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1.  Anomalous use of context during task preparation in schizophrenia: a magnetoencephalography study 
Biological psychiatry  2013;73(10):967-975.
Impaired ability to use contextual information to optimally prepare for tasks contributes to performance deficits in schizophrenia. We used magnetoencephalography (MEG) and an antisaccade task to investigate the neural basis of this deficit.
In patients with schizophrenia (n=25) and healthy controls (n=18) we examined the difference in preparatory activation to cues indicating an impending antisaccade or prosaccade. We analyzed activation for correct trials only and focused on the network for volitional ocular motor control – frontal eye field (FEF), dorsal anterior cingulate cortex (dACC), and the ventrolateral and dorsolateral prefrontal cortex (VLPFC, DLPFC).
Compared to controls, patients made more antisaccade errors and showed reduced differential preparatory activation in the dACC and increased differential preparatory activation in the VLPFC. In patients only, antisaccade error rates correlated with preparatory activation in the FEF, DLPFC, and VLPFC.
In schizophrenia, reduced differential preparatory activation of the dACC may reflect reduced signaling of the need for control. Greater preparatory activation in the VLPFC and the correlations of error rate with FEF, DLPFC, and VLPFC activation may reflect that patients who are more error-prone require stronger activation in these regions for correct performance. These findings provide the first evidence of abnormal task preparation, distinct from response generation, during volitional saccades in schizophrenia. We conclude that schizophrenia patients are impaired in using task cues to modulate cognitive control and that this contributes to deficits inhibiting prepotent but contextually inappropriate responses and to behavior that is stimulus-bound and error-prone rather than flexibly guided by context.
PMCID: PMC3641151  PMID: 23380717
schizophrenia; antisaccade; frontal eye field; anterior cingulate cortex; lateral prefrontal cortex; cognitive control
2.  Preparatory neural networks are impaired in adults with attention-deficit/hyperactivity disorder during the antisaccade task☆ 
NeuroImage : Clinical  2012;2:63-78.
Adults with attention-deficit/hyperactivity disorder (ADHD) often display executive function impairments, particularly in inhibitory control. The antisaccade task, which measures inhibitory control, requires one to suppress an automatic prosaccade toward a salient visual stimulus and voluntarily make an antisaccade in the opposite direction. ADHD patients not only have longer saccadic reaction times, but also make more direction errors (i.e., a prosaccade was executed toward the stimulus) during antisaccade trials. These deficits may stem from pathology in several brain areas that are important for executive control. Using functional MRI with a rapid event-related design, adults with combined subtype of ADHD (coexistence of attention and hyperactivity problems), who abstained from taking stimulant medication 20 h prior to experiment onset, and age-match controls performed pro- and antisaccade trials that were interleaved with pro- and anti-catch trials (i.e., instruction was presented but no target appeared, requiring no response). This method allowed us to examine brain activation patterns when participants either prepared (during instruction) or executed (after target appearance) correct pro or antisaccades. Behaviorally, ADHD adults displayed several antisaccade deficits, including longer and more variable reaction times and more direction errors, but saccade metrics (i.e., duration, velocity, and amplitude) were normal. When preparing to execute an antisaccade, ADHD adults showed less activation in frontal, supplementary, and parietal eye fields, compared to controls. However, activation in these areas was normal in the ADHD group during the execution of a correct antisaccade. Interestingly, unlike controls, adults with ADHD produced greater activation than controls in dorsolateral prefrontal cortex during antisaccade execution, perhaps as part of compensatory mechanisms to optimize antisaccade production. Overall, these data suggest that the saccade deficits observed in adults with ADHD do not result from an inability to execute a correct antisaccade but rather the failure to properly prepare (i.e., form the appropriate task set) for the antisaccade trial. The data support the view that the executive impairments, including inhibitory control, in ADHD adults are related to poor response preparation.
► The neural correlates of inhibitory control in adults with ADHD were examined. ► We used an interleaved pro and antisaccade task simultaneously with functional MRI. ► This enabled the dissociation of automatic versus voluntary control. ► Patients had less activity in fronto-parietal areas during antisaccade preparation. ► Overall, antisaccade deficits in ADHD adults likely arise from poor preparation.
PMCID: PMC3777763  PMID: 24179760
ADHD; Saccade; fMRI; Preparation; Inhibition
3.  Attention, intention, and strategy in preparatory control 
Neuropsychologia  2009;47(7):1670-1685.
The neural mechanisms underlying different forms of preparatory control were examined using event-related fMRI. Preparatory brain activation was monitored in relation to different types of advance information: (1) random task cues indicating which of two possible tasks to perform upon subsequent target presentation, (2) task-ambiguous target stimuli, or 3) targets for which the correct response could be pre-determined. Three types of activation pattern were observed in different brain regions. First, more posterior regions of lateral prefrontal cortex (LPFC) and parietal cortex were activated by both advance task cues and advance targets, but with increased and more sustained activation for the latter. Second, more anterior regions of LPFC and parietal cortex were selectively activated by advance targets. Importantly, in these regions preparatory activation was not further modulated by the availability of advance response information. In contrast, preparatory activation in a third set of brain regions, including medial frontal cortex, reflected the utilization of advance response information, but by only a subset of participants. These results suggest three types of preparatory control: attentional (stimulus-oriented), intentional (action-oriented), and a possibly strategic component that might determine inter-individual differences in response readiness. Notably, the absence of regions selectively or even preferentially activated during cue-based preparation argues against certain conceptualizations of task-selective attention under cued task switching conditions.
