Nonindependent and nonstationary response times in stopping and stepping saccade tasks

Saccade stop signal and target step tasks are used to investigate the mechanisms of cognitive control. Performance of these tasks can be explained as the outcome of a race between stochastic go and stop processes. The race model analyses assume that response times (RTs) measured throughout an experimental session are independent samples from stationary stochastic processes. This article demonstrates that RTs are neither independent nor stationary for humans and monkeys performing saccade stopping and target-step tasks. We investigate the consequences that this has on analyses of these data. Nonindependent and nonstationary RTs artificially flatten inhibition functions and account for some of the systematic differences in RTs following different types of trials. However, nonindependent and nonstationary RTs do not bias the estimation of the stop signal RT. These results demonstrate the robustness of the race model to some aspects of nonindependence and nonstationarity and point to useful extensions of the model.     

Changes in spinal reflex excitability in a countermanded timed response task

Subjects (N = 8) performed a timed response task in which they attempted to synchronize an impulsive foot-press response with the last in a series of four regularly spaced tones. In Experiment 1, the response was countermanded on one third of the trials (stop trials) by a stop signal that appeared at a predetermined delay after the third tone. No stop signal appeared on the remaining trials (go trials). All subjects showed a systematic transition from withholding the response on stop trials in which the stop signal appeared shortly after the third tone to executing the response on trials in which a single stop signal delay had been chosen so that a response would be made on about 50% of the stop trials. We elicited Hoffmann (H) reflexes from the soleus muscle on all trials to determine whether the reflexes were augmented on occasions on which a response was prepared but withheld. Mean H-reflex amplitudes on go trials and on stop trials on which the response was executed were similar and showed a marked augmentation beginning about 250 ms before response onset; mean H-reflex amplitudes on stop trials on which the response was withheld showed less pronounced augmentation. Inspection of individual H-reflex amplitudes revealed that on stop trials on which the response was withheld the reflexes could be augmented to the same extent as on trials on which the response was executed. This dissociation of H-reflex augmentation and response execution shows that H-reflex augmentation reflects a controlled response process. Ballistic response processes therefore must be limited to a brief duration.     

IMPULSIVITY AND INHIBITORY CONTROL

Abstract— We report an experiment testing the hypothesis that impulsive behavior reflects a deficit in the ability to inhibit prepotent responses Specifically, we examined whether impulsive people respond more slowly to signals to inhibit (stop signals) than non-impulsive people In this experiment, 136 undergraduate students completed an impulsivity questionnaire and then participated in a stop-signal experiment, in which they performed a choice reaction time (go) task and were asked to inhibit their responses to the go task when they heard a stop signal The delay between the go signal and the stop signal was determined by a tracking procedure designed to allow subjects to inhibit on 50% of the stop-signal trials. Reaction time to the go signal did not vary with impulsivity, but estimated stop-signal reaction time was longer in more impulsive subjects, consistent with the hypothesis and consistent with results from populations with pathological problems with impulse control.     

The interplay of stop signal inhibition and inhibition of return

Inhibition of return (IOR) refers to slower responding to a stimulus that appears in the same rather than a different location as that of a preceding stimulus. The goal of the present study was to examine the relationship between IOR and stop signal inhibition. Participants were presented with two stimuli (S1 and S2) on each trial. On half of the trials (go trials), participants were required to make a speeded button-press response to report the location of S1; on the other half of trials (stop trials), they were required to cancel the response to S1, as indicated by the appearance of a stop signal at a variable delay (stop signal delay, SSD) after the appearance of S1. Success in cancelling an S1 response varied directly as a function of the SSD: The longer the delay, the more difficult it was for participants to cancel the prepared response. We examined the magnitude of IOR in the S2 reaction times as a function of whether participants made a correct go response to S1, made an erroneous non-cancelled response to S1, or successfully cancelled a response to S1. Our results indicated that the presentation of a stop signal increased the magnitude of IOR, even when the S1 response was not successfully cancelled. However, this was true only when the to-be-cancelled response involved the same effectors as the response used to reveal IOR. These results suggest that there may be a motor component to IOR that is sensitive to the same inhibitory processes that are used to cancel responses in a stop signal paradigm.     

