Posterior parietal cortex activity predicts individual differences in visual short-term memory capacity

Humans show a severe capacity limit in the number of objects they can store in visual short-term memory (VSTM). We recently demonstrated with functional magnetic resonance imaging that VSTM storage capacity estimated in averaged group data correlated strongly with posterior parietal/superior occipital cortex activity (Todd & Marois, 2004). However, individuals varied widely in their VSTM capacity. Here, we examined the neural basis of these individual differences. A voxelwise, individualdifferences analysis revealed a significant correlation between posterior parietal cortex (PPC) activity and individuals’ VSTM storage capacity. In addition, a region-of-interest analysis indicated that other brain regions, particularly visual occipital cortex, may contribute to individual differences in VSTM capacity. Thus, although not ruling out contributions from other brain regions, the individual-differences approach supports a key role for the PPC in VSTM by demonstrating that its activity level predicts individual differences in VSTM storage capacity.     

Visual short-term memory for sequential arrays

The capacity of visual short-term memory (VSTM) for a single visual display has been investigated in past research, but VSTM for multiple sequential arrays has been explored only recently. In this study, we investigate the capacity of VSTM across two sequential arrays separated by a variable stimulus onset asynchrony (SOA). VSTM for spatial locations (Experiment 1), colors (Experiments 2-4), orientations (Experiments 3 and 4), and conjunction of color and orientation (Experiment 4) were tested, with the SOA across the two sequential arrays varying from 100 to 1,500 msec. We find that VSTM for the trailing array is much better than VSTM for the leading array, but when averaged across the two arrays VSTM has a constant capacity independent of the SOA. We suggest that multiple displays compete for retention in VSTM and that separating information into two temporally discrete groups does not enhance the overall capacity of VSTM.     

A visual short-term memory advantage for faces

What determines how much can be stored in visual short-term memory (VSTM)? Studies of VSTM have focused largely on stimulus-based properties such as the number or complexity of the items stored. Recent work also suggests that capacity is severely reduced for items within the same category. However, the importance for VSTM capacity of more qualitative differences in processing for different categories has not been investigated. For example, faces are processed more holistically than other objects. In Experiments 1 and 2, we show that the processing of faces, objects that are crucial socially and for which we possess considerable expertise, overcomes these limitations. More faces can be stored in VSTM than objects from other complex nonface categories. As in prior studies, at short encoding durations we found that capacity for faces was less than that for other categories. However, at longer encoding durations, capacity for faces exceeded that for nonface objects, and this advantage was specific to upright faces. Because inversion reduces holistic processing, the interaction of orientation with VSTM capacity—which occurred for faces but not objects in Experiment 3—suggests that it is holistic processing that confers an advantage for face VSTM when sufficient encoding time is allowed.     

Rapid development of feature binding in visual short-term memory

The binding of object identity (color) and location in visual short-term memory (VSTM) was examined in 6.5- to 12.5-month-old infants (N= 144). Although we previously found that by age 6.5 months, infants can represent both color and location in VSTM, in the present study we observed that 6.5-month-old infants could not remember trivially simple color-location combinations across a 300-ms delay. However, 7.5-month-old infants could bind color and location as effectively as 12.5-month-old infants. Control conditions confirmed that the failure of 6.5-month-old infants was not a result of perceptual or attentional limitations. This rapid development of VSTM binding between 6.5 and 7.5 months occurs during a period of rapid increase in VSTM storage capacity and just after a period of dramatic neuroanatomical changes in parietal cortex. Thus, the ability to bind features and the ability to store multiple objects may both depend on a process that is mediated by posterior parietal cortex and is perhaps related to focused attention.     

Understanding the object benefit in visual short-term memory: the roles of feature proximity and connectedness

Past research has identified visual objects as the units of information processing in visual short-term memory (VSTM) and has shown that two features from the same object can be remembered in VSTM as well (or almost as well) as one feature of that object and are much better remembered than the same two features from two spatially separated objects. It is not clear, however, what drives this object benefit in VSTM. Is it the shared spatial location (proximity), the connectedness among features of an object, or both? In six change detection experiments, both location/proximity and connectedness were found to be crucial in determining the magnitude of the object benefit in VSTM. Together, these results indicate that location/proximity and connectedness are essential elements in defining a coherent visual object representation in VSTM.     

