest that attention can be remarkably selective in its ability to enhance particular task-relevant features, and further reveal that increases in overall BOLD amplitude are not necessarily accompanied by improved processing of stimulus information.
doi:10.1523/JNEUROSCI.6153-09.2011
PMCID: PMC3134176  PMID: 21632942
fMRI; human; vision; attention; cortex; orientation
14.  Dissociating between object affordances and spatial compatibility effects using early response components 
Perception and action are tightly linked: objects may be perceived not only in terms of visual features, but also in terms of possibilities for action. Previous studies showed that when a centrally located object has a salient graspable feature (e.g., a handle), it facilitates motor responses corresponding with the feature's position. However, such so-called affordance effects have been criticized as resulting from spatial compatibility effects, due to the visual asymmetry created by the graspable feature, irrespective of any affordances. In order to dissociate between affordance and spatial compatibility effects, we asked participants to perform a simple reaction-time task to typically graspable and non-graspable objects with similar visual features (e.g., lollipop and stop sign). Responses were measured using either electromyography (EMG) on proximal arm muscles during reaching-like movements, or with finger key-presses. In both EMG and button press measurements, participants responded faster when the object was either presented in the same location as the responding hand, or was affordable, resulting in significant and independent spatial compatibility and affordance effects, but no interaction. Furthermore, while the spatial compatibility effect was present from the earliest stages of movement preparation and throughout the different stages of movement execution, the affordance effect was restricted to the early stages of movement execution. Finally, we tested a small group of unilateral arm amputees using EMG, and found residual spatial compatibility but no affordance, suggesting that spatial compatibility effects do not necessarily rely on individuals' available affordances. Our results show dissociation between affordance and spatial compatibility effects, and suggest that rather than evoking the specific motor action most suitable for interaction with the viewed object, graspable objects prompt the motor system in a general, body-part independent fashion.
doi:10.3389/fpsyg.2013.00591
PMCID: PMC3761160  PMID: 24027552
affordance; amputees; EMG; hand; stimulus-response
15.  The impact of fornix lesions in rats on spatial learning tasks sensitive to anterior thalamic and hippocampal damage 
Behavioural Brain Research  2015;278:360-374.
Highlights
•Fornix damage mildly impair spatial biconditional and passive place learning tasks.•Fornix lesions impair spatial go/no-go and alternation problems.•Fornix lesions impair tests making flexible demands on spatial memory.•Fornix connections are not always required for learning fixed spatial responses.
The present study sought to understand how the hippocampus and anterior thalamic nuclei are conjointly required for spatial learning by examining the impact of cutting a major tract (the fornix) that interconnects these two sites. The initial experiments examined the consequences of fornix lesions in rats on spatial biconditional discrimination learning. The rationale arose from previous findings showing that fornix lesions spare the learning of spatial biconditional tasks, despite the same task being highly sensitive to both hippocampal and anterior thalamic nuclei lesions. In the present study, fornix lesions only delayed acquisition of the spatial biconditional task, pointing to additional contributions from non-fornical routes linking the hippocampus with the anterior thalamic nuclei. The same fornix lesions spared the learning of an analogous nonspatial biconditional task that used local contextual cues. Subsequent tests, including T-maze place alternation, place learning in a cross-maze, and a go/no-go place discrimination, highlighted the impact of fornix lesions when distal spatial information is used flexibly to guide behaviour. The final experiment examined the ability to learn incidentally the spatial features of a square water-maze that had differently patterned walls. Fornix lesions disrupted performance but did not stop the rats from distinguishing the various corners of the maze. Overall, the results indicate that interconnections between the hippocampus and anterior thalamus, via the fornix, help to resolve problems with flexible spatial and temporal cues, but the results also signal the importance of additional, non-fornical contributions to hippocampal-anterior thalamic spatial processing, particularly for problems with more stable spatial solutions.
doi:10.1016/j.bbr.2014.10.016
PMCID: PMC4274319  PMID: 25453745
Biconditional learning; Cognitive map; Configural learning; Hippocampus; Incidental learning; Thalamus
16.  Visual Experience Determines the Use of External Reference Frames in Joint Action Control 
PLoS ONE  2013;8(3):e59008.