PMCID: PMC2674875  PMID: 19397862
4.  Disconnectivity of the Cortical Ocular Motor Control Network in Autism Spectrum Disorders 
Neuroimage  2012;61(4):1226-1234.
Response inhibition, or the suppression of prepotent but contextually inappropriate behaviors, is essential to adaptive, flexible responding. Individuals with autism spectrum disorders (ASD) consistently show deficient response inhibition during antisaccades. In our prior functional MRI study, impaired antisaccade performance was accompanied by reduced functional connectivity between the frontal eye field (FEF) and dorsal anterior cingulate cortex (dACC), regions critical to volitional ocular motor control. Here we employed magnetoencephalography (MEG) to examine the spectral characteristics of this reduced connectivity. We focused on coherence between FEF and dACC during the preparatory period of antisaccade and prosaccade trials, which occurs after the presentation of the task cue and before the imperative stimulus. We found significant group differences in alpha band mediated coherence. Specifically, neurotypical participants showed significant alpha band coherence between the right inferior FEF and right dACC and between the left superior FEF and bilateral dACC across antisaccade, prosaccade, and fixation conditions. Relative to the neurotypical group, ASD participants showed reduced coherence between these regions in all three conditions. Moreover, while neurotypical participants showed increased coherence between the right inferior FEF and the right dACC in preparation for an antisaccade compared to a prosaccade or fixation, ASD participants failed to show a similar increase in preparation for the more demanding antisaccade. These findings demonstrate reduced long-range functional connectivity in ASD, specifically in the alpha band. The failure in the ASD group to increase alpha band coherence with increasing task demand may reflect deficient top-down recruitment of additional neural resources in preparation to perform a difficult task.
PMCID: PMC3376722  PMID: 22433660
Autism; ASD; Connectivity; Alpha Synchrony; MEG
5.  The impact of preparatory activities on medical school selection outcomes: a cross-sectional survey of applicants to the university of Adelaide medical school in 2007 
BMC Medical Education  2013;13:159.
Selection into medical school is highly competitive with more applicants than places. Little is known about the preparation that applicants undertake for this high stakes process. The study aims to determine what preparatory activities applicants undertake and what difficulties they encounter for each stage of the application process to medical school and in particular what impact these have on the outcome.
A cross-sectional survey of 1097 applicants who applied for a place in the University of Adelaide Medical School in 2007 and participated in the UMAT (Undergraduate Medicine and Health Sciences Admission Test) and oral assessment components of the selection process. The main outcome measures were an offer of an interview and offer of a place in the medical school and were analysed using logistic regression.
The odds of a successful outcome increased with each additional preparatory activity undertaken for the UMAT (odds ratio 1.22, 95% confidence interval 1.11 to 1.33; P < 0.001) and the oral assessment (1.36, 1.19 to 1.55; P < 0.001) stage of selection. The UMAT preparatory activities associated with the offer of an interview were attendance of a training course by a private organisation (1.75, 1.35 to 2.27: P < 0.001), use of online services of a private organisation (1.58, 1.23 to 2.04; P < 0.001), and familiarising oneself with the process (1.52, 1.15 to 2.00; p = 0.021). The oral assessment activities associated with an offer of a place included refining and learning a personal resume (9.73, 2.97 to 31.88; P < 0.001) and learning about the course structure (2.05, 1.29 to 3.26; P = 0.022).
For the UMAT, applicants who found difficulties with learning for this type of test (0.47, 0.35 to 0.63: P < 0.001), with the timing of UMAT in terms of school exams (0.48, 0.5 to 0.66; P < 0.001) and with the inability to convey personal skills with the UMAT (0.67, 0.52 to 0.86; P = 0.026) were significantly less likely to be offered an interview.
Medical schools make an enormous effort to undertake a selection process that is fair and equitable and which selects students most appropriate for medical school and the course they provide. Our results indicate that performance in the selection processes can be improved by training. However, if these preparatory activities may be limited to those who can access them, the playing field is not even and increasing equity of access to medical schools will not be achieved.
PMCID: PMC4220562  PMID: 24289521
Medical school selection; Admissions; Equity; Preparation
6.  Incentive effect on inhibitory control in adolescents with early-life stress: an antisaccade study 
Child abuse & neglect  2012;36(3):217-225.
Early-life stress (ES) such as adoption, change of caregiver, or experience of emotional neglect [0]may influence the way in which affected individuals respond to emotional stimuli of positive or negative valence. These modified responses may stem from a direct alteration of how emotional stimuli are coded, and/or the cognitive function implicated in emotion modulation, such as self-regulation or inhibition. These ES effects have been probed on tasks either targeting reward and inhibitory function. Findings revealed deficits in both reward processing and inhibitory control in ES youths. However, no work has yet examined whether incentives can improve automatic response or inhibitory control in ES youths.
To determine whether incentives would only improve self-regulated voluntary actions or generalize to automated motoric responses, participants were tested on a mixed eye movement task that included reflex-like prosaccades and voluntary controlled antisaccade eye movements. Seventeen adopted children (10 females, mean age 11.3 years) with a documented history of neglect and 29 typical healthy youths (16 females, mean age 11.9 years) performed the mixed prosaccade/antisaccade task during monetary incentive conditions or during no-incentive conditions.
