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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.  Reactive control processes contributing to residual switch cost and mixing cost across the adult lifespan 
In task-switching paradigms, performance is better when repeating the same task than when alternating between tasks (switch cost) and when repeating a task alone rather than intermixed with another task (mixing cost). These costs remain even after extensive practice and when task cues enable advanced preparation (residual costs). Moreover, residual reaction time mixing cost has been consistently shown to increase with age. Residual switch and mixing costs modulate the amplitude of the stimulus-locked P3b. This mixing effect is disproportionately larger in older adults who also prepare more for and respond more cautiously on these “mixed” repeat trials (Karayanidis et al., 2011). In this paper, we analyze stimulus-locked and response-locked P3 and lateralized readiness potentials to identify whether residual switch and mixing cost arise from the need to control interference at the level of stimulus processing or response processing. Residual mixing cost was associated with control of stimulus-level interference, whereas residual switch cost was also associated with a delay in response selection. In older adults, the disproportionate increase in mixing cost was associated with greater interference at the level of decision-response mapping and response programming for repeat trials in mixed-task blocks. These findings suggest that older adults strategically recruit greater proactive and reactive control to overcome increased susceptibility to post-stimulus interference. This interpretation is consistent with recruitment of compensatory strategies to compensate for reduced repetition benefit rather than an overall decline on cognitive flexibility.
doi:10.3389/fpsyg.2014.00383
PMCID: PMC4012181  PMID: 24817859
cognitive control; aging; lifespan; ERP; evidence accumulation models
3.  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
4.  Cognitive and Electrophysiological Correlates of the Bilingual Stroop Effect 
The color word Stroop effect in bilinguals is commonly half the magnitude when the written and naming languages are different (between) than when they are the same (within). This between-within language Stroop difference (BWLS) is likened to a response set effect, with greater response conflict for response relevant than irrelevant words. The nature of the BWLS was examined using a bilingual Stroop task. In a given block (Experiment 1), color congruent and incongruent words appeared in the naming language or not (single), or randomly in both languages (mixed). The BWLS effect was present for both balanced and unbalanced bilinguals, but only partially supported a response set explanation. As expected, color incongruent trials during single language blocks, lead to slower response times within than between languages. However, color congruent trials during mixed language blocks led to slower times between than within languages, indicating that response-irrelevant stimuli interfered with processing. In Experiment 2, to investigate the neural timing of the BWLS effect, event related potentials were recorded while balanced bilinguals named silently within and between languages. Replicating monolingual findings, an N450 effect was observed with larger negative amplitude for color incongruent than congruent trials (350–550 ms post-stimulus onset). This effect was equivalent within and between languages, indicating that color words from both languages created response conflict, contrary to a strict response set effect. A sustained negativity (SN) followed with larger amplitude for color incongruent than congruent trials, resolving earlier for between than within language Stroop. This effect shared timing (550–700 ms), but not morphology or scalp distribution with the commonly reported sustained potential. Finally, larger negative amplitude (200–350 ms) was observed between than within languages independent of color congruence. This negativity, likened to a no-go N2, may reflect processes of inhibitory control that facilitate the resolution of conflict at the SN, while the N450 reflects parallel processing of distracter words, independent of response set (or language). In sum, the BWLS reflects brain activity over time with contributions from language and color conflict at different points.
doi:10.3389/fpsyg.2012.00081
PMCID: PMC3317261  PMID: 22485099
bilingual; Stroop; response conflict; between language interference; N450; N2; event related potential; language dominance
5.  Age Differences in the Neuroelectric Adaptation to Meaningful Sounds 
PLoS ONE  2013;8(7):e68892.
Much of what we know regarding the effect of stimulus repetition on neuroelectric adaptation comes from studies using artificially produced pure tones or harmonic complex sounds. Little is known about the neural processes associated with the representation of everyday sounds and how these may be affected by aging. In this study, we used real life, meaningful sounds presented at various azimuth positions and found that auditory evoked responses peaking at about 100 and 180 ms after sound onset decreased in amplitude with stimulus repetition. This neural adaptation was greater in young than in older adults and was more pronounced when the same sound was repeated at the same location. Moreover, the P2 waves showed differential patterns of domain-specific adaptation when location and identity was repeated among young adults. Background noise decreased ERP amplitudes and modulated the magnitude of repetition effects on both the N1 and P2 amplitude, and the effects were comparable in young and older adults. These findings reveal an age-related difference in the neural processes associated with adaptation to meaningful sounds, which may relate to older adults’ difficulty in ignoring task-irrelevant stimuli.