Vision plays a crucial role in human interaction by facilitating the coordination of one's own actions with those of others in space and time. While previous findings have demonstrated that vision determines the default use of reference frames, little is known about the role of visual experience in coding action-space during joint action. Here, we tested if and how visual experience influences the use of reference frames in joint action control. Dyads of congenitally-blind, blindfolded-sighted, and seeing individuals took part in an auditory version of the social Simon task, which required each participant to respond to one of two sounds presented to the left or right of both participants. To disentangle the contribution of external—agent-based and response-based—reference frames during joint action, participants performed the task with their respective response (right) hands uncrossed or crossed over one another. Although the location of the auditory stimulus was completely task-irrelevant, participants responded overall faster when the stimulus location spatially corresponded to the required response side than when they were spatially non-corresponding: a phenomenon known as the social Simon effect (SSE). In sighted participants, the SSE occurred irrespective of whether hands were crossed or uncrossed, suggesting the use of external, response-based reference frames. Congenitally-blind participants also showed an SSE, but only with uncrossed hands. We argue that congenitally-blind people use both agent-based and response-based reference frames resulting in conflicting spatial information when hands are crossed and, thus, canceling out the SSE. These results imply that joint action control functions on the basis of external reference frames independent of the presence or (transient/permanent) absence of vision. However, the type of external reference frames used for organizing motor control in joint action seems to be determined by visual experience.
doi:10.1371/journal.pone.0059008
PMCID: PMC3594222  PMID: 23536848
17.  Attention to the Color of a Moving Stimulus Modulates Motion-Signal Processing in Macaque Area MT: Evidence for a Unified Attentional System 
Directing visual attention to spatial locations or to non-spatial stimulus features can strongly modulate responses of individual cortical sensory neurons. Effects of attention typically vary in magnitude, not only between visual cortical areas but also between individual neurons from the same area. Here, we investigate whether the size of attentional effects depends on the match between the tuning properties of the recorded neuron and the perceptual task at hand. We recorded extracellular responses from individual direction-selective neurons in the middle temporal area (MT) of rhesus monkeys trained to attend either to the color or the motion signal of a moving stimulus. We found that effects of spatial and feature-based attention in MT, which are typically observed in tasks allocating attention to motion, were very similar even when attention was directed to the color of the stimulus. We conclude that attentional modulation can occur in extrastriate cortex, even under conditions without a match between the tuning properties of the recorded neuron and the perceptual task at hand. Our data are consistent with theories of object-based attention describing a transfer of attention from relevant to irrelevant features, within the attended object and across the visual field. These results argue for a unified attentional system that modulates responses to a stimulus across cortical areas, even if a given area is specialized for processing task-irrelevant aspects of that stimulus.
doi:10.3389/neuro.06.012.2009
PMCID: PMC2773174  PMID: 19893762
attention; visual objects; neuronal representation; motion; color; middle temporal area MT
18.  Hippocampal Lesions can Enhance Discrimination Learning Despite Normal Sensitivity to Interference From Incidental Information 
Hippocampus  2011;22(7):1553-1566.
Spatial properties of stimuli are sometimes encoded even when incidental to the demands of a particular learning task. Incidental encoding of spatial information may interfere with learning by (i) causing a failure to generalize learning between trials in which a cue is presented in different spatial locations and (ii) adding common spatial features to stimuli that predict different outcomes. Hippocampal lesions have been found to facilitate acquisition of certain tasks. This facilitation may occur because hippocampal lesions impair incidental encoding of spatial information that interferes with learning. To test this prediction mice with lesions of the hippocampus were trained on appetitive simple simultaneous discrimination tasks using inserts in the goal arms of a T-maze. It was found that hippocampal lesioned mice were facilitated at learning the discriminations, but they were sensitive to changes in spatial information in a manner that was similar to control mice. In a second experiment it was found that both control and hippocampal lesioned mice showed equivalent incidental encoding of egocentric spatial properties of the inserts, but both groups did not encode the allocentric information. These results demonstrate that mice show incidental encoding of egocentric spatial information that decreases the ability to solve simultaneous discrimination tasks. The normal egocentric spatial encoding in hippocampal lesioned mice contradicts theories of hippocampal function that suggest that the hippocampus is necessary for incidental learning per se, or is required for modulating stimulus representations based on the relevancy of information. The facilitated learning suggests that the hippocampal lesions can enhance learning of the same qualitative information as acquired by control mice. © 2011 Wiley Periodicals, Inc.