Across both saccade types, ES adolescents responded more slowly than controls. As expected, control participants committed fewer errors on antisaccades during the monetary incentive condition relative to the no-incentive condition. By contrast, ES youths failed to show this incentive-related improvement on inhibitory control. No significant incentive effects were found with prepotent prosaccades trials in either group. Finally, co-morbid psychopathology did not modulate the findings.
These data suggest that youths with experience of early stress exhibit deficient modulation of inhibitory control by reward processes, in tandem with a reward-independent deficit in preparation for both automatic and controlled responses. These data may be relevant to interventions in ES youths.
PMCID: PMC3320703  PMID: 22425696
reward; antisaccade; cognitive control; early adversity; stress
It is shown in this paper that homologous immune sera are able to neutralize the B. typhosus skin-preparatory factors. The neutralization experiments were performed on a large number of rabbits, at least ten rabbits which showed positive control reactions being used for the titration of each serum. The rabbits into which the mixtures of B. typhosus culture filtrates with immune sera were injected can be divided into the following categories: those showing complete neutralization in highest dilutions (HN), those showing complete neutralization only in lower dilutions (LN) and those showing no neutralization (NN). The results indicate that the potency of a given serum as measured by the method outlined above has a direct relation to the reactions obtained in these groups of rabbits. For practical purposes the highest dilution of the serum which gives complete neutralization of the B. typhosus skin-preparatory factors (HN titer) may be taken as the actual titer of the serum as expressed in terms of their neutralization. The occurrence of a phenomenon suggestive of the prozone reaction is demonstrated. It also appears that the filtrates possess an antigenicity equal to that of dead and live bacteria. The studies on normal sera bring out the fact that normal sera fail to neutralize the B. typhosus skin-preparatory factors unless agglutinins can be demonstrated for B. typhosus. No normal sera have thus far been obtained which neutralized the skin-preparatory factors yet contained no B. typhosus agglutinins, but there were sera which contained these agglutinins but failed to neutralize the skin-preparatory factors. Some of the normal animals whose sera failed to neutralize the skin-preparatory factors were subsequently injected with B. typhosus culture filtrate and responded with neutralizing sera of high titer. Several heterologous sera were also investigated namely, scarlet fever, erysipelas, Shiga bacillus, Flexner bacillus, Mt. Desert bacillus, P. coli and B. avicida. These did not neutralize the B. typhosus skin-preparatory factors. Paratyphosus A and B sera on the other hand produced neutralization in various proportions. And the rabbits into which the serum-filtrate mixtures were injected could also be divided according to the results obtained into the same three groups as those with B. typhosus sera. It is not known yet whether this neutralization is a group reaction or whether the skin-preparatory factors are identical with those of B. typhosus. It would appear from these studies that a method is available for the quantitative titration of substances in the serum which neutralize the skin-preparatory factors of local skin reactivity to B. typhosus culture filtrates. It should be emphasized that it is possible to control the individual susceptibility of rabbits to this phenomenon. The method should permit of considerable accuracy in the quantitative titration of the neutralizing properties of a serum when a standardized procedure is developed. Experiments are under way to determine whether the method can be applied to the preparation of therapeutic sera. Work is also in progress to determine the effect of specific antisera upon B. typhosus skin-reacting factors introduced by the intravenous route.
PMCID: PMC2131566  PMID: 19869566
8.  Electrophysiological evidence for abnormal preparatory states and inhibitory processing in adult ADHD 
Attention deficit hyperactivity disorder (ADHD) is a common neurodevelopmental disorder that starts in childhood and frequently persists in adults. Several theories postulate deficits in ADHD that have effects across many executive functions or in more narrowly defined aspects, such as response inhibition. Electrophysiological studies on children, however, indicate that ADHD is not associated with a core deficit of response inhibition, as abnormal inhibitory processing is typically preceded or accompanied by other processing deficits. It is not yet known if this pattern of abnormal processing is evident in adult ADHD.
The objective of this paper was to investigate event-related potential indices of preparatory states and subsequent response inhibition processing in adults with ADHD. Two cued continuous performance tasks were presented to 21 adults meeting current criteria for adult ADHD and combined type ADHD in childhood, and 20 controls.
The ADHD group exhibited significantly weaker orienting attention to cues, cognitive preparation processes and inhibitory processing. In addition, we observed a strong correlation between the resources allocated to orienting to cues and the strength of the subsequent response strength control processes, suggesting that orienting deficits partly predict and determine response control deficits in ADHD.
These findings closely resemble those previously found in children with ADHD, which indicate that there is not a core response inhibition deficit in ADHD. These findings therefore suggest the possibility of developmental stability into adulthood of the underlying abnormal processes in ADHD.
PMCID: PMC2988695  PMID: 21029446
9.  Left Posterior Parietal Cortex Participates in Both Task Preparation and Episodic Retrieval 
Neuroimage  2009;46(4):1209-1221.