The capacity of visual short-term memory within and between hemifields

Visual short-term memory (VSTM) and attention are both thought to have a capacity limit of four items [e.g. Luck, S. J., & Vogel, E. K. (1997). The capacity of visual working memory for features and conjunctions. Nature, 309, 279-281; Pylyshyn, Z. W., & Storm, R. W. (1988). Tracking multiple independent targets: evidence for a parallel tracking mechanism. Spatial Vision, 3, 179-197.]. Using the multiple object visual tracking paradigm (MOT), it has recently been shown that twice as many items can be simultaneously attended when they are separated between two visual fields compared to when they are all presented within the same hemifield [Alvarez, G. A., & Cavanagh, P. (2004). Independent attention resources for the left and right visual hemifields (Abstract). Journal of Vision, 4(8), 29a.]. Does VSTM capacity also increase when the items to be remembered are distributed between the two visual fields? The current paper investigated this central issue in two different tasks, namely a color and spatial location change detection task, in which the items were displayed either in the two visual fields or in the same hemifield. The data revealed that only memory capacity for spatial locations and not colors increased when the items were separated between the two visual fields. These findings support the view of VSTM as a chain of capacity limited operations where the spatial selection of stimuli, which dominates in both spatial location VSTM and MOT, occupies the first place and shows independence between the two fields.     

Separate and shared sources of dual-task cost in stimulus identification and response selection

There is often strong interference if a second target stimulus (T2) is presented before processing of a prior target stimulus (T1) is complete. In the "Psychological Refractory Period" (PRP) paradigm, responses are speeded and interference manifests as increased response time for T2. In the "Attentional Blink" (AB) paradigm, stimuli are masked and responses unspeeded; interference manifests as reduced T2 accuracy. While different causes have usually been considered for PRP and AB phenomena, recent evidence has supported a unified account based on a single, shared restriction on concurrent processing. Here we show that a full assessment of separate and shared resource limitations requires direct comparison of hybrid PRP/AB trials with corresponding pure PRP and AB cases. Randomizing trial types in such a comparison also brings substantial benefit in addressing possible changes in task preparation or readiness. The data from two such experiments--combining speeded auditory (SA) and unspeeded visual (UV) task events--provide clear evidence for both separate and shared resource limitations. Often interference is strongest for T1 and T2 events of the same type, reflecting predominantly different limitations in SA and UV processing. With modest increases in demand, however, interference between different event types can also be made arbitrarily large, reflecting arbitrarily important shared limitations. For even such simple tasks as these, T--T2 interference reflects a combination of relatively local and relatively global sources.     

Visual attention capacity: a review of TVA-based patient studies

Psychophysical studies have identified two distinct limitations of visual attention capacity: processing speed and apprehension span. Using a simple test, these cognitive factors can be analyzed by Bundesen's Theory of Visual Attention (TVA). The method has strong specificity and sensitivity, and measurements are highly reliable. As the method is theoretically founded, it also has high validity. TVA-based assessment has recently been used to investigate a broad range of neuropsychological and neurological conditions. We present the method, including the experimental paradigm and practical guidelines to patient testing, and review existing TVA-based patient studies organized by lesion anatomy. Lesions in three anatomical regions affect visual capacity: The parietal lobes, frontal cortex and basal ganglia, and extrastriate cortex. Visual capacity thus depends on large, bilaterally distributed anatomical networks that include several regions outside the visual system. The two visual capacity parameters are functionally separable, but seem to rely on largely overlapping brain areas.     

Organization of visual short-term memory

The authors examined the organization of visual short-term memory (VSTM). Using a change-detection task, they reported that VSTM stores relational information between individual items. This relational processing is mediated by the organization of items into spatial configurations. The spatial configuration of visual objects is important for VSTM of spatial locations, colors, and shapes. When color VSTM is compared with location VSTM, spatial configuration plays an integral role because configuration is important for color VSTM, whereas color is not important for location VSTM. The authors also examined the role of attention and found that the formation of configuration is modulated by both top-down and bottom-up attentional factors. In summary, the authors proposed that VSTM stores the relational information of individual visual items on the basis of global spatial configuration.     

Is visual short-term memory object based? Rejection of the “strong-object” hypothesis

Is the capacity of visual short-term memory (VSTM) limited by the number of objects or by the number of features? VSTM for objects with either one feature or two color features was tested. Results show that capacity was limited primarily by the number of colors to be memorized, not by the number of objects. This result held up with variations in color saturation, blocked or mixed conditions, duration of memory image, and absence or presence of verbal load. However, conjoining features into objects improved VSTM capacity when size-orientation and color-orientation conjunctions were tested. Nevertheless, the number of features still mattered. When feature heterogeneity was controlled, VSTM for conjoined objects was worse than VSTM for objects made of single features. Our results support a weak-object hypothesis of VSTM capacity that suggests that VSTM is limited by both the number of objects and the feature composition of those objects.     