Developmental trends in simple and selective inhibition of compatible and incompatible responses

This study examined age-related change in the ability to inhibit responses using two varieties of the stop signal paradigm. Three age groups (29 7-year-olds, 24 10-year-olds, and 28 young adults) performed first on a visual choice reaction task in which the spatial mapping between the go signal and response was varied between blocks. The choice task was then complicated by randomly inserting a visual stop signal on 30% of the trials. In the simple stop signal paradigm, the stop signal required the inhibition of the planned response. In the selective stop signal paradigm, the stop signal required response inhibition only when the stop signal was presented at the same side as the instructed response to the go signal. The results showed that simple stopping was faster than selective stopping and that selective, but not simple, stopping of incompatible responses was slower than stopping of compatible responses. Brinley plot analysis yielded linear functions relating children's latencies to adults' latencies. Analysis of shared variance indicated that developmental change in the speed of selective stopping continued to be significant even when the effect associated with simple stopping was removed. This pattern of findings is discussed vis-à-vis notions of global versus specific developmental trends in the speed of information processing.     

Restraint and Cancellation: Multiple Inhibition Deficits in Attention Deficit Hyperactivity Disorder

We used variations of the stop signal task to study two components of motor response inhibition—the ability to withhold a strong response tendency (restraint) and the ability to cancel an ongoing action (cancellation)—in children with a diagnosis of attention deficit hyperactivity disorder (ADHD) and in non-ADHD controls of similar age (ages 7–14 years). The goal was to determine if restraint and cancellation were related and if both were deficient in ADHD. The stop signal task involved a choice reaction time task (go task) which required a rapid response. The demand for inhibitory control was invoked through the presentation of a stop signal on a subset of go trials which required that the ongoing response be suspended. The stop signal was presented either concurrently with the go signal (restraint version) or after a variable delay (cancellation version). In Study 1, we compared ADHD and control children on the cancellation version of the stop task; in Study 2, we compared ADHD and controls on the restraint version. In Study 3, a subset of ADHD and control participants completed both tasks so that we could examine convergence of these dimensions of inhibition. Compared to control participants, ADHD participants showed a deficit both in the ability to cancel and to restrain a speeded motor response. Performance on the restraint version was significantly correlated with performance on the cancellation version in controls, but not in ADHD participants. We conclude that ADHD is associated with deficits in both restraint and cancellation subcomponents of inhibition.     

Reduced response readiness delays stop signal inhibition

This study examines the effect of response readiness on the stopping of motor responses. Thirteen subjects performed a primary task requiring a speeded choice reaction on go trials and response inhibition on nogo trials. An occasional cue informed subjects that a nogo trial was imminent but left them uncertain about the number of go trials separating the cue and the upcoming nogo trial. This setup was meant to create test episodes of reduced response readiness (i.e., trial sequences initiated by the cue and terminated by the nogo signal) and control episodes, in which subjects were ready to execute a speeded choice reaction (i.e., trial sequences consisting only of go trials). During both episodes, a visual stop signal could occasionally and unpredictably follow go signal onset, instructing subjects to withhold their response to the go signal. Choice reactions on go trials were delayed during test episodes relative to control episodes. Most importantly, stop reactions were delayed, not facilitated, during test episodes compared to control episodes. These findings were taken to suggest that reduced readiness gives rise to more forceful responses that are then more difficult to inhibit.     

Effects of stop-signal probability in the stop-signal paradigm: the N2/P3 complex further validated

The aim of this study was to examine the effects of frequency of occurrence of stop signals in the stop-signal paradigm. Presenting stop signals less frequently resulted in faster reaction times to the go stimulus and a lower probability of inhibition. Also, go stimuli elicited larger and somewhat earlier P3 responses when stop signals occurred less frequently. Since the amplitude effect was more pronounced on trials when go signals were followed by fast than slow reactions, it probably reflected a stronger set to produce fast responses. N2 and P3 components to stop signals were observed to be larger and of longer latency when stop signals occurred less frequently. The amplitude enhancement of these N2 and P3 components were more pronounced for unsuccessful than for successful stop-signal trials. Moreover, the successfully inhibited stop trials elicited a frontocentral P3 whereas unsuccessfully inhibited stop trials elicited a more posterior P3 that resembled the classical P3b. P3 amplitude in the unsuccessfully inhibited condition also differed between waveforms synchronized with the stop signal and waveforms synchronized with response onset whereas N2 amplitude did not. Taken together these findings suggest that N2 reflected a greater significance of failed inhibitions after low probability stop signals while P3 reflected continued processing of the erroneous response after response execution.     