Optimal memory retrieval depends not only on the fidelity of stored information, but also on the attentional state of the subject. Factors such as mental preparedness to engage in stimulus processing can facilitate or hinder memory retrieval. The current study used functional magnetic resonance imaging (fMRI) to distinguish preparatory brain activity before episodic and semantic retrieval tasks from activity associated with retrieval itself. A catch-trial imaging paradigm permitted separation of neural responses to preparatory task cues and memory probes. Episodic and semantic task preparation engaged a common set of brain regions, including the bilateral intraparietal sulcus (IPS), left fusiform gyrus (FG), and the pre-supplementary motor area (pre-SMA). In the subsequent retrieval phase, the left IPS was among a set of frontoparietal regions that responded differently to old and new stimuli. In contrast, the right IPS responded to preparatory cues with little modulation during memory retrieval. The findings support a strong left-lateralization of retrieval success effects in left parietal cortex, and further indicate that left IPS performs operations that are common to both task preparation and memory retrieval. Such operations may be related to attentional control, monitoring of stimulus relevance, or retrieval.
PMCID: PMC3339571  PMID: 19285142
10.  Neural Processes of Preparatory Control for Stop Signal Inhibition 
Human brain mapping  2011;33(12):2785-2796.
This study investigated the preparatory control of motor inhibition and motor execution using a stop signal task (SST) and functional magnetic resonance imaging (fMRI). In the SST, a frequent “go” signal triggered a prepotent response and a less frequent “stop” signal prompted the inhibition of this response. Preparatory control of motor inhibition and execution in the stop signal trials were examined by contrasting brain activation between stop success and stop error trials during the fore-period, in which participants prepared to respond to go or to stop. Results from 91 healthy adults showed greater activation in the right prefrontal cortex and inferior parietal lobule during preparatory motor inhibition. Preparatory motor execution activated bilateral putamen, primary motor cortices, posterior cingulate cortex, ventromedial prefrontal cortex, and superior temporal/intraparietal sulci. Furthermore, the extents of these inhibition and execution activities were inversely correlated across subjects. On the basis of a median split of the stop signal reaction time (SSRT), subjects with short SSRT showed greater activity in the right orbital frontal cortex during preparatory inhibition. These new findings suggest that the go and stop processes interact prior to target presentation in the SST, in accord with recent computational models of stop signal inhibition.
PMCID: PMC3293936  PMID: 21976392
stop signal; inhibitory control; proactive; prefrontal; race model; go/no-go
11.  Does Explicit Expectation Really Affect Preparation? 
Expectation enables preparation for an upcoming event and supports performance if the anticipated situation occurs, as manifested in behavioral effects (e.g., decreased RT). However, demonstrating coincidence between expectation and preparation is not sufficient for attributing a causal role to the former. The content of explicit expectation may simply reflect the present preparation state. We targeted this issue by experimentally teasing apart demands for preparation and explicit expectations. Expectations often originate from our experience: we expect that events occurring with a high frequency in the past are more likely to occur again. In addition to expectation, other task demands can feed into action preparation. In four experiments, frequency-based expectation was pitted against a selective response deadline. In a three-choice reaction time task, participants responded to stimuli that appeared with varying frequency (60, 30, 10%). Trial-by-trial stimulus expectations were either captured via verbal predictions or induced by visual cues. Predictions as well as response times quickly conformed to the variation in stimulus frequency. After two (of five) experimental blocks we forced participants by selective time pressure to respond faster to a less frequent stimulus. Therefore, participants had to prepare for one stimulus (medium frequency) while often explicitly expecting a different one (high frequency). Response times for the less frequent stimulus decreased immediately, while explicit expectations continued to indicate the (unchanged) presentation frequencies. Explicit expectations were thus not just reflecting preparation. In fact, participants responded faster when the stimulus matched the trial-wise expectations, even when task demands discouraged their use. In conclusion, we argue that explicit expectation feeds into preparatory processes instead of being a mere by-product.
PMCID: PMC3521289  PMID: 23248606
explicit expectation; action control; anticipation; preparation; task goals
12.  Distributed representations of the "preparatory set" in the frontal oculomotor system: a TMS study 
BMC Neuroscience  2008;9:89.
The generation of saccades is influenced by the level of "preparatory set activity" in cortical oculomotor areas. This preparatory activity can be examined using the gap-paradigm in which a temporal gap is introduced between the disappearance of a central fixation target and the appearance of an eccentric target.
Ten healthy subjects made horizontal pro- or antisaccades in response to lateralized cues after a gap period of 200 ms. Single-pulse transcranial magnetic stimulation (TMS) was applied to the dorsolateral prefrontal cortex (DLPFC), frontal eye field (FEF), or supplementary eye field (SEF) of the right hemisphere 100 or 200 ms after the disappearance of the fixation point. Saccade latencies were measured to probe the disruptive effect of TMS on saccade preparation. In six individuals, we gave realistic sham TMS during the gap period to mimic auditory and somatosensory stimulation without stimulating the cortex.
TMS to DLPFC, FEF, or SEF increased the latencies of contraversive pro- and antisaccades. This TMS-induced delay of saccade initiation was particularly evident in conditions with a relatively high level of preparatory set activity: The increase in saccade latency was more pronounced at the end of the gap period and when participants prepared for prosaccades rather than antisaccades. Although the "lesion effect" of TMS was stronger with prefrontal TMS, TMS to FEF or SEF also interfered with the initiation of saccades. The delay in saccade onset induced by real TMS was not caused by non-specific effects because sham stimulation shortened the latencies of contra- and ipsiversive anti-saccades, presumably due to intersensory facilitation.
Our results are compatible with the view that the "preparatory set" for contraversive saccades is represented in a distributed cortical network, including the contralateral DLPFC, FEF and SEF.
