Working Memory Load Selectively Influences Response Inhibition in a Stop Signal Task.

Inhibitory control is a key executive function and has been studied extensively using the stop signal task. By applying a simple race model that posits an independent race between a GO process responsible for initiation of responses and a STOP process responsible for inhibition of responses, one can estimate how long it takes an individual to inhibit an ongoing response, the stop signal reaction time. Here, we examined how stop signal reaction time can be affected by working memory. Participants engaged in a dual task; they completed a stop signal task under low and high working memory load conditions. Working memory capacity was also measured. We found that the STOP process was lengthened in the high, compared to the low, working memory load condition, as evidenced by differences in stop signal reaction time. The GO process was unaffected and working memory capacity could not account for differences across the load conditions. These results indicate that inhibitory control can be influenced by placing demands on working memory.

The Effects of Videogaming with a Brain-Computer Interface on Mood and Physiological Arousal.

Objective: In recent years, immersive videogame technologies such as virtual reality have been shown to affect psychological welfare in such way that they can be applied to clinical psychology treatments. However, the effects of videogaming with other immersive gaming apparatuses such as commercial electroencephalography (EEG)-based brain-computer interfaces (BCIs) on psychological welfare have not been extensively researched. Thus, we aimed at providing early insights into some of these effects by looking at how videogaming with a commercial EEG-based BCI would impact mood and physiological arousal. Materials and Methods: A total of 26 participants were sampled. Participants were randomly assigned to either a BCI condition or a traditional condition wherein they played an action videogame with a commercial EEG-based BCI or a standard keyboard and mouse interface for 20 minutes. In both conditions, participants filled out the profile of mood states to assess mood and the perceived stress scale to control for stress. We also measured heart rate, heart rate variability as measured by the root mean square of successive differences, and galvanic skin response (GSR) amplitude differences. Results: Participants in the BCI condition overall reported a significantly higher total mood disturbance (P < 0.05), tension (P < 0.05), confusion (P < 0.05), and significantly less vigor (P < 0.05). We also found that participants in the BCI condition had significantly lower GSR amplitude differences between gaming and baseline (P < 0.05). Conclusion: The results suggest that the use of commercial EEG-based BCIs for playing with videogames can induce greater frustration and negative moods than playing with a traditional keyboard and mouse interface, possibly limiting their use in clinical psychology settings.

Visual-spatial attention aids the maintenance of object representations in visual working memory.

Theories have proposed that the maintenance of object representations in visual working memory is aided by a spatial rehearsal mechanism. In this study, we used two different approaches to test the hypothesis that overt and covert visual-spatial attention mechanisms contribute to the maintenance of object representations in visual working memory. First, we tracked observers' eye movements while they remembered a variable number of objects during change-detection tasks. We observed that during the blank retention interval, participants spontaneously shifted gaze to the locations that the objects had occupied in the memory array. Next, we hypothesized that if attention mechanisms contribute to the maintenance of object representations, then drawing attention away from the object locations during the retention interval should impair object memory during these change-detection tasks. Supporting this prediction, we found that attending to the fixation point in anticipation of a brief probe stimulus during the retention interval reduced change-detection accuracy, even on the trials in which no probe occurred. These findings support models of working memory in which visual-spatial selection mechanisms contribute to the maintenance of object representations.

Neural basis of adaptive response time adjustment during saccade countermanding.

Humans and macaque monkeys adjust their response time adaptively in stop-signal (countermanding) tasks, responding slower after stop-signal trials than after control trials with no stop signal. We investigated the neural mechanism underlying this adaptive response time adjustment in macaque monkeys performing a saccade countermanding task. Earlier research showed that movements are initiated when the random accumulation of presaccadic movement-related activity reaches a fixed threshold. We found that a systematic delay in response time after stop-signal trials was accomplished not through a change of threshold, baseline, or accumulation rate, but instead through a change in the time when activity first began to accumulate. The neurons underlying movement initiation have been identified with stochastic accumulator models of response time performance. Therefore, this new result provides surprising new insights into the neural instantiation of stochastic accumulator models and the mechanisms through which executive control can be exerted.

Response inhibition and response monitoring in a saccadic countermanding task in schizophrenia.

BACKGROUND: Cognitive control deficits are pervasive in individuals with schizophrenia (SZ) and are reliable predictors of functional outcome, but the specificity of these deficits and their underlying neural mechanisms have not been fully elucidated. The objective of the present study was to determine the nature of response inhibition and response monitoring deficits in SZ and their relationship to symptoms and social and occupational functioning with a behavioral paradigm that provides a translational approach to investigating cognitive control. METHODS: Seventeen patients with SZ and 16 demographically matched healthy control subjects participated in a saccadic countermanding task. Performance on this task is approximated as a race between movement generation and inhibition processes; this race model provides an estimate of the time needed to cancel a planned movement. Response monitoring can be assessed by reaction time adjustments on the basis of trial history. RESULTS: Saccadic reaction time was normal, but patients required more time to inhibit a planned saccade. The latency of the inhibitory process was associated with the severity of negative symptoms and poorer occupational functioning. Both groups slowed down significantly after correctly cancelled and erroneously noncancelled stop signal trials, but patients slowed down more than control subjects after correctly inhibited saccades. CONCLUSIONS: These results suggest that SZ is associated with a difficulty in inhibiting planned movements and an inflated response adjustment effect after inhibiting a saccade. Furthermore, behavioral results are consistent with potential abnormalities in frontal and supplementary eye fields in patients with SZ.

