Edward K. Vogel

The capacity of visual working memory for features and conjunctions

Short-term memory storage can be divided into separate subsystems for verbal information and visual information, and recent studies have begun to delineate the neural substrates of these working-memory systems. Although the verbal storage system has been well characterized, the storage capacity of visual working memory has not yet been established for simple, suprathreshold features or for conjunctions of features. Here we demonstrate that it is possible to retain information about only four colours or orientations in visual working memory at one time. However, it is also possible to retain both the colour and the orientation of four objects, indicating that visual working memory stores integrated objects rather than individual features. Indeed, objects defined by a conjunction of four features can be retained in working memory just as well as single-feature objects, allowing sixteen individual features to be retained when distributed across four objects. Thus, the capacity of visual working memory must be understood in terms of integrated objects rather than individual features, which places significant constraints on cognitive and neurobiological models of the temporary storage of visual information.

Neural activity predicts individual differences in visual working memory capacity

Contrary to our rich phenomenological visual experience, our visual short-term memory system can maintain representations of only three to four objects at any given moment. For over a century, the capacity of visual memory has been shown to vary substantially across individuals, ranging from 1.5 to about 5 objects. Although numerous studies have recently begun to characterize the neural substrates of visual memory processes, a neurophysiological index of storage capacity limitations has not yet been established. Here, we provide electrophysiological evidence for lateralized activity in humans that reflects the encoding and maintenance of items in visual memory. The amplitude of this activity is strongly modulated by the number of objects being held in the memory at the time, but approaches a limit asymptotically for arrays that meet or exceed storage capacity. Indeed, the precise limit is determined by each individuals memory capacity, such that the activity from low-capacity individuals reaches this plateau much sooner than that from high-capacity individuals. Consequently, this measure provides a strong neurophysiological predictor of an individuals capacity, allowing the demonstration of a direct relationship between neural activity and memory capacity.

Neural measures reveal individual differences in controlling access to working memory

The capacity of visual short-term memory is highly limited, maintaining only three to four objects simultaneously. This extreme limitation necessitates efficient mechanisms to select only the most relevant objects from the immediate environment to be represented in memory and to restrict irrelevant items from consuming capacity. Here we report a neurophysiological measure of this memory selection mechanism in humans that gauges an individuals efficiency at excluding irrelevant items from being stored in memory. By examining the moment-by-moment contents of visual memory, we observe that selection efficiency varies substantially across individuals and is strongly predicted by the particular memory capacity of each person. Specifically, high capacity individuals are much more efficient at representing only the relevant items than are low capacity individuals, who inefficiently encode and maintain information about the irrelevant items present in the display. These results provide evidence that under many circumstances low capacity individuals may actually store more information in memory than high capacity individuals. Indeed, this ancillary allocation of memory capacity to irrelevant objects may be a primary source of putative differences in overall storage capacity.

Event-related potential studies of attention

Over the past 30 years, recordings of event-related potentials (ERPs) from normal individuals have played an increasingly important role in our understanding of the mechanisms of attention. This article reviews some of the recent ERP studies of attention, focusing on studies that isolate the operation of attention in specific cognitive subsystems such as perception, working memory, and response selection. Several conclusions are drawn. First, under some conditions attention modulates the initial feedforward volley of neural activity in intermediate visual processing areas. Second, these early effects can be observed for both the voluntary allocation of attention and for the automatic capture of attention following a peripheral visual transient. Third, these effects are present not only when attention is directed to a location in 2-dimensional space, but also when attention is directed to one of two spatially overlapping surfaces. Fourth, attention does not modulate sensory activity unless sensory systems are overloaded; when sensory systems are not taxed, attention may instead operate to influence memory or response processes. That is, attention operates to mitigate information overload in whichever cognitive subsystems are overloaded by a particular combination of stimuli and task.

The visual N1 component as an index of a discrimination process.