PMCID: PMC2564971  PMID: 18801205
13.  Changes in corticospinal excitability and the direction of evoked movements during motor preparation: A TMS study 
BMC Neuroscience  2008;9:51.
Preparation of the direction of a forthcoming movement has a particularly strong influence on both reaction times and neuronal activity in the primate motor cortex. Here, we aimed to find direct neurophysiologic evidence for the preparation of movement direction in humans. We used single-pulse transcranial magnetic stimulation (TMS) to evoke isolated thumb-movements, of which the direction can be modulated experimentally, for example by training or by motor tasks. Sixteen healthy subjects performed brisk concentric voluntary thumb movements during a reaction time task in which the required movement direction was precued. We assessed whether preparation for the thumb movement lead to changes in the direction of TMS-evoked movements and to changes in amplitudes of motor-evoked potentials (MEPs) from the hand muscles.
When the required movement direction was precued early in the preparatory interval, reaction times were 50 ms faster than when precued at the end of the preparatory interval. Over time, the direction of the TMS-evoked thumb movements became increasingly variable, but it did not turn towards the precued direction. MEPs from the thumb muscle (agonist) were differentially modulated by the direction of the precue, but only in the late phase of the preparatory interval and thereafter. MEPs from the index finger muscle did not depend on the precued direction and progressively decreased during the preparatory interval.
Our data show that the human corticospinal movement representation undergoes progressive changes during motor preparation. These changes are accompanied by inhibitory changes in corticospinal excitability, which are muscle specific and depend on the prepared movement direction. This inhibition might indicate a corticospinal braking mechanism that counteracts any preparatory motor activation.
PMCID: PMC2453131  PMID: 18559096
14.  Functionally dissociating temporal and motor components of response preparation in left intraparietal sulcus 
Neuroimage  2011;54(2-3):1221-1230.
To optimise speed and accuracy of motor behaviour, we can prepare not only the type of movement to be made but also the time at which it will be executed. Previous cued reaction-time paradigms have shown that anticipating the moment in time at which this response will be made (“temporal orienting”) or selectively preparing the motor effector with which an imminent response will be made (motor intention or “motor orienting”) recruits similar regions of left intraparietal sulcus (IPS), raising the possibility that these two preparatory processes are inextricably co-activated. We used a factorial design to independently cue motor and temporal components of response preparation within the same experimental paradigm. By differentially cueing either ocular or manual response systems, rather than spatially lateralised responses within just one of these systems, potential spatial confounds were removed. We demonstrated that temporal and motor orienting were behaviourally dissociable, each capable of improving performance alone. Crucially, fMRI data revealed that temporal orienting activated the left IPS even if the motor effector that would be used to execute the response was unpredictable. Moreover, temporal orienting activated left IPS whether the target required a saccadic or manual response, and whether this response was left- or right-sided, thus confirming the ubiquity of left IPS activation for temporal orienting. Finally, a small region of left IPS was also activated by motor orienting for manual, though not saccadic, responses. Despite their functional independence therefore, temporal orienting and manual motor orienting nevertheless engage partially overlapping regions of left IPS, possibly reflecting their shared ontogenetic roots.
Research Highlights
►Anticipating when and how to respond can be functionally and neurally dissociated. ►Temporal orienting (when) engages left IPS for both oculomotor and manual responses. ►Motor orienting (how) engages left IPS only for manual, not oculomotor, responses. ►Manual action circuits may be functionally recycled for anticipating when to respond.
PMCID: PMC3025354  PMID: 20868756
Motor attention; Motor preparation; Motor intention; Temporal preparation; Supramarginal gyrus
15.  Longitudinal Growth Curves of Brain Function Underlying Inhibitory Control through Adolescence 
The Journal of Neuroscience  2013;33(46):18109-18124.
Neuroimaging studies suggest that developmental improvements in inhibitory control are primarily supported by changes in prefrontal executive function. However, studies are contradictory with respect to how activation in prefrontal regions changes with age, and they have yet to analyze longitudinal data using growth curve modeling, which allows characterization of dynamic processes of developmental change, individual differences in growth trajectories, and variables that predict any interindividual variability in trajectories. In this study, we present growth curves modeled from longitudinal fMRI data collected over 302 visits (across ages 9 to 26 years) from 123 human participants. Brain regions within circuits known to support motor response control, executive control, and error processing (i.e., aspects of inhibitory control) were investigated. Findings revealed distinct developmental trajectories for regions within each circuit and indicated that a hierarchical pattern of maturation of brain activation supports the gradual emergence of adult-like inhibitory control. Mean growth curves of activation in motor response control regions revealed no changes with age, although interindividual variability decreased with development, indicating equifinality with maturity. Activation in certain executive control regions decreased with age until adolescence, and variability was stable across development. Error-processing activation in the dorsal anterior cingulate cortex showed continued increases into adulthood and no significant interindividual variability across development, and was uniquely associated with task performance. These findings provide evidence that continued maturation of error-processing abilities supports the protracted development of inhibitory control over adolescence, while motor response control regions provide early-maturing foundational capacities and suggest that some executive control regions may buttress immature networks as error processing continues to mature.
PMCID: PMC3828464  PMID: 24227721
16.  Costs of a Predictable Switch Between Simple Cognitive Tasks Following Severe Closed-Head Injury 
Neuropsychology  2006;20(6):675-684.