doi:10.1371/journal.pone.0068892
PMCID: PMC3723892  PMID: 23935900
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.  Searching from the top down: Ageing and attentional guidance during singleton detection 
Previous investigations of adult age differences in visual search suggest that an age-related decline may exist in attentional processes dependent on the observer's knowledge of task-relevant features (top–down processing). The present experiments were conducted to examine age-related changes in top–down attentional guidance during a highly efficient form of search, singleton detection. In Experiment 1 reaction times to detect targets were lower when target features were constant (feature condition) than when target features were allowed to vary between trials (mixed condition), and this reaction time benefit was similar for younger and older adults. Experiments 2 and 3 investigated possible interactions between top–down and bottom–up (stimulus-driven) processes. Experiment 2 demonstrated that search times for both age groups could be improved when targets varied on an additional feature from distractors (double-feature condition) but only when top–down control was available (feature search). In Experiment 3, the availability of top–down guidance enabled both younger and older adults to override the distracting effects of a noninformative spatial location cue. These findings indicate that top–down attentional control mechanisms interact with bottom–up processes to guide search for targets, and that in the context of singleton detection these mechanisms of top–down control are preserved for older adults.
PMCID: PMC1805468  PMID: 15881292
8.  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
9.  Predictive knowledge of stimulus relevance does not influence top-down suppression of irrelevant information in older adults 
Our ability to focus attention on task-relevant stimuli and ignore irrelevant distractions is reflected by differential enhancement and suppression of neural activity in sensory cortices. Previous research has shown that older adults exhibit a deficit in suppressing task-irrelevant information, the magnitude of which is associated with a decline in working memory performance. However, it remains unclear if a failure to suppress is a reflection of an inability of older adults to rapidly assess the relevance of information upon stimulus presentation when they are not aware of the relevance beforehand. To address this, we recorded the electroencephalogram (EEG) in healthy older participants (aged 60–80 years) while they performed two different versions of a selective face/scene working memory task, both with and without prior knowledge as to when relevant and irrelevant stimuli would appear. Each trial contained two faces and two scenes presented sequentially followed by a nine second delay and a probe stimulus. Participants were given the following instructions: remember faces (ignore scenes), remember scenes (ignore faces), remember the xth and yth stimuli (where x and y could be 1st, 2nd, 3rd or 4th), or passively view all stimuli. Working memory performance remained consistent regardless of task instructions. Enhanced neural activity was observed at posterior electrodes to attended stimuli, while neural responses that reflected the suppression of irrelevant stimuli was absent for both tasks. The lack of significant suppression at early stages of visual processing was revealed by P1 amplitude and N1 latency modulation indices. These results reveal that prior knowledge of stimulus relevance does not modify early neural processing during stimulus encoding and does not improve working memory performance in older adults. These results suggest that the inability to suppress irrelevant information early in the visual processing stream by older adults is related to mechanisms specific to top-down suppression.
doi:10.1016/j.cortex.2009.08.003
PMCID: PMC2826515  PMID: 19744649
suppression; EEG; aging; working memory; selective attention
10.  Can Older Adults Enhance Task-Switching Performance by Verbal Self-Instructions? The Influence of Working-Memory Load and Early Learning 
In this study we examined (a) whether verbal self-instructions can enhance task-switching performance in younger and older adults, and (b) whether verbal self-instruction benefits on task switching are smaller when memory demands on keeping track of the task sequence are reduced by spatial task cueing. Task-switching ability was measured as the difference in performance between single-task and mixed-task blocks (termed mixing costs), in which participants switched between two tasks A and B. One group of participants performed the switching tasks with spatial task cues, indicating which of the two tasks has to be performed, thereby reducing demands on the endogenous control of serial task order (low task-sequencing load). The other group switched between tasks without external task cues (high task-sequencing load). To investigate the influence of verbal self-instructions on task switching, participants either named aloud the next task during task preparation (task-naming condition) or they did not verbalize (control condition). Results indicated that age differences in verbalization benefits on mixing costs depend on early learning whereby benefits were generally larger when subjects had some prior practice in task switching alone, and that verbalization benefits did not differ between the two task-sequencing load conditions. These findings suggest that task naming is a suitable cognitive intervention for enhancing the control of task switching in younger and older adults, even if memory load is reduced, and that for the efficient application of this strategy it first has to be coordinated with task switching, which is easier when task switching is already practiced.
doi:10.3389/fnagi.2010.00147
PMCID: PMC2992032  PMID: 21120146
aging; task switching; verbalization benefits; working-memory load; task-sequencing function; endogenous task control
11.  Development of cognitive control and executive functions from 4 to 13 years: Evidence from manipulations of memory, inhibition, and task switching 
Neuropsychologia  2006;44(11):2037-2078.
