Richard P. Heitz

An automated version of the operation span task

We present an easy-to-administer and automated version of a popular working memory (WM) capacity task (operation span; Ospan) that is mouse driven, scores itself, and requires little intervention on the part of the experimenter. It is shown that this version of Ospan correlates well with other measures of WM capacity and has both good internal consistency (alpha=.78) and test-retest reliability (.83). In addition, the automated version of Ospan (Aospan) was shown to load on the same factor as two other WM measures. This WM capacity factor correlated with a factor composed of fluid abilities measures. The utility of the Aospan was further demonstrated by analyzing response times (RTs) that indicated that RT measures obtained in the task accounted for additional variance in predicting fluid abilities. Our results suggest that Aospan is a reliable and valid indicator of WM capacity that can be applied to a wide array of research domains.

Complex working memory span tasks and higher-order cognition: A latent-variable analysis of the relationship between processing and storage

Complex span tasks, assumed by many to measure an individuals working memory capacity, are predictive of several aspects of higher-order cognition. However, the underlying cause of the relationships between “processing-and-storage” tasks and cognitive abilities is still hotly debated nearly 30 years after the tasks were first introduced. The current study utilised latent constructs across verbal, numerical, and spatial content domains to examine a number of questions regarding the predictive power of complex span tasks. In particular, the relations among processing time, processing accuracy, and storage accuracy from the complex span tasks were examined, in combination with their respective relationships with fluid intelligence. The results point to a complicated pattern of unique and shared variance among the constructs. Implications for various theories of working memory are discussed.

Neurally constrained modeling of perceptual decision making.

Stochastic accumulator models account for response time in perceptual decision-making tasks by assuming that perceptual evidence accumulates to a threshold. The present investigation mapped the firing rate of frontal eye field (FEF) visual neurons onto perceptual evidence and the firing rate of FEF movement neurons onto evidence accumulation to test alternative models of how evidence is combined in the accumulation process. The models were evaluated on their ability to predict both response time distributions and movement neuron activity observed in monkeys performing a visual search task. Models that assume gating of perceptual evidence to the accumulating units provide the best account of both behavioral and neural data. These results identify discrete stages of processing with anatomically distinct neural populations and rule out several alternative architectures. The results also illustrate the use of neurophysiological data as a model selection tool and establish a novel framework to bridge computational and neural levels of explanation.

Focusing the spotlight: individual differences in visual attention control.

A time-course analysis of visual attention focusing (attentional constraint) was conducted in groups of participants with high and low working memory spans, a dimension the authors have argued reflects the ability to control attention. In 4 experiments, participants performed the Eriksen flanker paradigm under increasing levels of speed stress. Conditional accuracy functions were derived to measure the time course of attentional constraint. The data showed that accuracy rates rose toward asymptote at different rates, with participants with high working memory spans reaching peak performance before participants with low working memory spans. The authors interpret these data in terms of a rate of attention constraint model.

The speed-accuracy tradeoff: history, physiology, methodology, and behavior

There are few behavioral effects as ubiquitous as the speed-accuracy tradeoff (SAT). From insects to rodents to primates, the tendency for decision speed to covary with decision accuracy seems an inescapable property of choice behavior. Recently, the SAT has received renewed interest, as neuroscience approaches begin to uncover its neural underpinnings and computational models are compelled to incorporate it as a necessary benchmark. The present work provides a comprehensive overview of SAT. First, I trace its history as a tractable behavioral phenomenon and the role it has played in shaping mathematical descriptions of the decision process. Second, I present a “users guide” of SAT methodology, including a critical review of common experimental manipulations and analysis techniques and a treatment of the typical behavioral patterns that emerge when SAT is manipulated directly. Finally, I review applications of this methodology in several domains.

Neural mechanisms of saccade target selection: gated accumulator model of the visual–motor cascade

We review a new computational model developed to understand how evidence about stimulus salience in visual search is translated into a saccade command. The model uses the activity of visually responsive neurons in the frontal eye field as evidence for stimulus salience that is accumulated in a network of stochastic accumulators to produce accurate and timely saccades. We discovered that only when the input to the accumulation process was gated could the model account for the variability in search performance and predict the dynamics of movement neuron discharge rates. This union of cognitive modeling and neurophysiology indicates how the visual–motor transformation can occur, and provides a concrete mapping between neuron function and specific cognitive processes.