The authors used a predictable, externally cued task-switching paradigm to investigate executive control in a severe closed-head injury (CHI) population. Eighteen individuals with severe CHI and 18 controls switched between classifying whether a digit was odd or even and whether a letter was a consonant or vowel on every 4th trial. The target stimuli appeared in a circle divided into 8 equivalent parts. Presentation of the stimuli rotated clockwise. Participants performed the switching task at both a short (200 ms) and a long (1,000 ms) preparatory interval. Although the participants with CHI exhibited slower response times and greater switch costs, similar to controls, additional preparatory time reduced the switch costs, and the switch costs were limited to the 1st trial in the run. These findings indicate that participants with severe CHI were able to take advantage of time to prepare for the task switch, and the executive control processes involved in the switch costs were completed before the 1st trial of the run ended.
PMCID: PMC1779821  PMID: 17100512
closed-head injury; traumatic brain injury; executive functions; task switching; set shifting
17.  Effects of Magnitude and Magnitude Predictability of Postural Perturbations on Preparatory Cortical Activity in Older Adults with and without Parkinson’s Disease 
The goal of this study was to identify whether impaired cortical preparation may relate to impaired scaling of postural responses of people with Parkinson’s disease (PD). We hypothesized that impaired scaling of postural responses in participants with PD would be associated with impaired set-dependent cortical activity in preparation for perturbations of predictable magnitudes. Participants performed postural responses to backward surface translations. We examined the effects of perturbation magnitude (predictable small vs. predictable large) and predictability of magnitude (predictable vs. unpredictable-in-magnitude) on postural responses (center-of-pressure (CoP) displacements) and on preparatory electroencephalographic (EEG) measures of contingent negative variation (CNV) and alpha and beta event-related desynchronization (ERD). Our results showed that unpredictability of perturbation magnitude, but not the magnitude of the perturbation itself, was associated with increased CNV amplitude at the CZ electrode in both groups. While control participants scaled their postural responses to the predicted magnitude of the perturbation, their condition-related changes in CoP displacements were not correlated with condition-related changes in EEG preparatory activity (CNV or ERD). In contrast, participants with PD did not scale their postural responses to the predicted magnitude of the perturbation, but they did demonstrate greater beta ERD in the condition of predictably small-magnitude perturbations and greater beta ERD than the control participants at the CZ electrode. In addition, increased beta ERD in PD was associated with decreased adaptability of postural responses, suggesting that preparatory cortical activity may have a more direct influence on postural response scaling for people with PD than for control participants.
PMCID: PMC3472959  PMID: 22936099
Parkinson’s disease; electroencephalography; contingent negative variation; event related desynchronization; posture; preparation
18.  Preparatory Allocation of Attention and Adjustments in Conflict Processing 
NeuroImage  2007;35(2):949-958.
Attentional control involves the ability to allocate preparatory attention to improve subsequent stimulus processing and response selection. There is behavioral evidence to support the hypothesis that increased expectancy of stimulus and response conflict may decrease the subsequent experience of conflict during task performance. We used a cued Flanker and event-related fMRI design to separate processes involved in preparation from those involved in resolving conflict, and to identify the brain systems involved in these processes as well as the association between preparatory activity levels and activity related to subsequent conflict processing. Our results demonstrate that preparatory attentional allocation following a cue to the upcoming level of conflict is mediated by a network involving Dorsolateral Prefrontal Cortex (DLPFC) and the Intraparietal Sulcus (IPS). Informed preparation for conflict processing was associated with decreased Anterior Cingulate Cortex/preSupplementary Motor Area (ACC/preSMA) and IPS activity during the flanker target presentation, supporting their roles in conflict processing and visuospatial attention during the flanker task. Ventrolateral Prefrontal Cortex/Orbitofrontal Cortex (VLPFC/OFC) was active when specific strategic task rule and outcome information was available.
PMCID: PMC2859823  PMID: 17258912
19.  EEG alpha activity reflects motor preparation rather than the mode of action selection 
Alpha-band activity (8–13 Hz) is not only suppressed by sensory stimulation and movements, but also modulated by attention, working memory and mental tasks, and could be sensitive to higher motor control functions. The aim of the present study was to examine alpha oscillatory activity during the preparation of simple left or right finger movements, contrasting the external and internal mode of action selection. Three preparation conditions were examined using a precueing paradigm with S1 as the preparatory and S2 as the imperative cue: Full, laterality instructed by S1; Free, laterality freely selected and None, laterality instructed by S2. Time-frequency (TF) analysis was performed in the alpha frequency range during the S1–S2 interval, and alpha motor-related amplitude asymmetries (MRAA) were also calculated. The significant MRAA during the Full and Free conditions indicated effective external and internal motor response preparation. In the absence of specific motor preparation (None), a posterior alpha event-related desynchronization (ERD) dominated, reflecting the main engagement of attentional resources. In Full and Free motor preparation, posterior alpha ERD was accompanied by a midparietal alpha event-related synchronization (ERS), suggesting a concomitant inhibition of task-irrelevant visual activity. In both Full and Free motor preparation, analysis of alpha power according to MRAA amplitude revealed two types of functional activation patterns: (1) a motor alpha pattern, with predominantly midparietal alpha ERS and large MRAA corresponding to lateralized motor activation/visual inhibition and (2) an attentional alpha pattern, with dominating right posterior alpha ERD and small MRAA reflecting visuospatial attention. The present results suggest that alpha oscillatory patterns do not resolve the selection mode of action, but rather distinguish separate functional strategies of motor preparation.