Response inhibition during perceptual decision making in humans and macaques

Response inhibition in stop signal tasks has been explained as the outcome of a race between GO and STOP processes (e.g., Logan, 1981). Response choice in two-alternative perceptual categorization tasks has been explained as the outcome of an accumulation of evidence for the alternative responses. To begin unifying these two powerful investigation frameworks, we obtained data from humans and macaque monkeys performing a stop signal task with responses guided by perceptual categorization and variable degrees of difficulty, ranging from low to high accuracy. Comparable results across species reinforced the validity of this animal model. Response times and errors increased with categorization difficulty. The probability of failing to inhibit responses on stop signal trials increased with stop signal delay, and the response times for failed stop signal trials were shorter than those for trials with no stop signal. Thus, the Logan race model could be applied to estimate the duration of the stopping process. We found that the duration of the STOP process did not vary across a wide range of discrimination accuracies. This is consistent with the functional, and possibly mechanistic, independence of choice and inhibition mechanisms.     

Stopping eye and hand movements: are the processes independent?

To explore how eye and hand movements are controlled in a stop task, we introduced effector uncertainty by instructing subjects to initiate and occasionally inhibit eye, hand, or eye + hand movements in response to a color-coded foveal or tone-coded auditory stop signal. Regardless of stop signal modality, stop signal reaction time was shorter for eye movements than for hand movements, but notably did not vary with knowledge about which movement to cancel. Most errors on eye + hand stopping trials were combined eye + hand movements. The probability and latency of signal respond eye and hand movements corresponded to predictions of Logan and Cowan's (1984) race model applied to each effector independently.     

Executive control of gaze by the frontal lobes

Executive control requires controlling the initiation of movements, judging the consequences of actions, and adjusting performance accordingly. We have investigated the role of different areas in the frontal lobe in executive control expressed by macaque monkeys performing a saccade stop signal task. Certain neurons in the frontal eye field respond to visual stimuli, and others control the production of saccadic eye movements. Neurons in the supplementary eye field do not control directly the initiation of saccades but, instead, signal the production of errors, the anticipation and delivery of reinforcement, and the presence of response conflict. Neurons in the anterior cingulate cortex signal the production of errors and the anticipation and delivery of reinforcement, but not the presence of response conflict. Intracranial local field potentials in the anterior cingulate cortex of monkeys indicate that these medial frontal signals can contribute to event-related potentials related to performance monitoring. Electrical stimulation of the supplementary eye field improves performance in the task by elevating saccade latency. An interactive race model shows how interacting units produce behavior that can be described as the outcome of a race between independent processes and how conflict between gaze-holding and gaze-shifting neurons can be used to adjust performance.     

On the ability to inhibit simple and choice reaction time responses: a model and a method

This article reports four experiments on the ability to inhibit responses in simple and choice reaction time (RT) tasks. Subjects responding to visually presented letters were occasionally presented with a stop signal (a tone) that told them not to respond on that trial. The major dependent variables were (a) the probability of inhibiting a response when the signal occurred, (b) mean and standard deviation (SD) of RT on no-signal trials, (c) mean RT on trials on which the signal occurred but subjects failed to inhibit, and (d) estimated RT to the stop signal. A model was proposed to estimated RT to the stop signal and to account for the relations among the variables. Its main assumption is that the RT process and the stopping process race, and response inhibition depends on which process finishes first. The model allows us to account for differences in response inhibition between tasks in terms of transformations of stop-signal delay that represent the relative finishing times of the RT process and the stopping process. The transformations specified by the model were successful in group data and in data from individual subjects, regardless of how delays were selected. The experiments also compared different methods of selecting stop-signal delays to equate the probability of inhibition in the two tasks.     

The effect of reinforcement variables on inhibition in children with ADHD.

Behavioral inhibition, often cited as a central deficit in children with ADHD, has been shown to change in response to reinforcement. In this preliminary investigation, children with ADHD (n= 20) and matched controls (ages 7 to 12) completed a new version of the stop signal paradigm, the Fire Fighter Game, a measure of inhibition of a prepotent motor response, under three conditions: (1) no reinforcement; (2) immediate reinforcement; and (3) delayed reinforcement. In all conditions, the stop signal reaction time (SSRT) of children with ADHD was consistently longer than controls. Both groups improved significantly with reinforcement, and there was no strong evidence that immediate reinforcement was more effective than delayed reinforcement. However, it appeared that the reason for the changes in SSRT in response to reinforcement differed between the groups. Children in the control group responded faster on go trials, whereas children with ADHD improved their ability to inhibit responding with shorter stop delays. The relevance of these findings is discussed in terms of current theories of ADHD.     

Inhibitory control in mind and brain: an interactive race model of countermanding saccades

The stop-signal task has been used to study normal cognitive control and clinical dysfunction. Its utility is derived from a race model that accounts for performance and provides an estimate of the time it takes to stop a movement. This model posits a race between go and stop processes with stochastically independent finish times. However, neurophysiological studies demonstrate that the neural correlates of the go and stop processes produce movements through a network of interacting neurons. The juxtaposition of the computational model with the neural data exposes a paradox-how can a network of interacting units produce behavior that appears to be the outcome of an independent race? The authors report how a simple, competitive network can solve this paradox and provide an account of what is measured by stop-signal reaction time.     