Parallel consolidation of simple features into visual short-term memory

Although considerable research has examined the storage limits of visual short-term memory (VSTM), little is known about the initial formation (i.e., the consolidation) of VSTM representations. A few previous studies have estimated the capacity of consolidation to be one item at a time. Here we used a sequential-simultaneous manipulation to reexamine the limits of consolidating items into VSTM. Participants viewed briefly presented and masked color patches (targets), which were shown either sequentially or simultaneously. A probe color followed the targets and participants decided whether it matched one of the targets or was a novel color. In four experiments, we consistently found equal performance for sequential and simultaneous presentations for two targets. Worse performance in the simultaneous than the sequential condition was observed for larger set sizes (three and four). Contrary to previous results, suggesting a severe capacity limit of one item, our results indicate that consolidation into VSTM can occur in parallel and without capacity limits for at least two items.     

Is visual short-term memory object based? Rejection of the "strong-object" hypothesis

Is the capacity of visual short-term memory (VSTM) limited by the number of objects or by the number of features? VSTM for objects with either one feature or two color features was tested. Results show that capacity was limited primarily by the number of colors to be memorized, not by the number of objects. This result held up with variations in color saturation, blocked or mixed conditions, duration of memory image, and absence or presence of verbal load. However, conjoining features into objects improved VSTM capacity when size-orientation and color-orientation conjunctions were tested. Nevertheless, the number of features still mattered. When feature heterogeneity was controlled, VSTM for conjoined objects was worse than VSTM for objects made of single features. Our results support a weak-object hypothesis of VSTM capacity that suggests that VSTM is limited by both the number of objects and the feature composition of those objects.     

Enhanced visual short-term memory in action video game players

Visual short-term memory (VSTM) is critical for acquiring visual knowledge and shows marked individual variability. Previous work has illustrated a VSTM advantage among action video game players (Boot et al. Acta Psychologica 129:387–398, 2008). A growing body of literature has suggested that action video game playing can bolster visual cognitive abilities in a domain-general manner, including abilities related to visual attention and the speed of processing, providing some potential bases for this VSTM advantage. In the present study, we investigated the VSTM advantage among video game players and assessed whether enhanced processing speed can account for this advantage. Experiment 1, using simple colored stimuli, revealed that action video game players demonstrate a similar VSTM advantage over nongamers, regardless of whether they are given limited or ample time to encode items into memory. Experiment 2, using complex shapes as the stimuli to increase the processing demands of the task, replicated this VSTM advantage, irrespective of encoding duration. These findings are inconsistent with a speed-of-processing account of this advantage. An alternative, attentional account, grounded in the existing literature on the visuo-cognitive consequences of video game play, is discussed.     

Visual short-term memory load strengthens selective attention

Perceptual load theory accounts for many attentional phenomena; however, its mechanism remains elusive because it invokes underspecified attentional resources. Recent dual-task evidence has revealed that a concurrent visual short-term memory (VSTM) load slows visual search and reduces contrast sensitivity, but it is unknown whether a VSTM load also constricts attention in a canonical perceptual load task. If attentional selection draws upon VSTM resources, then distraction effects—which measure attentional “spill-over”—will be reduced as competition for resources increases. Observers performed a low perceptual load flanker task during the delay period of a VSTM change detection task. We observed a reduction of the flanker effect in the perceptual load task as a function of increasing concurrent VSTM load. These findings were not due to perceptual-level interactions between the physical displays of the two tasks. Our findings suggest that perceptual representations of distractor stimuli compete with the maintenance of visual representations held in memory. We conclude that access to VSTM determines the degree of attentional selectivity; when VSTM is not completely taxed, it is more likely for task-irrelevant items to be consolidated and, consequently, affect responses. The “resources” hypothesized by load theory are at least partly mnemonic in nature, due to the strong correspondence they share with VSTM capacity.     

Visual short-term memory benefit for objects on different 3-D surfaces

Visual short-term memory (VSTM) plays an important role in visual cognition. Although objects are located on different 3-dimensional (3-D) surfaces in the real world, how VSTM capacity may be influenced by the presence of multiple 3-D surfaces has never been examined. By manipulating binocular disparities of visual displays, the authors found that more colored objects could be held in VSTM when they were placed on 2 rather than on 1 planar 3-D surfaces. This between-surface benefit in VSTM was present only when binding of objects' colors to their 3-D locations was required (i.e., when observers needed to remember which color appeared where). When binding was not required, no between-surface benefit in VSTM was observed. This benefit in VSTM could not be attributed to the number of spatial locations attended within a given surface. It was not due to a general perceptual grouping effect either, because grouping by motion and grouping by different regions of the same surface did not yield the same benefit. This increment in capacity indicates that VSTM benefits from the placement of objects in a 3-D scene.     