PMCID: PMC3418545  PMID: 22912607
motor preparation; motor selection; externally-cued action; internally-cued action; alpha-band activity; motor-related amplitude asymmetry
20.  Evidence for two concurrent inhibitory mechanisms during response preparation 
Inhibitory mechanisms are critically involved in goal-directed behaviors. To gain further insight into how such mechanisms shape motor representations during response preparation, motor evoked potentials (MEPs) elicited by transcranial magnetic stimulation (TMS) and H-reflexes were recorded from left hand muscles during choice reaction time tasks. The imperative signal, which indicated the required response, was always preceded by a preparatory cue. During the post-cue delay period, left MEPs were suppressed when the left hand had been cued for the forthcoming response, suggestive of a form of inhibition specifically directed at selected response representations. H-reflexes were also suppressed on these trials, indicating that the effects of this inhibition extend to spinal circuits. In addition, left MEPs were suppressed when the right hand was cued, but only when left hand movements were a possible response option before the onset of the cue. Notably, left hand H-reflexes were not modulated on these trials, consistent with a cortical locus of inhibition that lowers the activation of task-relevant, but non-selected responses. These results suggest the concurrent operation of two inhibitory mechanisms during response preparation: one decreases the activation of selected responses at the spinal level, helping to control when selected movements should be initiated by preventing their premature release; a second, upstream mechanism helps to determine what response to make during a competitive selection process.
PMCID: PMC2852647  PMID: 20220014
cognitive control; competition; corticospinal excitability; decision-making; response selection; transcranial magnetic stimulation; H-reflex
A phenomenon of local skin reactivity to B. typhosus culture filtrates is described in this report. The reactivity was induced by skin injections of the filtrate followed 24 hours later by an intravenous injection of the same filtrate. The local response consisted of severe hemorrhagic necrosis and was fully developed 4 to 5 hours after the second injection. About 78 to 79 per cent of the rabbits employed were susceptible to this phenomenon. Different areas of the skin of the abdomen, when similarly treated, responded with equal severity to the intravenous injection. There were variations in the size of different areas in the same animals. The intensity and size of the local hemorrhagic reactions were not related to the intensity of the erythema produced by the preparatory skin injections. Following intravenous injection very severe hemorrhagic reactions were obtained, in those areas which reacted negatively in this respect to the preparatory skin injections. Evidently, the local trauma produced by the preparatory skin injections was not responsible for the localization of the toxic factors introduced by the intravenous route. It was necessary to allow a short interval of time between the skin preparatory injections and the intravenous injection, for the reproduction of the phenomenon. An incubation period of 2 hours was insufficient. An interval of 24 hours was invariably sufficient. The ability to react disappeared in 48 hours after the preliminary skin injections. Repeated direct injections of the filtrate into the same areas of the skin, with an interval of 24 hours between the injections, did not result in reactions similar to the above described hemorrhagic necrosis. The second skin injection was followed by reddening, some swelling and a local accumulation of polymorphonuclear neutrophil leucocytes which showed no signs of necrobiosis. There was no rupture of blood vessels. Skin injections followed after a suitable interval by intravenous injection, were necessary for the reproduction of the severe local hemorrhagic response. Skin reactivity to B. typhosus culture filtrate injected intravenously was not induced by turpentine in various dilutions, sterile tryptic digest broth, culture filtrates of 4 strains of streptococci or by the Streptococcus erysipelatis toxin. It was possible to titrate the skin preparatory factors. Dilutions of the filtrate up to 1:64 were able to induce the local skin reactivity. But, whereas dilutions up to 1:4 invariably prepared the skin so that very severe hemorrhagic reactions followed the second injection in susceptible animals, dilutions from 1:8 to 1:64 were uncertain and their preparatory effect varied in different animals. The skin preparatory factors showed considerable heat resistance. The heat resistance varied with the strains employed. One strain produced factors totally resistant to heating in the autoclave for 1 hour. However, there was definite and unquestionable inactivation of these factors as derived from other strains when the filtrate was diluted 1:2 and heated in the autoclave. Various hydrogen ion concentrations in the range from 9.0 to 4.0 had no effect upon the skin preparatory factors. Heat resistance was not modified by the various pH within this range. The mechanism of the phenomenon described has not been fully studied as yet. An experimental comparison of it with the manifestations of bacterial allergy of the skin is necessary. There are certain features, however, which considered together, distinguish this phenomenon from the known phenomena of bacterial hypersusceptibility. These features are: local reactivity; the short incubation period necessary to induce the local reactivity; the short duration of the state of reactivity; the ability to induce local reactivity by a single skin injection; the severity of the reaction; and the necessity to make the second injection of the toxic agent by the intravenous route. Studies on the relation of specific antisera to the phenomenon described are under way.