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.

Proactive inhibitory control and attractor dynamics in countermanding action: a spiking neural circuit model.

Flexible behavior depends on the brain's ability to suppress a habitual response or to cancel a planned movement whenever needed. Such inhibitory control has been studied using the countermanding paradigm in which subjects are required to withhold an imminent movement when a stop signal appears infrequently in a fraction of trials. To elucidate the circuit mechanism of inhibitory control of action, we developed a recurrent network model consisting of spiking movement (GO) neurons and fixation (STOP) neurons, based on neurophysiological observations in the frontal eye field and superior colliculus of behaving monkeys. The model places a premium on the network dynamics before the onset of a stop signal, especially the experimentally observed high baseline activity of fixation neurons, which is assumed to be modulated by a persistent top-down control signal, and their synaptic interaction with movement neurons. The model simulated observed neural activity and fit behavioral performance quantitatively. In contrast to a race model in which the STOP process is initiated at the onset of a stop signal, in our model whether a movement will eventually be canceled is determined largely by the proactive top-down control and the stochastic network dynamics, even before the appearance of the stop signal. A prediction about the correlation between the fixation neural activity and the behavioral outcome was verified in the neurophysiological data recorded from behaving monkeys. The proposed mechanism for adjusting control through tonically active neurons that inhibit movement-producing neurons has significant implications for exploring the basis of impulsivity associated with psychiatric disorders.

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.

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.

Influence of history on saccade countermanding performance in humans and macaque monkeys.

The stop-signal or countermanding task probes the ability to control action by requiring subjects to withhold a planned movement in response to an infrequent stop signal which they do with variable success depending on the delay of the stop signal. We investigated whether performance of humans and macaque monkeys in a saccade countermanding task was influenced by stimulus and performance history. In spite of idiosyncrasies across subjects several trends were evident in both humans and monkeys. Response time decreased after successive trials with no stop signal. Response time increased after successive trials with a stop signal. However, post-error slowing was not observed. Increased response time was observed mainly or only after cancelled (signal inhibit) trials and not after noncancelled (signal respond) trials. These global trends were based on rapid adjustments of response time in response to momentary fluctuations in the fraction of stop signal trials. The effects of trial sequence on the probability of responding were weaker and more idiosyncratic across subjects when stop signal fraction was fixed. However, both response time and probability of responding were influenced strongly by variations in the fraction of stop signal trials. These results indicate that the race model of countermanding performance requires extension to account for these sequential dependencies and provide a basis for physiological studies of executive control of countermanding saccade performance.

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.

Ocular tracking as a measure of auditory motion perception.

Motion is a potent sub-modality of vision. Motion cues alone can be used to segment images into figure and ground and break camouflage. Specific patterns of motion support vivid percepts of form, guide locomotion by specifying directional heading and the passage of objects, and in case of an impending collision, the time to impact. Visual motion also drives smooth pursuit eye movements (SPEMs) that serve to stabilize the retinal image of objects in motion. In contrast, the auditory system does not appear to be particularly sensitive to motion. We review the ambiguous status of auditory motion processing from the psychophysical and electrophysiological perspectives. We then report the results of two experiments that use ocular tracking performance as an objective measure of the perception of auditory motion in humans. We examine ocular tracking of auditory motion, visual motion, combined auditory + visual motion and imagined motion in both the frontal plane and in depth. The results demonstrate that ocular tracking of auditory motion is no better than ocular tracking of imagined motion. These results are consistent with the suggestion that, unlike the visual system, the human auditory system is not endowed with low-level motion sensitive elements. We hypothesize however, that auditory information may gain access to a recently described high-level motion processing system that is heavily dependent on 'top-down' influences, including attention.

Smooth pursuit under stimulus-response uncertainty.

Simple reaction times (RTs) are typically faster than choice reaction times and increase with uncertainty according to Hick's law. Here we show that smooth pursuit eye movement RTs show no effect of SR uncertainty while joystick tracking shows a step change between SRT and CRT, but no significant increases beyond two choices. The results suggest there is a benefit to pre-programming joystick tracking but not for smooth pursuit eye movements (SPEMs).

Saccades operate in violation of Hick's law.

Hick's law states that response times (RTs) increase in proportion to the logarithm of the number of potential stimulus-response (S-R) alternatives. We hypothesized that time-consuming processes associated with response selection contribute significantly to this effect. We also hypothesized that the latency of saccades might not conform to Hick's law since visually guided saccades can be automatically selected using topographically organized pathways that convert spatially coded visual activity into spatially coded motor commands. We evaluated these hypotheses by examining three response modalities for their compliance with Hick's law: saccades directed to a visual target (prosaccades), saccades directed away from the target (antisaccades) and manual responses in which each digit was associated with a specific target location (key-press responses). Both antisaccades and key-press responses conformed to Hick's law but saccade latencies were completely unaffected by S-R uncertainty. The significance of these findings is considered in terms of the processes of response selection and premotor programming.