Many previous studies have demonstrated that the visual N1 component is larger for attended-location stimuli than for unattended-location stimuli. This difference is observed typically only for tasks involving a discrimination of the attended-location stimuli, suggesting that the N1 wave reflects a discrimination process that is applied to the attended location. The present study tested this hypothesis by examining the N1 component elicited by attended stimuli under conditions that either required or did not require the subject to perform a discrimination. Specifically, the N1 elicited by foveal stimuli during choice-reaction time (RT) tasks was compared with the N1 elicited by identical stimuli during simple-RT tasks. In three experiments, a larger posterior N1 was observed in choice-RT tasks than in simple-RT tasks, even when several potential confounds were eliminated (e.g., arousal and motor preparation). This N1 discrimination effect was observed even when no motor response was required and was present for both color- and form-based discriminations. Moreover, this discrimination effect was equally large for easy and difficult discriminations, arguing against a simple resource-based explanation of the present results. Instead, the results of this study are consistent with the hypothesis that the visual N1 component reflects the operation of a discrimination process within the focus of attention.

Interactions between attention and working memory

Studies of attention and working memory address the fundamental limits in our ability to encode and maintain behaviorally relevant information, processes that are critical for goal-driven processing. Here we review our current understanding of the interactions between these processes, with a focus on how each construct encompasses a variety of dissociable phenomena. Attention facilitates target processing during both perceptual and postperceptual stages of processing, and functionally dissociated processes have been implicated in the maintenance of different kinds of information in working memory. Thus, although it is clear that these processes are closely intertwined, the nature of these interactions depends upon the specific variety of attention or working memory that is considered.

Visual Working Memory Represents a Fixed Number of Items Regardless of Complexity

Does visual working memory represent a fixed number of objects, or is capacity reduced as object complexity increases? We measured accuracy in detecting changes between sample and test displays and found that capacity estimates dropped as complexity increased. However, these apparent capacity reductions were strongly correlated with increases in sample-test similarity (r = .97), raising the possibility that change detection was limited by errors in comparing the sample and test, rather than by the number of items that were maintained in working memory. Accordingly, when sample-test similarity was low, capacity estimates for even the most complex objects were equivalent to the estimate for the simplest objects (r = .88), suggesting that visual working memory represents a fixed number of items regardless of complexity. Finally, a correlational analysis suggested a two-factor model of working memory ability, in which the number and resolution of representations in working memory correspond to distinct dimensions of memory ability.

Stimulus-Specific Delay Activity in Human Primary Visual Cortex

Working memory (WM) involves maintaining information in an on-line state. One emerging view is that information in WM is maintained via sensory recruitment, such that information is stored via sustained activity in the sensory areas that encode the to-be-remembered information. Using functional magnetic resonance imaging, we observed that key sensory regions such as primary visual cortex (V1) showed little evidence of sustained increases in mean activation during a WM delay period, though such amplitude increases have typically been used to determine whether a region is involved in on-line maintenance. However, a multivoxel pattern analysis of delay-period activity revealed a sustained pattern of activation in V1 that represented only the intentionally stored feature of a multifeature object. Moreover, the pattern of delay activity was qualitatively similar to that observed during the discrimination of sensory stimuli, suggesting that WM representations in V1 are reasonable “copies” of those evoked during pure sensory processing.

The time course of consolidation in visual working memory.

How long does it take to form a durable representation in visual working memory? Several theorists have proposed that this consolidation process is very slow. Here, we measured the time course of consolidation. Observers performed a change-detection task for colored squares, and shortly after the presentation of the first array, pattern masks were presented at the locations of each of the colored squares to disrupt representations that had not yet been consolidated. Performance on the memory task was impaired when the delay between the colored squares and the masks was short, and this effect became larger when the number of colored squares was increased. The rate of consolidation was approximately 50 ms per item, which is considerably faster than previous proposals.

Visual working memory capacity: from psychophysics and neurobiology to individual differences.

Visual working memory capacity is of great interest because it is strongly correlated with overall cognitive ability, can be understood at the level of neural circuits, and is easily measured. Recent studies have shown that capacity influences tasks ranging from saccade targeting to analogical reasoning. A debate has arisen over whether capacity is constrained by a limited number of discrete representations or by an infinitely divisible resource, but the empirical evidence and neural network models currently favor a discrete item limit. Capacity differs markedly across individuals and groups, and recent research indicates that some of these differences reflect true differences in storage capacity whereas others reflect variations in the ability to use memory capacity efficiently.