Orienting attention to locations in mental representations

Many cognitive processes depend on our ability to hold information in mind, often well beyond the offset of the original sensory input. The capacity of this visual short-term memory (VSTM) is limited to around three to four items. Recent research has demonstrated that the content of VSTM can be modulated by top-down attentional biases. This has been demonstrated using retrodictive spatial cues, termed "retro-cues," which orient subjects' attention to spatial locations within VSTM. In the present article, we tested whether the use of these cues is modulated by memory load and cue delay. There are a number of important conclusions: (1) Top-down biases can operate on very brief iconic traces as well as on older VSTM representations (Exp. 1). (2) When operating within capacity, subjects use the cue to prioritise where they initiate their memory search, rather than to discard uncued items (Exps. 2 and 3). (3) When capacity is exceeded, there is little benefit to top-down biasing relative to a neutral condition; however, unattended items are lost, with there being a substantial cost of invalid spatial cueing (Exp. 3). (4) These costs and benefits of orienting spatial attention differ across iconic memory and VSTM representations when VSTM capacity is exceeded (Exp. 4).     

Improving visual short-term memory by sequencing the stimulus array

When multiple objects are presented briefly and simultaneously in a visual array, visual short-term memory (VSTM) can maintain only a limited number of these items. The present research report reveals that splitting the to-be-remembered items into two sequential arrays significantly increases VSTM performance relative to the simultaneous presentation of the same items. A memory benefit also emerges when the full object array is flashed twice (repeated) rather than being presented continuously for the same duration. Moreover, the sequential and repetition benefits are specifically pronounced for individuals with low performance for simultaneously presented items. Our results suggest that the conventional, simultaneous presentation mode may underestimate VSTM performance due to attentional limitations and/or competition between stimulus representations. In contrast, temporal segregation of the stimulus input may help participants maximize their performance and utilize their full VSTM capacity.     

What are the units of visual short-term memory, objects or spatial locations?

We investigated whether the capacity of visual short-term memory (VSTM) is defined by number of objects or number of spatial locations. Previous work is consistent with either alternative. To distinguish these factors, we used overlapping stimuli that allowed us to independently manipulate the number of spatial locations while holding constant the number of objects and features to be encoded (Duncan, 1984; Vecera & Farah, 1994). In Experiment 1, the number of spatial locations had no effect on VSTM, suggesting that VSTM is object based. Experiments 2 and 3 ruled out alternative explanations based on perceptual segregation difficulty or decision noise factors. Our results provide additional support to Luck and Vogel's (1997) demonstration that integrated objects form the units of VSTM capacity.     

What kind of memory supports visual marking?

In visual search tasks, if a set of items is presented for 1 s before another set of new items (containing the target) is added, search can be restricted to the new set. The process that eliminates old items from search is visual marking. This study investigates the kind of memory that distinguishes the old items from the new items during search. Using an accuracy paradigm in which perfect marking results in 100% accuracy and lack of marking results in near chance performance, the authors show that search can be restricted to new items not by visual short-term memory (VSTM) of old locations but by a limited capacity and slow-decaying VSTM of new locations and a high capacity and fast-decaying memory for asynchrony.     

Hemifield asymmetries differentiate VSTM for single- and multiple-feature objects

Visual short-term memory (VSTM) is a capacity-limited system for maintaining visual information across brief durations. Limits in the amount of information held in memory reflect processing constraints in the intraparietal sulcus (IPS), a region of the frontoparietal network also involved in visual attention. During VSTM and visual attention, areas of IPS demonstrate hemispheric asymmetries. Whereas the left hemisphere represents information in only the right hemifield, the right hemisphere represents information across the visual field. In visual attention, hemispheric asymmetries are associated with differences in behavioral performance across the visual field. In order to assess the degree of hemifield asymmetries in VSTM, we measured memory performance across the visual field for both single- and two-feature objects. Consistent with theories of right-hemisphere dominance, there was a memory benefit for single-feature items in the left visual hemifield. However, when the number of features increased, the behavioral bias reversed, demonstrating a benefit for remembering two-feature objects in the right hemifield. On an individual basis, the cost of remembering an additional feature in the hemifields was correlated, suggesting that the shift in hemifield biases reflected a redistribution of resources across the visual field. Furthermore, we demonstrate that these results cannot be explained by differences in perceptual or decision-making load. Our results are consistent with a flexible resource model of VSTM in which attention and/or working memory demands result in representation of items in the right hemifield by both the left and right hemispheres.