Redundancy gain in the stop-signal paradigm: implications for the locus of coactivation in simple reaction time

The authors carried out 2 experiments designed to cast light on the locus of redundancy gain in simple visual reaction time by using a stop-signal paradigm. In Experiment 1, the authors found that single visual stimuli were more easily inhibited than double visual stimuli by an acoustic stop signal. This result is in keeping with the idea that redundancy gain occurs prior to the ballistic stage of the stop-signal task. In Experiment 2, the authors found that the response to an acoustic go signal was more easily inhibited by a double than by a single visual stop signal. This result provides conclusive evidence for a redundancy gain in the stop process--in a process that does not involve a motor response but rather its inhibition.     

Inhibitory effects on response force in the stop-signal paradigm

The forcefulness of key press responses was measured in stop-all and selective stopping versions of the stop-signal paradigm. When stop signals were presented too late for participants to succeed in stopping their responses, response force was nonetheless reduced relative to trials in which no stop signal was presented. This effect shows that peripheral motor aspects of primary task responses can still be influenced by inhibition even when the stop signal arrives too late to prevent the response. It thus requires modification of race models in which responses in the presence of stop signals are either stopped completely or produced normally, depending on whether the responding or stopping process finishes first.     

The modification of an already-programmed response: a new interpretation of Henry and Harrison (1961)

In an influential study, Henry and Harrison (1961) examined the capability of subjects to inhibit an already-programmed response. Their experiment showed that when a stop signal was presented only 100 ms after the imperative go signal; a subject could not inhibit the movement. The inferences were that rapid ballistic actions are programmed and the program cannot be altered in a limited amount of time (see Schmidt, 1988). In the present note, we describe some forgotten data from the Henry and Harrison paper and, with a trend analysis, demonstrate that the stop signal does have an influence on the movement in a somewhat continuous fashion.     

The duration of response inhibition in the stop-signal paradigm varies with response force

In a previous study, we have found that the speed of stopping a response is delayed when response readiness is reduced by cuing the probability of no-go trials [Acta Psychol. 111 (2002) 155]. Other investigators observed that responses are more forceful when the probability to respond is low than when it is high (e.g. [Quart. J. Exp. Psychol. A 50 (1997) 405]). In this study, the hypothesis was tested that low probability responses are more forceful than high probability responses and that these responses are more difficult to stop. Subjects performed on a choice reaction task and on three tasks with respectively 100%, 80%, and 50% response probabilities. Stop signals were presented on 30% of the trials, instructing subjects to withhold their response. Response force on non-signal (go) trials and the duration of response inhibition on signal (stop) trials increased as response probability decreased. This pattern of findings was interpreted to support the hypothesis predicting that stopping is more difficult when response readiness is low than when it is high.     

Don’t look! don’t touch! inhibitory control of eye and hand movements

Inhibitory control of eye and hand movements was compared in the stop-signal task. Subjects moved their eyes to the right or left or pressed keys on the right or left in response to visual stimuli. The stimuli were either central (angle brackets pointing left or right) or peripheral (plus signs turning into Xs left or right of fixation), and the task was either pro (respond on the same side as the stimulus) or anti (respond on the opposite side). Occasionally, a stop signal was presented, which instructed subjects to inhibit their responses to the go stimulus. Stop-signal reaction times (SSRTs) were faster overall for eye movements than for hand movements, and they were affected differently by stimulus conditions (central vs. peripheral) and task (pro vs. anti), suggesting that the eyes and hands are inhibited by different processes operating under similar principles (i.e., a race between stop and go processes).     

Post-stop-signal adjustments: inhibition improves subsequent inhibition

Performance in the stop-signal paradigm involves a balance between going and stopping, and one way that this balance is struck is through shifting priority away from the go task, slowing responses after a stop signal, and improving the probability of inhibition. In 6 experiments, the authors tested whether there is a corresponding shift in priority toward the stop task, speeding reaction time to the stop signal. Consistent with this hypothesis, stop-signal reaction time (SSRT) decreased on the trial immediately following a stop signal in each experiment. Experiments 2-4 used 2 very different stop signals within a modality, and stopping improved when the stop stimulus repeated and alternated. Experiments 5 and 6 presented stop signals in different modalities and showed that SSRT improved only when the stop stimulus repeated within a modality. These results demonstrate within-modality post-stop-signal speeding of response inhibition.