Eric H. Schumacher

Verbal working memory load affects regional brain activation as measured by pet

We report an experiment that assesses the effect of variations in memory load on brain activations that mediate verbal working memory. The paradigm that forms the basis of this experiment is the n-back task in which subjects must decide for each letter in a series whether it matches the one presented n items back in the series. This task is of interest because it recruits processes involved in both the storage and manipulation of information in working memory. Variations in task difficulty were accomplished by varying the value of n. As n increased, subjects showed poorer behavioral performance as well as monotonically increasing magnitudes of brain activation in a large number of sites that together have been identified with verbal working-memory processes. By contrast, there was no reliable increase in activation in sites that are unrelated to working memory. These results validate the use of parametric manipulation of task variables in neuroimaging research, and they converge with the subtraction paradigm used most often in neuroimaging. In addition, the data support a model of working memory that includes both storage and executive processes that recruit a network of brain areas, all of which are involved in task performance.

The Role of Parietal Cortex in Verbal Working Memory

Neuroimaging studies of normal subjects and studies of patients with focal lesions implicate regions of parietal cortex in verbal working memory (VWM), yet the precise role of parietal cortex in VWM remains unclear. Some evidence (Paulesu et al., 1993; Awh et al., 1996) suggests that the parietal cortex mediates the storage of verbal information, but these studies and most previous ones included encoding and retrieval processes as well as storage and rehearsal of verbal information. A recent positron emission tomography (PET) study by Fiez et al. (1996) isolated storage and rehearsal from other VWM processes and did not find reliable activation in parietal cortex. This result suggests that parietal cortex may not be involved in VWM storage, contrary to previous proposals. However, we report two behavioral studies indicating that some of the verbal material used by Fiez et al. (1996) may not have required phonological representations in VWM. In addition, we report a PET study that isolated VWM encoding, retrieval, and storage and rehearsal processes in different PET scans and used material likely to require phonological codes in VWM. After subtraction of appropriate controls, the encoding condition revealed no reliable activations; the retrieval condition revealed reliable activations in dorsolateral prefrontal, anterior cingulate, posterior parietal, and extrastriate cortices, and the storage condition revealed reliable activations in dorsolateral prefrontal, inferior frontal, premotor, and posterior parietal cortices, as well as cerebellum. These results suggest that parietal regions are part of a network of brain areas that mediate the short-term storage and retrieval of phonologically coded verbal material.

Spatial versus object working memory: Pet investigations

We used positron emission tomography (PET) to answer the following question: Is working memory a unitary storage system, or does it instead include different storage buffers for different kinds of information? In Experiment 1, PET measures were taken while subjects engaged in either a spatial-memory task (retain the position of three dots for 3 sec) or an object-memory task (retain the identity of two objects for 3 sec). The results manifested a striking double dissociation, as the spatial task activated only right-hemisphere regions, whereas the object task activated primarily left-hemisphere regions. The spatial (right-hemisphere) regions included occipital, parietal, and prefrontal areas, while the object (left-hemisphere) regions included inferotemporal and parietal areas. Experiment 2 was similar to Experiment 1 except that the stimuli and trial events were identical for the spatial and object tasks; whether spatial or object memory was required was manipulated by instructions. The PET results once more showed a double dissociation, as the spatial task activated primarily right-hemisphere regions (again including occipital, parietal and prefrontal areas), whereas the object task activated only left-hemisphere regions (again including inferotemporal and parietal areas). Experiment 3 was a strictly behavioral study, which produced another double dissociation. It used the same tasks as Experiment 2, and showed that a variation in spatial similarity affected performance in the spatial but not the object task, whereas a variation in shape similarity affected performance in the object but not the spatial task. Taken together, the results of the three experiments clearly imply that different working-memory buffers are used for storing spatial and object information.

Virtually Perfect Time Sharing in Dual-Task Performance: Uncorking the Central Cognitive Bottleneck

A fundamental issue for psychological science concerns the extent to which people can simultaneously perform two perceptual-motor tasks. Some theorists have hypothesized that such dual-task performance is severely and persistently constrained by a central cognitive “bottleneck,” whereas others have hypothesized that skilled procedural decision making and response selection for two or more tasks can proceed at the same time under adaptive executive control. The three experiments reported here support this latter hypothesis. Their results show that after relatively modest amounts of practice, at least some participants achieve virtually perfect time sharing in the dual-task performance of basic choice reaction tasks. The results also show that observed interference between tasks can be modulated by instructions about differential task priorities and personal preferences for daring (concurrent) or cautious (successive) scheduling of tasks. Given this outcome, future research should investigate exactly when and how such sophisticated skills in dual-task performance are acquired.

PET Evidence for an Amodal Verbal Working Memory System

Current models of verbal working memory assume that modality-specific representations are translated into phonological representations before entering the working memory system. We report an experiment that tests this assumption. Positron emission tomography measures were taken while subjects performed a verbal working memory task. Stimuli were presented either visually or aurally, and a visual or auditory search tasks, respectively, was used as a control. Results revealed an almost complete overlap between the active memory areas regardless of input modality. These areas included dorsolateral frontal, Brocas area, SMA, and premotor cortex in the left hemisphere; bilateral superior and posterior parietal cortices and anterior cingulate; and right cerebellum. These results correspond well with previous research and suggest that verbal working memory is modality independent and is mediated by a circuit involving frontal, parietal, and cerebellar mechanisms.