Predictions concerning development, interrelations, and possible independence of working memory, inhibition, and cognitive flexibility were tested in 325 participants (roughly 30 per age from 4 to 13 years and young adults; 50% female). All were tested on the same computerized battery, designed to manipulate memory and inhibition independently and together, in steady state (single-task blocks) and during task-switching, and to be appropriate over the lifespan and for neuroimaging (fMRI). This is one of the first studies, in children or adults, to explore: (a) how memory requirements interact with spatial compatibility and (b) spatial incompatibility effects both with stimulus-specific rules (Simon task) and with higher-level, conceptual rules. Even the youngest children could hold information in mind, inhibit a dominant response, and combine those as long as the inhibition required was steady-state and the rules remained constant. Cognitive flexibility (switching between rules), even with memory demands minimized, showed a longer developmental progression, with 13-year-olds still not at adult levels. Effects elicited only in Mixed blocks with adults were found in young children even in single-task blocks; while young children could exercise inhibition in steady state it exacted a cost not seen in adults, who (unlike young children) seemed to re-set their default response when inhibition of the same tendency was required throughout a block. The costs associated with manipulations of inhibition were greater in young children while the costs associated with increasing memory demands were greater in adults. Effects seen only in RT in adults were seen primarily in accuracy in young children. Adults slowed down on difficult trials to preserve accuracy; but the youngest children were impulsive; their RT remained more constant but at an accuracy cost on difficult trials. Contrary to our predictions of independence between memory and inhibition, when matched for difficulty RT correlations between these were as high as 0.8, although accuracy correlations were less than half that. Spatial incompatibility effects and global and local switch costs were evident in children and adults, differing only in size. Other effects (e.g., asymmetric switch costs and the interaction of switching rules and switching response-sites) differed fundamentally over age.
doi:10.1016/j.neuropsychologia.2006.02.006
PMCID: PMC1513793  PMID: 16580701
Task switching; Inhibition; Working memory; Simon effect; Asymmetric switch costs; Global and local switch costs; Stimulus-response compatibility; Development; Children; Frontal lobe
12.  Training Reveals the Sources of Stroop and Flanker Interference Effects 
PLoS ONE  2013;8(10):e76580.
In the field of cognitive control, dimensional overlap and pathway automaticity are generally believed to be critical for the generation of congruency effects. However, their specific roles in the generation of congruency effects are unclear. In two experiments, with the 4∶2 mapping design, we investigated this issue by examining the training-related effects on congruency effects (the Stroop interference effect and the Flanker interference effect in Experiments 1 and 2, respectively) normally expressed as incongruent minus congruent difference and on their subcomponents (the stimulus interference and response interference). Experiment 1 revealed that the stimulus interference in the Stroop task, wherein the task-relevant (printed color of word) and the task-irrelevant (semantics of word) dimensions of the stimuli were processed in different pathways, was present during early training but was virtually eliminated at the late stage of training. This indicates that the two dimensions overlap at the early stage but separate at the late stage. In contrast, Experiment 2 showed that the response interference in a variant of the Flanker task, wherein the task-relevant (central color word printed in black font) and the task-irrelevant (flanking color words printed in black font) dimensions of the stimuli were processed in the same pathway, was enhanced after training. This indicates that the enhanced automaticity of irrelevant-dimension processing induces stronger response competition, which therefore results in the larger response interference. Taken together, the present study demonstrates that (1) dimensional overlap is necessary for the generation of congruency effects, (2) pathway automaticity can affect the size of congruency effects, and (3) training enhances the degree of automatic processing in a given pathway.
doi:10.1371/journal.pone.0076580
PMCID: PMC3795719  PMID: 24146892
13.  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
14.  Processing Irrelevant Location Information: Practice and Transfer Effects in a Simon Task 
PLoS ONE  2013;8(7):e64993.
