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.     

Spatial working memory maintenance: Does attention play a role? A visual search study

Recent studies have proposed that a common mechanism may underlie spatial attention and spatial working memory. One proposal is that spatial working memory is maintained by attention-based rehearsal [Awh, E., Jonides, J., & Reuter-Lorenz, P. A. (1998). Rehearsal in spatial working memory. Journal of Experimental Psychology: Human Perception and Performance, 24(3), 780-790], and so a spatial attention shift during the retention interval of a spatial location should impair its memory performance. In the present study, participants engaged in single-item, parallel or serial search tasks while remembering a spatial location. Although memory tended to bias all searches, the need for an attentional shift during the retention interval impaired memory performance only in single-item search, but not in other searches. These findings suggest that previous evidence for the attention-based rehearsal account does not generalize to visual search. Results are discussed with regard to the relationship between spatial attention and spatial working memory.     

No functional role of attention-based rehearsal in maintenance of spatial working memory representations

The present study systematically examined the role of attention in maintenance of spatial representations in working memory as proposed by the attention-based rehearsal hypothesis [Awh, E., Jonides, J., & Reuter-Lorenz, P. A. (1998). Rehearsal in spatial working memory. Journal of Experimental Psychology - Human Perception and Performance, 24(3), 780-790]. Three main issues were examined. First, Experiments 1-3 demonstrated that inhibition and not facilitation of visual processing is often observed at the memorized location during the retention interval. This inhibition was caused by keeping a location in memory and not by the exogenous nature of the memory cue. Second, Experiment 4 showed that inhibition of the memorized location does not lead to any significant impairment in memory accuracy. Finally, Experiment 5 connected current results to the previous findings and demonstrated facilitation of processing at the memorized location. Importantly, facilitation of processing did not lead to more accurate memory performance. The present results challenge the functional role of attention in maintenance of spatial working memory representations.     

Overt is no better than covert when rehearsing visuo-spatial information in working memory

In the present study, we examined whether eye movements facilitate retention of visuo-spatial information in working memory. In two experiments, participants memorised the sequence of the spatial locations of six digits across a retention interval. In some conditions, participants were free to move their eyes during the retention interval, but in others they either were required to remain fixated or were instructed to move their eyes exclusively to a selection of the memorised locations. Memory performance was no better when participants were free to move their eyes during the memory interval than when they fixated a single location. Furthermore, the results demonstrated a primacy effect in the eye movement behaviour that corresponded with the memory performance. We conclude that overt eye movements do not provide a benefit over covert attention for rehearsing visuo-spatial information in working memory.     

Remembering a location makes the eyes curve away

Working memory is a system that keeps limited information on-line for immediate access by cognitive processes. This type of active maintenance is important for everyday life activities. The present study shows that maintaining a location in spatial working memory affects the trajectories of saccadic eye movements toward visual targets, as the eyes deviate away from the remembered location. This finding provides direct evidence for a strong overlap between spatial working memory and the eye movement system. We argue that curvature is the result of the need to inhibit memory-based eye movement activity in the superior colliculus, in order to allow an accurate saccade to the visual target. Whereas previous research has shown that the eyes may deviate away from visually presented stimuli that need to be ignored, we show that the eyes also curve away from remembered stimuli.     

Rehearsal in Spatial Working Memory: Evidence From Neuroimaging

A variety of biological evidence has identified a frontal-parietal circuit underlying spatial working memory for visual stimuli. But the question remains, how do these neural regions accomplish the goal of maintaining location information on-line? We tested the hypothesis that the active rehearsal of spatial information in working memory is accomplished by means of focal shifts of spatial selective attention to memorized locations. Spatial selective attention has been shown to cause changes in the early visual processing of stimuli that appear in attended locations. Thus, the hypothesis of attention-based rehearsal predicts similar modulations of visual processing at memorized locations. We used functional magnetic resonance imaging to observe posterior visual activations during the performance of a spatial working memory task. In line with the hypothesis, spatial rehearsal led to enhanced activation in the early visual areas contralateral to the memorized locations.     

The time course of spatial memory distortions

Four experiments investigated the memory distortions for the location of a dot in relation to two horizontally aligned landmarks. In Experiment 1, participants reproduced from memory a dot location with respect to the two landmarks. Their performance showed a systematic pattern of distortion that was consistent across individual participants. The three subsequent experiments investigated the time course of spatial memory distortions. Using a visual discrimination task, we were able to map the emergence of spatial distortions within the first 800 msec of the retention interval. After retention intervals as brief as 50 msec, a distortion was already present. In all but one experiment, the distortion increased with longer retention intervals. This early onset of spatial memory distortions might reflect the almost immediate decay of detailed spatial information and the early influence of an enduring spatial memory representation, which encodes spatial information in terms of the perceived structure of space.     

The effects of a task-irrelevant visual event on spatial working memory

In the present experiment, we investigated whether the memory of a location is affected by the occurrence of an irrelevant visual event. Participants had to memorize the location of a dot. During the retention interval, a task-irrelevant stimulus was presented with abrupt onset somewhere in the visual field. Results showed that the spatial memory representation was affected by the occurrence of the external irrelevant event relative to a control condition in which there was no external event. Specifically, the memorized location was shifted toward the location of the task-irrelevant stimulus. This effect was only present when the onset was close in space to the memory representation. These findings suggest that the “internal” spatial map used for keeping a location in spatial working memory and the “external” spatial map that is affected by exogenous events in the outside world are either the same or tightly linked.     