doi:10.1002/hipo.20995
PMCID: PMC3506972  PMID: 22161993
spatial learning; memory; mouse; lesion; conjunctive representations
19.  Influence of Task-Relevant and Task-Irrelevant Feature Continuity on Selective Auditory Attention 
Past studies have explored the relative strengths of auditory features in a selective attention task by pitting features against one another and asking listeners to report the words perceived in a given sentence. While these studies show that the continuity of competing features affects streaming, they did not address whether the influence of specific features is modulated by volitionally directed attention. Here, we explored whether the continuity of a task-irrelevant feature affects the ability to selectively report one of two competing speech streams when attention is specifically directed to a different feature. Sequences of simultaneous pairs of spoken digits were presented in which exactly one digit of each pair matched a primer phrase in pitch and exactly one digit of each pair matched the primer location. Within a trial, location and pitch were randomly paired; they either were consistent with each other from digit to digit or were switched (e.g., the sequence from the primer's location changed pitch across digits). In otherwise identical blocks, listeners were instructed to report digits matching the primer either in location or in pitch. Listeners were told to ignore the irrelevant feature, if possible, in order to perform well. Listener responses depended on task instructions, proving that top–down attention alters how a subject performs the task. Performance improved when the separation of the target and masker in the task-relevant feature increased. Importantly, the values of the task-irrelevant feature also influenced performance in some cases. Specifically, when instructed to attend location, listeners performed worse as the separation between target and masker pitch increased, especially when the spatial separation between digits was small. These results indicate that task-relevant and task-irrelevant features are perceptually bound together: continuity of task-irrelevant features influences selective attention in an automatic, obligatory manner, consistent with the idea that auditory attention operates on objects.
doi:10.1007/s10162-011-0299-7
PMCID: PMC3254717  PMID: 22124889
psychophysics; streaming; top–down; bottom–up; object-based attention
20.  Dynamic Grouping of Hippocampal Neural Activity During Cognitive Control of Two Spatial Frames 
PLoS Biology  2010;8(6):e1000403.
Hippocampal neurons represent two concurrent streams of spatial information by transiently organizing into subpopulations of coactive neurons and can reflect the most behaviorally relevant information at any given time.
Cognitive control is the ability to coordinate multiple streams of information to prevent confusion and select appropriate behavioral responses, especially when presented with competing alternatives. Despite its theoretical and clinical significance, the neural mechanisms of cognitive control are poorly understood. Using a two-frame place avoidance task and partial hippocampal inactivation, we confirmed that intact hippocampal function is necessary for coordinating two streams of spatial information. Rats were placed on a continuously rotating arena and trained to organize their behavior according to two concurrently relevant spatial frames: one stationary, the other rotating. We then studied how information about locations in these two spatial frames is organized in the action potential discharge of ensembles of hippocampal cells. Both streams of information were represented in neuronal discharge—place cell activity was organized according to both spatial frames, but almost all cells preferentially represented locations in one of the two spatial frames. At any given time, most coactive cells tended to represent locations in the same spatial frame, reducing the risk of interference between the two information streams. An ensemble's preference to represent locations in one or the other spatial frame alternated within a session, but at each moment, location in the more behaviorally relevant spatial frame was more likely to be represented. This discharge organized into transient groups of coactive neurons that fired together within 25 ms to represent locations in the same spatial frame. These findings show that dynamic grouping, the transient coactivation of neural subpopulations that represent the same stream of information, can coordinate representations of concurrent information streams and avoid confusion, demonstrating neural-ensemble correlates of cognitive control in hippocampus.