How humans produce cognitively driven fine motor movements is a question of fundamental importance in how we interact with the world around us. For example, we are exposed to a constant stream of information and we must select the information that is most relevant by which to guide our actions. In the present study, we employed a well-known behavioral assay called the Simon task to better understand how humans are able to learn to filter out irrelevant information. We trained subjects for four days with a visual stimulus presented, alternately, in central and lateral locations. Subjects responded with one hand moving a joystick in either the left or right direction. They were instructed to ignore the irrelevant location information and respond based on color (e.g. red to the right and green to the left). On the fifth day, an additional testing session was conducted where the task changed and the subjects had to respond by shape (e.g. triangle to the right and rectangle to the left). They were instructed to ignore the color and location, and respond based solely on the task relevant shape. We found that the magnitude of the Simon effect decreases with training, however it returns in the first few trials after a break. Furthermore, task-defined associations between response direction and color did not significantly affect the Simon effect based on shape, and no significant associative learning from the specific stimulus-response features was found for the centrally located stimuli. We discuss how these results are consistent with a model involving route suppression/gating of the irrelevant location information. Much of the learning seems to be driven by subjects learning to suppress irrelevant location information, however, this seems to be an active inhibition process that requires a few trials of experience to engage.
doi:10.1371/journal.pone.0064993
PMCID: PMC3701670  PMID: 23861735
15.  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
16.  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
17.  Age-Related Changes in Attentional Selection: Quality of Task Set or Degradation of Task Set Across Time? 
Psychology and aging  2013;28(3):744-753.
The present study explores the nature of attentional selection in younger and older adults. Following de Jong, Berendsen, and Cools (1999, Acta Psychologica, 101, 379–394), we manipulated the response to stimulus interval (RSI) in two attentional selection paradigms to examine if there are age-related differences in the quality of task set and/or the maintenance of task set across time. In Experiment 1, we found that the interference effect in a spatial interference task was (a) overall larger in older adults compared to younger adults, and (b) smaller at the short RSI (200 ms) compared to the long RSI (2000 ms), and (c) there was no hint of an interaction between age and RSI. The second experiment explored the same variables in a Stroop color interference paradigm. Again, older adults produced a disproportionately larger interference effect than younger adults, the interference effect was smaller at the short RSI compared to the long RSI, and there was no evidence of an interaction between age and RSI. In both experiments the larger interference effect could not be attributed to age-related general slowing and there was evidence from Vincentile analyses of increasing interference and age effects at the slower response latencies. These results indicate that attentional selection deficits in these two experiments were due to a breakdown in the quality of the task set as opposed to age-related differences in the maintenance of the task set across time.
doi:10.1037/a0033159
PMCID: PMC3979426  PMID: 23834491
18.  Performance Dip in Motor Response Induced by Task-Irrelevant Weaker Coherent Visual Motion Signals 
Cerebral Cortex (New York, NY)  2011;22(8):1887-1893.
The Performance Dip is a newly characterized behavioral phenomenon, where, paradoxically, a weaker task-irrelevant visual stimulus causes larger disturbances on the accuracy of a main letter identification task than a stronger stimulus does. Understanding mechanisms of the Performance Dip may provide insight into unconsciousness behavior. Here, we investigated the generalization of the Performance Dip. Specifically, we tested whether the Performance Dip occurs in a motion-related Simon task, and if so, whether the Performance Dip involves the same brain region, that is, the dorsolateral prefrontal cortex (DLPFC), previously implicated in the Performance Dip, or the supplementary motor area (SMA) and pre-SMA, implicated in a motion-related Simon Task. Subjects made manual directional responses according to the color of stochastic moving dots while ignoring the global direction of moving dots, which could be either congruent or incongruent to the response appropriate to the main task. We found that weak incongruent task-irrelevant stimuli caused a Performance Dip, in which the SMA and pre-SMA, rather than DLPFC, played critical roles. Our results suggest a possible common brain mechanism across different neural circuits, in which weak, but not strong, task-irrelevant information is free from inhibition and intrudes into neural circuits relevant to the main task.