Neural Basis of the Set-Size Effect in Frontal Eye Field: Timing of Attention During Visual Search

Visual search for a target object among distractors often takes longer when more distractors are present. To understand the neural basis of this capacity limitation, we recorded activity from visually responsive neurons in the frontal eye field (FEF) of macaque monkeys searching for a target among distractors defined by form (randomly oriented T or L). To test the hypothesis that the delay of response time with increasing number of distractors originates in the delay of attention allocation by FEF neurons, we manipulated the number of distractors presented with the search target. When monkeys were presented with more distractors, visual target selection was delayed and neuronal activity was reduced in proportion to longer response time. These findings indicate that the time taken by FEF neurons to select the target contributes to the variation in visual search efficiency.

Homologous mechanisms of visuospatial working memory maintenance in macaque and human: Properties and sources

Although areas of frontal cortex are thought to be critical for maintaining information in visuospatial working memory, the event-related potential (ERP) index of maintenance is found over posterior cortex in humans. In the present study, we reconcile these seemingly contradictory findings. Here, we show that macaque monkeys and humans exhibit the same posterior ERP signature of working memory maintenance that predicts the precision of the memory-based behavioral responses. In addition, we show that the specific pattern of rhythmic oscillations in the alpha band, recently demonstrated to underlie the human visual working memory ERP component, is also present in monkeys. Next, we concurrently recorded intracranial local field potentials from two prefrontal and another frontal cortical area to determine their contribution to the surface potential indexing maintenance. The local fields in the two prefrontal areas, but not the cortex immediately posterior, exhibited amplitude modulations, timing, and relationships to behavior indicating that they contribute to the generation of the surface ERP component measured from the distal posterior electrodes. Rhythmic neural activity in the theta and gamma bands during maintenance provided converging support for the engagement of the same brain regions. These findings demonstrate that nonhuman primates have homologous electrophysiological signatures of visuospatial working memory to those of humans and that a distributed neural network, including frontal areas, underlies the posterior ERP index of visuospatial working memory maintenance.

Biophysical Support for Functionally Distinct Cell Types in the Frontal Eye Field

Numerous studies have described different functional cell types in the frontal eye field (FEF), but the reliability of the distinction between these types has been uncertain. Studies in other brain areas have described specific differences in the width of action potentials recorded from different cell types. To substantiate the functionally defined cell types encountered in FEF, we measured the width of spikes of visual, movement, and visuomovement types of FEF neurons in macaque monkeys. We show that visuomovement neurons had the thinnest spikes, consistent with a role in local processing. Movement neurons had the widest spikes, consistent with their role in sending eye movement commands to subcortical structures such as the superior colliculus. Visual neurons had wider spikes than visuomovement neurons, consistent with their role in receiving projections from occipital and parietal cortex. These results show how structure and function of FEF can be linked to guide inferences about neuronal architecture.

A touch screen based Stop Signal Response Task in rhesus monkeys for studying impulsivity associated with chronic cocaine self-administration

Among a range of cognitive deficits, human cocaine addicts display increased impulsivity and decreased performance monitoring. In order to establish an animal model that can be used to study the underlying neurobiology of these deficits associated with addiction, we have developed a touch screen based Stop Signal Response Task for rhesus monkeys. This task is essentially identical to the clinically used Stop Signal Task employed for diagnostic and research purposes. In this task, impulsivity is reflected in the amount of time needed to inhibit a response after it has been initiated, the Stop Signal Response Time (SSRT). Performance monitoring is reflected by the slowing of response times following Stop trials (Post-Stop Slowing, PSS). Herein we report on the task structure, the staged methods for training animals to perform the task, and a comparison of performance values for control and cocaine experienced animals. Relative to controls, monkeys that had self-administered cocaine, followed by 18 months abstinence, displayed increased impulsivity (increased SSRT values), and decreased performance monitoring (decreased PSS values). Our results are consistent with human data, and thereby establish an ideal animal model for studying the etiology and underlying neurobiology of cocaine-induced impulse control and performance monitoring deficits.