Author Summary
Understanding the world and making optimal decisions requires using the most relevant information while at the same time ignoring irrelevant information, a psychological phenomenon known as “cognitive control.” How the same population of neurons deals with multiple streams of information simultaneously is poorly understood. In this study, we investigated the underlying neural mechanisms of cognitive control in a network of hippocampal neurons known to represent space. We implanted electrodes into the hippocampus of rats and recorded the action potential discharge of many neurons at the same time. The recordings were made while rats that were foraging on a rotating disk used cognitive control to coordinate spatial information from different spatial frames. We found that at each moment, discharge preferentially represented location in one or the other spatial frame. Importantly, we were able to influence the behavioral relevance of these spatial frames, and we found that discharge alternated between signaling location in one or the other frames in accord with its current behavioral importance. The timing of when these neurons were active was also related to their function, such that neurons collectively represented locations in the same spatial frame if they were coactive within a few tens of milliseconds to seconds. We conclude that cognitive control is mediated by a dynamic functional grouping. Neural activity distributed across many neurons transiently organizes into functional groups by coactive firing that represents a coherent stream of information.
doi:10.1371/journal.pbio.1000403
PMCID: PMC2889929  PMID: 20585373
21.  Multisensory interactions follow the hands across the midline: Evidence from a non-spatial visual-tactile congruency task 
Brain research  2006;1077(1):108-115.
Crossing the hands over, whether across the body midline or with respect to each other, leads to measurable changes in spatial compatibility, spatial attention, and frequently to a general decrement in discrimination performance for tactile stimuli. The majority of multisensory crossed hands effects, however, have been demonstrated with explicit or implicit spatial discrimination tasks, raising the question of whether non-spatial discrimination tasks also show spatial effects when the hands are crossed. We designed a novel, non-spatial tactile discrimination task to address this issue. Participants made speeded discriminations of single-versus double-pulse vibrotactile targets, while trying to ignore simultaneous visual distractor stimuli, in both hands uncrossed, and hands crossed postures. Tactile discrimination performance was significantly affected by the visual distractors (demonstrating a significant crossmodal congruency effect), and was affected most by visual distractors in the same external location as the tactile target (i.e., spatial modulation), regardless of the posture (uncrossed or crossed) of the hands (i.e., spatial ‘re-mapping’ of visual-tactile interactions). Finally, crossing the hands led to a general performance decrement with visual distractors, but not in a control task with unimodal visual or tactile judgements. These results demonstrate, for the first time, significant spatial and postural modulations of crossmodal congruency effects in a non-spatial discrimination task.
doi:10.1016/j.brainres.2005.11.010
PMCID: PMC1482253  PMID: 16483553
Multisensory; Crossmodal congruency; Crossed hands; Somatosensation; Vision
22.  A task-irrelevant stimulus attribute affects perception and short-term memory 
Memory & cognition  2009;37(8):1088-1102.
Selective attention protects cognition against intrusions of task-irrelevant stimulus attributes. This protective function was tested in coordinated psychophysical and memory experiments. Stimuli were superimposed, horizontally and vertically oriented gratings of varying spatial frequency; only one orientation was task relevant. Experiment 1 demonstrated that a task-irrelevant spatial frequency interfered with visual discrimination of the task-relevant spatial frequency. Experiment 2 adopted a two-item Sternberg task, using stimuli that had been scaled to neutralize interference at the level of vision. Despite being visually neutralized, the task-irrelevant attribute strongly influenced recognition accuracy and associated reaction times (RTs). This effect was sharply tuned, with the task-irrelevant spatial frequency having an impact only when the task-relevant spatial frequencies of the probe and study items were highly similar to one another. Model-based analyses of judgment accuracy and RT distributional properties converged on the point that the irrelevant orientation operates at an early stage in memory processing, not at a later one that supports decision making.