A functional MRI study of the influence of practice on component processes of working memory

Previous neuroimaging studies have shown that neural activity changes with task practice. The types of changes reported have been inconsistent, however, and the neural mechanisms involved remain unclear. In this study, we investigated the influence of practice on different component processes of working memory (WM) using a face WM task. Event-related functional magnetic resonance imaging (fMRI) methodology allowed us to examine signal changes from early to late in the scanning session within different task stages (i.e., encoding, delay, retrieval), as well as to determine the influence of different levels of WM load on neural activity. We found practice-related decreases in fMRI signal and effects of memory load occurring primarily during encoding. This suggests that practice improves encoding efficiency, especially at higher memory loads. The decreases in fMRI signal we observed were not accompanied by improved behavioral performance; in fact, error rate increased for high WM load trials, indicating that practice-related changes in activation may occur during a scanning session without behavioral evidence of learning. Our results suggest that practice influences particular component processes of WM differently, and that the efficiency of these processes may not be captured by performance measures alone.

Neural Evidence for Representation-Specific Response Selection

Response selection is the mental process of choosing representations for appropriate motor behaviors given particular environmental stimuli and ones current task situation and goals. Many cognitive theories of response selection postulate a unitary process. That is, one central response-selection mechanism chooses appropriate responses in most, if not all, task situations. However, neuroscience research shows that neural processing is often localized based on the type of information processed. Our current experiments investigate whether response selection is unitary or stimulus specific by manipulating response-selection difficulty in two functional magnetic resonance imaging experiments using spatial and nonspatial stimuli. The same participants were used in both experiments. We found spatial response selection involves the right prefrontal cortex, the bilateral premotor cortex, and the dorsal parietal cortical regions (precuneus and superior parietal lobule). Nonspatial response selection, conversely, involves the left prefrontal cortex and the more ventral posterior cortical regions (left middle temporal gyrus, left inferior parietal lobule, and right extrastriate cortex). Our brain activation data suggest a cognitive model for response selection in which different brain networks mediate the choice of appropriate responses for different types of stimuli. This model is consistent with behavioral research suggesting that responseselection processing may be more flexible and adaptive than originally proposed.

Adaptive executive control: Flexible multiple-task performance without pervasive immutable response-selection bottlenecks

Abstract A new theoretical framework, the EPIC (Executive-Process/Interactive-Control) architecture, provides the basis for accurate detailed computational models of human multiple-task performance. Contrary to the traditional response-selection bottleneck hypothesis, EPICs cognitive processor can select responses and do other procedural operations simultaneously for multiple concurrent tasks. Using this capacity together with flexible executive control of peripheral perceptual-motor components, EPIC computational models account well for various patterns of mean reaction times, systematic individual differences in multiple-task performance, and influences of special training on peoples task-coordination strategies. These diverse phenomena, and EPICs success at modeling them, raise strong doubts about the existence of a pervasive immutable response-selection bottleneck in the human information-processing system. The present research therefore helps further characterize the nature of discrete versus continuous information processing.

Neural implementation of response selection in humans as revealed by localized effects of stimulus–response compatibility on brain activation

Response selection, which involves choosing representations for appropriate motor behaviors given ones current situation, is a fundamental mental process central to a wide variety of human performance, yet the neural mechanisms underlying this mental process remain unclear. Research using nonhuman primates implicates ventral prefrontal and lateral premotor cortices in this process. In contrast, human neuroimaging research also highlights the role of dorsal prefrontal, anterior cingulate, and superior parietal cortices in response selection. This inconsistency may stem from the difficulty of isolating response selection within the constraints of cognitive subtraction methodology utilized in neuroimaging. We overcome this limitation by using an experimental procedure designed to selectively influence discrete mental processing stages and analyses that are less reliant on the assumptions of cognitive subtraction. We varied stimulus contrast to affect stimulus encoding and stimulus–response compatibility to affect response selection. Brain activation data suggest processing specific to response selection in superior parietal and dorsal prefrontal cortices, and not ventral prefrontal cortex. Anterior cingulate and lateral premotor cortices may also be involved in response selection, or these regions may mediate other response processes. Hum. Brain Mapping 17:193–201, 2002.

Audience preferences are predicted by temporal reliability of neural processing

Naturalistic stimuli evoke highly reliable brain activity across viewers. Here we record neural activity from a group of naive individuals while viewing popular, previously-broadcast television content for which the broad audience response is characterized by social media activity and audience ratings. We find that the level of inter-subject correlation in the evoked encephalographic responses predicts the expressions of interest and preference among thousands. Surprisingly, ratings of the larger audience are predicted with greater accuracy than those of the individuals from whom the neural data is obtained. An additional functional magnetic resonance imaging study employing a separate sample of subjects shows that the level of neural reliability evoked by these stimuli covaries with the amount of blood-oxygenation-level-dependent (BOLD) activation in higher-order visual and auditory regions. Our findings suggest that stimuli which we judge favourably may be those to which our brains respond in a stereotypical manner shared by our peers.