PMCID: PMC2131457  PMID: 19869481
22.  ‘Put on your poker face’: neural systems supporting the anticipation for expressive suppression and cognitive reappraisal 
It is a unique human ability to regulate negative thoughts and feelings. Two well-investigated emotion-regulation strategies (ERSs), cognitive reappraisal and expressive suppression, are associated with overlapping prefrontal neural correlates, but differ temporally during the emotion-generation process. Although functional imaging studies have mainly investigated these ERS as a reaction to an emotion-inducing event, the intention to regulate upcoming negative emotions might already be associated with differences in neural activity. Hence, event-related functional magnetic resonance imaging was recorded in 42 participants while they completed an emotion-regulation paradigm. During this task, participants were instructed to proactively prepare to use a specific ERS knowing that a negative, high-arousing image would appear after the preparation period. As expected, the results demonstrated prefrontal and parietal activation while participants were suppressing or reappraising their emotions (family-wise error (FWE)-corrected). The intention to suppress emotions was associated with increased activation in the right inferior frontal gyrus, bilateral putamen, pre-supplementary motor area and right supramarginal gyrus (FWE-corrected). This enhanced proactive inhibitory control: (i) predicted decreased motoric activity during the actual suppression of emotional expressions and (2) trended toward a significant association with how successfully participants suppressed their emotions. However, neural correlates of preparatory control for cognitive reappraisal were not observed, possibly because contextual cues about the upcoming emotional stimulus are necessary to proactively start to cognitively reinterpret the situation.
PMCID: PMC3831557  PMID: 22956675
emotion regulation; cognitive reappraisal; expressive suppression; proactive and reactive control
23.  Anterior cingulate neurons represent errors and preparatory attention within the same behavioral sequence 
The anterior cingulate cortex (ACC) has been implicated in both preparatory attention (i.e., selecting behaviorally relevant stimuli) and in detecting errors. We recorded from the rat ACC and medial prefrontal cortex (mPFC), which is functionally homologous with the primate dorsolateral PFC, during an attention task. The 3-choice serial reaction time task requires a rat to orient toward and divide attention between 3 brief (300 msec duration) light stimuli presented in random order across nose poke holes in an operant chamber. In both the ACC and mPFC, we found that neural activity was related to the level of preparatory (pre-cue) attention and subsequent correct or incorrect choice, in that the magnitude of the single units' response to the cue was lower on incorrect trials and was not different from baseline on unattended trials. This preparatory neural activity consisted of both excitatory and inhibitory phasic responses. The number of units responding to the cue was similarly graded, in that fewer units exhibited phasic responses to the cue on incorrect and unattended trials, compared to correct trials. Although preparatory activity was found in both the ACC and mPFC, activity after incorrect nose pokes, which may be related to error detection, were only observed in the ACC. Thus, during the same behavioral sequence, the ACC encodes both error-related events and preparatory attention, whereas the mPFC only participates in preparatory attention. The finding of substantial inhibitory activity during the preparatory period suggests a critical role for inhibition of pyramidal cells in PFC-mediated cognitive functions.
PMCID: PMC2730728  PMID: 19458213
attention-deficit hyperactivity disorder; dopamine; prefrontal cortex; schizophrenia; single unit; rat
24.  ERP correlates of shared control mechanisms involved in saccade preparation and in covert attention 
Brain Research  2007;1135(8-3):154-166.
We investigated whether attention shifts and eye movement preparation are mediated by shared control mechanisms, as claimed by the premotor theory of attention. ERPs were recorded in three tasks where directional cues presented at the beginning of each trial instructed participants to direct their attention to the cued side without eye movements (Covert task), to prepare an eye movement in the cued direction without attention shifts (Saccade task) or both (Combined task). A peripheral visual Go/Nogo stimulus that was presented 800 ms after cue onset signalled whether responses had to be executed or withheld. Lateralised ERP components triggered during the cue–target interval, which are assumed to reflect preparatory control mechanisms that mediate attentional orienting, were very similar across tasks. They were also present in the Saccade task, which was designed to discourage any concomitant covert attention shifts. These results support the hypothesis that saccade preparation and attentional orienting are implemented by common control structures. There were however systematic differences in the impact of eye movement programming and covert attention on ERPs triggered in response to visual stimuli at cued versus uncued locations. It is concluded that, although the preparatory processes underlying saccade programming and covert attentional orienting may be based on common mechanisms, they nevertheless differ in their spatially specific effects on visual information processing.
PMCID: PMC2248222  PMID: 17198687
Spatial attention; Eye movements; Attentional control; Vision; Event-related brain potentials
25.  ERP correlates of shared control mechanisms involved in saccade preparation and in covert attention 
Brain research  2007;1135(1):154-166.
We investigated whether attention shifts and eye movement preparation are mediated by shared control mechanisms, as claimed by the premotor theory of attention. ERPs were recorded in three tasks where directional cues presented at the beginning of each trial instructed participants to direct their attention to the cued side without eye movements (Covert task), to prepare an eye movement in the cued direction without attention shifts (Saccade task), or both (Combined task). A peripheral visual Go/Nogo stimulus that was presented 800 ms after cue onset signalled whether responses had to be executed or withheld. Lateralised ERP components triggered during the cue-target interval, which are assumed to reflect preparatory control mechanisms that mediate attentional orienting, were very similar across tasks. They were also present in the Saccade task, which was designed to discourage any concomitant covert attention shifts. These results support the hypothesis that saccade preparation and attentional orienting are implemented by common control structures. There were however systematic differences in the impact of eye movement programming and covert attention on ERPs triggered in response to visual stimuli at cued versus uncued locations. It is concluded that although the preparatory processes underlying saccade programming and covert attentional orienting may be based on common mechanisms, they nevertheless differ in their spatially specific effects on visual information processing.
PMCID: PMC2248222  PMID: 17198687
Spatial attention; Eye movements; Attentional control; Vision; Event-related brain potentials

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