doi:10.1093/cercor/bhr270
PMCID: PMC3388893  PMID: 21940704
fMRI; performance dip; pre-SMA/SMA; task-irrelevant stimulus; visual motion stimulus
19.  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
20.  Inhibitory Motor Control in Old Age: Evidence for De-Automatization? 
To examine age-related effects on high-level consciously controlled and low-level automatically controlled inhibitory processes, the Simon task was combined with the masked prime task in a hybrid procedure. Young and older adults responded to the identity of targets (left/right key-press to left-/right-pointing arrows) that appeared on the left/right of the screen and were preceded by left-/right-pointing backward-masked arrow primes at fixation. Responses were faster and more accurate when the target was congruent with its location than incongruent (Simon effect), and when the target was incompatible with the prime than compatible (negative compatibility effect; NCE). The Simon effect was disproportionately larger, and the NCE disproportionately delayed, in older adults compared to young adults, indicating both high- and low-level inhibitory control deficits with aging. Moreover, the two effects were additive in young adults, but interactive in older adults, providing support for the dedifferentiation hypothesis of aging. Specifically, older adults’ prime-related inhibitory control appeared improved on incongruent relative to congruent trials, suggesting that impaired automatic control was substituted by high-level, non-automatic processes.
doi:10.3389/fpsyg.2011.00132
PMCID: PMC3122077  PMID: 21734899
aging; inhibition; cognitive control; masked priming; negative compatibility effect; Simon effect; dedifferentiation
21.  Conflict Processing in the Rat Brain: Behavioral Analysis and Functional μPET Imaging Using [18F]Fluorodeoxyglucose 
Conflicts in spatial stimulus–response tasks occur when the task-relevant feature of a stimulus implies a response toward a certain location which does not match the location of stimulus presentation. This conflict leads to increased error rates and longer reaction times, which has been termed Simon effect. A model of dual route processing (automatic and intentional) of stimulus features has been proposed, predicting response conflicts if the two routes are incongruent. Although there is evidence that the prefrontal cortex, notably the anterior cingulate cortex (ACC), plays a crucial role in conflict processing, the neuronal basis of dual route architecture is still unknown. In this study, we pursue a novel approach using positron emission tomography (PET) to identify relevant brain areas in a rat model of an auditory Simon task, a neuropsychological interference task, which is commonly used to study conflict processing in humans. For combination with PET we used the metabolic tracer [18F]fluorodeoxyglucose, which accumulates in metabolically active brain cells during the behavioral task. Brain areas involved in conflict processing are supposed to be activated when automatic and intentional route processing lead to different responses (dual route model). Analysis of PET data revealed specific activation patterns for different task settings applicable to the dual route model as established for response conflict processing. The rat motor cortex (M1) may be part of the automatic route or involved in its facilitation, while premotor (M2), prelimbic, and ACC seemed to be essential for inhibiting the incorrect, automatic response, indicating conflict monitoring functions. Our findings and the remarkable similarities to the pattern of activated regions reported during conflict processing in humans demonstrate that our rodent model opens novel opportunities to investigate the anatomical basis of conflict processing and dual route architecture.
doi:10.3389/fnbeh.2012.00004
PMCID: PMC3275787  PMID: 22363272
prefrontal cortex; rodent model; cognitive conflict; Simon task
22.  The Effects of Foreknowledge and Task-Set Shifting as Mirrored in Cue- and Target-Locked Event-Related Potentials 
PLoS ONE  2012;7(11):e49486.
The present study examined the use of foreknowledge in a task-cueing protocol while manipulating sensory updating and executive control in both, informatively and non-informatively pre-cued trials. Foreknowledge, sensory updating (cue switch effects) and task-switching were orthogonally manipulated in order to address the question of whether, and to which extent, the sensory processing of cue changes can partly or totally explain the final task switch costs. Participants responded faster when they could prepare for the upcoming task and if no task-set updating was necessary. Sensory cue switches influenced cue-locked ERPs only when they contained conceptual information about the upcoming task: frontal P2 amplitudes were modulated by task-relevant cue changes, mid-parietal P3 amplitudes by the anticipatory updating of stimulus-response mappings, and P3 peak latencies were modulated by task switching. Task preparation was advantageous for efficient stimulus-response re-mapping at target-onset as mirrored in target N2 amplitudes. However, N2 peak latencies indicate that this process is faster for all repeat trials. The results provide evidence to support a very fast detection of task-relevance in sensory (cue) changes and argue against the view of task repetition benefits as secondary to purely perceptual repetition priming. Advanced preparation may have a stronger influence on behavioral performance and target-locked brain activity than the local effect of repeating or switching the task-set in the current trial.