doi:10.3758/MC.37.8.1088
PMCID: PMC2836167  PMID: 19933454
23.  When co-action eliminates the Simon effect: disentangling the impact of co-actor's presence and task sharing on joint-task performance 
This study aimed at assessing whether the mere belief of performing a task with another person, who is in charge of the complementary part of the task, is sufficient for the so-called joint Simon effect to occur. In all three experiments of the study, participants sat alone in a room and underwent two consecutive Go/NoGo tasks that were identical except for the instructions. In Experiment 1, participants performed the task first individually (baseline task), and then either co-acting with another person who responded from an unknown location to the NoGo stimuli (joint task) or imaging themselves responding to the NoGo stimuli (imaginative task). Relative to the baseline, the instructions of the imaginative task made the Simon effect occur, while those of the joint task were ineffective in eliciting the effect. This result suggests that sharing a task with a person who is known to be in charge of the complementary task, but is not physically present, is not sufficient to induce the representation of an alternative response able to produce interference, which happens instead when the instructions explicitly require to imagine such a response. Interestingly, we observed that when the Simon effect was already present in the baseline task (i.e., when the response alternative to the Go response was represented in the individual task due to non-social factors), it disappeared in the joint task. We propose that, when no information about the co-actor's position is available, the division of labor between the participant and co-actor allows participants to filter out the possible (incidental) representation of the alternative response from their task representation, thus eliminating potential sources of interference. This account is supported by the results of Experiments 2 and 3 and suggests that under certain circumstances task-sharing may reduce the interference produced by the irrelevant information, rather than increase it.
doi:10.3389/fpsyg.2013.00844
PMCID: PMC3833097  PMID: 24312066
spatial compatibility; joint Simon effect; joint action; social interaction; social cognition; division of labor; referential coding; action co-representation
24.  Independent Processing of Stimulus-Stimulus and Stimulus-Response Conflicts 
PLoS ONE  2014;9(2):e89249.
The dimensional overlap (DO) model proposes distinct mechanisms for stimulus-stimulus (S-S) and stimulus-response (S-R) conflict effects. Many studies have examined the independence of S-S and S-R conflict effects in the color-word Stroop and Simon tasks. However, confounds exist between the distinction of DO (i.e., S-S dimensional overlap compared with S-R dimensional overlap) and the distinction of stimulus attributes (e.g., color compared with spatial location; semantic compared with nonsemantic information), which may hinder interpretation of the independence of S-S and S-R conflicts. A spatial Stroop (word) task and a spatial Stroop (arrow) task were combined with a Simon task in Experiments 1 and 2, respectively to eliminate these confounds of stimulus attributes. The results showed that S-S and S-R conflicts affected performance additively. There was no significant correlation across participants. These findings lend further support to independent processing of S-S and S-R conflicts as it is outlined in the taxonomy of DO.
doi:10.1371/journal.pone.0089249
PMCID: PMC3928426  PMID: 24558489
25.  The influence of prior practice and handedness on the orthogonal Simon effect 
When stimuli are arranged vertically and responses horizontally, right-handed participants respond faster with right responses to stimuli presented above fixation and with left responses to stimuli presented below fixation, even when stimulus position is task-irrelevant (orthogonal Simon effect). The aim of the present work was twofold. First, we assessed whether the orthogonal Simon effect evident in right-handed participants is present also for left-handed participants (Experiment 1). Second, we investigated whether for both groups of participants the orthogonal Simon effect is influenced by the stimulus-response (S-R) mapping used for an orthogonal spatial S-R compatibility task performed 5 min before (Experiment 2). Our results showed that the orthogonal Simon effect significantly differed in the two groups, with left-handers showing an advantage for the up-left/down-right mapping (Experiment 1). Interestingly, the orthogonal Simon effect was strongly influenced by prior practice regardless of the participants’ handedness (Experiment 2). These results suggest that the short-term S-R associations acquired during practice can override the long-term, hardwired associations established on the basis of handedness.
doi:10.3389/fpsyg.2014.00039
PMCID: PMC3916779  PMID: 24570665
orthogonal Simon effect; orthogonal spatial compatibility; handedness; practice paradigm; S-R associations

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