doi:10.1371/journal.pone.0049486
PMCID: PMC3495879  PMID: 23152912
23.  An Investigation of Response and Stimulus Modality Transfer Effects after Dual-Task Training in Younger and Older 
It has been shown that dual-task training leads to significant improvement in dual-task performance in younger and older adults. However, the extent to which training benefits to untrained tasks requires further investigation. The present study assessed (a) whether dual-task training leads to cross-modality transfer in untrained tasks using new stimuli and/or motor responses modalities, (b) whether transfer effects are related to improved ability to prepare and maintain multiple task-set and/or enhanced response coordination, (c) whether there are age-related differences in transfer effects. Twenty-three younger and 23 older adults were randomly assigned to dual-task training or control conditions. All participants were assessed before and after training on three dual-task transfer conditions; (1) stimulus modality transfer (2) response modality transfer (3) stimulus and response modalities transfer task. Training group showed larger improvement than the control group in the three transfer dual-task conditions, which suggests that training leads to more than specific learning of stimuli/response associations. Attentional costs analyses showed that training led to improved dual-task cost, only in conditions that involved new stimuli or response modalities, but not both. Moreover, training did not lead to a reduced task-set cost in the transfer conditions, which suggests some limitations in transfer effects that can be expected. Overall, the present study supports the notion that cognitive plasticity for attentional control is preserved in late adulthood.
doi:10.3389/fnhum.2012.00129
PMCID: PMC3355323  PMID: 22629239
cognitive plasticity; cognitive training; transfer; divided attention; executive function; aging
24.  Age-related deficits in efficiency of low-level lateral inhibition 
Background: In a masked prime task using a 0 ms prime-target inter-stimulus-interval, responses on trials where prime and target match (compatible trials) are usually faster and more accurate than responses where prime and target mismatch (incompatible trials). This positive compatibility effect (PCE) comprises both behavioral benefits on compatible relative to neutral trials, and behavioral costs on incompatible relative to neutral trials. Comparing performance in 2- vs. 4-alternative-response versions of the task indicates that benefits are due to direct priming (i.e., pre-activation) of a motor response, whereas costs reflect an inhibition of the alternative response tendency. The present study employs this paradigm to test the hypothesis that normal aging is associated with a selective deficit in inhibitory function, affecting both low-level motor and higher-level executive control. Experiment and Results: Testing 20 young and 20 older healthy adults, we found that (1) overall, prime-induced benefits were of similar magnitude across age groups, but inhibition-based costs were smaller in older compared to young adults; (2) increasing the number of response alternatives caused the same pattern of unaltered benefits and reduced costs in both age groups; and (3) costs, but not benefits, in the 2-alternative condition were significantly predicted by scores on the digit symbol substitution task (DSST), independently of age and other background variables. Interpretation: Results demonstrate the possibility of isolating an inhibitory component in low-level perceptuo-motor control. Importantly, this component shows an age-related decline in the absence of a corresponding decline of visuo-motor excitability, and appears to be linked to performance on a higher-level processing speed task. We hypothesize that aging might affect the brain's ability to establish precise short-term lateral inhibitory links, and that even in young adults, the efficiency of such links is a significant contributing factor in higher-level cognitive performance.
doi:10.3389/fnhum.2012.00102
PMCID: PMC3338071  PMID: 22557955
aging; inhibition; motor control; positive compatibility effect; masked priming
25.  Reducing and Restoring Stimulus-Response Compatibility Effects by Decreasing the Discriminability of Location Words 
Acta psychologica  2008;130(1):95-102.
In two experiments, we compared level of activation and temporal overlap accounts of compatibility effects in the Simon task by reducing the discriminability of spatial and non-spatial features of a target location word. Participants made keypress responses to the non-spatial or spatial feature of centrally-presented location words. The discriminability of the spatial feature of the word (Experiment 1), or of both the spatial and non-spatial feature (Experiment 2), was manipulated. When the spatial feature of the word was task-irrelevant, lowering the discriminability of this feature reduced the compatibility effect. The compatibility effect was restored when the discriminability of both the task-relevant and task-irrelevant features were reduced together. Results provide further evidence for the temporal overlap account of compatibility effects. Furthermore, compatibility effects when the spatial information was task-relevant and those when the spatial information was task-irrelevant were moderately correlated with each other, suggesting a common underlying mechanism in both versions.
doi:10.1016/j.actpsy.2008.10.005
PMCID: PMC2677062  PMID: 19041085
Simon effect; stimulus-response compatibility; temporal overlap; automatic activation

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