Spatio-temporal working-memory and short-term object-location tasks use different memory mechanisms

Spatial short-term memory for objects' locations was investigated in a spatial relocation task. During maintenance, dynamic visual noise or spatial tapping were administered as visual or spatial secondary tasks, respectively. Because memory for location should tap the visual component of working memory, a visual but not a spatial secondary task should impair location memory. In fact, neither of the tasks impaired memory (Experiment 1), although the expected dissociation between visual and spatial components was clearly confirmed for a spatio-temporal main task (Corsi test) (Experiment 2). We then contrasted location memory for pictures of objects and of nonsense figures under visual interference. Real objects were relocated much better than nonsense figures, and visual noise was again ineffective (Experiment 3). When spatial tapping was combined with the same material (Experiment 3a), again no influence on memory for locations of objects was observed and only a small influence on remembering nonsense figures. We suggest that the Corsi and the relocation VSWM-tasks use different memory mechanisms. The configuration of objects is reconstructed from perceptual records in an episodic buffer, provided by the same structures that enable visual memory after longer intervals. Rehearsal is not necessary for the persistence of these traces. In contrast, in the Corsi task remembering, a temporal sequence across homogeneous locations needs spatio-temporal marking and therefore active rehearsal of the locations by shifting spatial attention. A spatially demanding secondary task during retention interrupts this rehearsal.     

Attention on our mind: The role of spatial attention in visual working memory

The current study shows that spatial visual attention is used to retrieve information from visual working memory. Participants had to keep four colored circles in visual working memory. While keeping this information in memory we asked whether one of the colors was present in the array. While retrieving this information, on some trials a probe dot was presented. When this probe dot was presented at the location of the color that had to be retrieved, participants responded faster than when it was presented at another location. Our findings further elaborate the role of visual attention in working memory: not only is attention the mechanism by which information is stored into working memory, it is also the mechanism by which information is retrieved from visual working memory.     

视觉工作记忆中注意模板的表征—来自EROS的证据

采用事件相关光信号技术,分析再认任务和搜索任务下脑区的动态激活,探讨视觉工作记忆中注意模板的表征机制。实验要求被试在记忆项目后,分别完成再认任务和搜索任务。结果发现,与再认任务相比,视觉搜索任务下记忆项诱发前额叶、枕叶更高程度的激活,且前额叶的激活增强早于枕叶。研究表明,与用于再认任务的普通记忆项相比,用于视觉搜索任务的注意模板的潜在表征机制可能是：先激活前额叶以增强注意控制,并对枕叶的活动加以调节,增强注意模板表征的复述。     

Working memory load reduces the late positive potential and this effect is attenuated with increasing anxiety

Emotion regulation decreases the processing of arousing stimuli, as indexed by the late positive potential (LPP), an electrocortical component that varies in amplitude with emotional arousal. Emotion regulation increases activity in the prefrontal areas associated with cognitive control, including the dosolateral prefrontal cortex (DLPFC). The present study manipulated working memory load, known to activate the DLPFC, and recorded the LPP elicited by aversive and neutral IAPS pictures presented during the retention interval. The LPP was larger on low-load compared to high-load trials, and on trials with aversive compared to neutral pictures. These LPP data suggest that emotional content and working memory load have opposing effects on attention to distracting stimuli. State anxiety was associated with reduced modulation of the LPP by working memory load. Results are discussed in terms of competition for attention between emotion and cognition and suggest a relationship between DLPFC activation and the allocation of attentional resources to distracting visual stimuli–a relationship that may be disrupted with increasing anxiety.     

Working memory, long-term memory, and medial temporal lobe function

Early studies of memory-impaired patients with medial temporal lobe (MTL) damage led to the view that the hippocampus and related MTL structures are involved in the formation of long-term memory and that immediate memory and working memory are independent of these structures. This traditional idea has recently been revisited. Impaired performance in patients with MTL lesions on tasks with short retention intervals, or no retention interval, and neuroimaging findings with similar tasks have been interpreted to mean that the MTL is sometimes needed for working memory and possibly even for visual perception itself. We present a reappraisal of this interpretation. Our main conclusion is that, if the material to be learned exceeds working memory capacity, if the material is difficult to rehearse, or if attention is diverted, performance depends on long-term memory even when the retention interval is brief. This fundamental notion is better captured by the terms subspan memory and supraspan memory than by the terms short-term memory and long-term memory. We propose methods for determining when performance on short-delay tasks must depend on long-term (supraspan) memory and suggest that MTL lesions impair performance only when immediate memory and working memory are insufficient to support performance. In neuroimaging studies, MTL activity during encoding is influenced by the memory load and correlates positively with long-term retention of the material that was presented. The most parsimonious and consistent interpretation of all the data is that subspan memoranda are supported by immediate memory and working memory and are independent of the MTL.     

Overlapping mechanisms of attention and spatial working memory

Spatial selective attention and spatial working memory have largely been studied in isolation. Studies of spatial attention have provided clear evidence that observers can bias visual processing towards specific locations, enabling faster and better processing of information at those locations than at unattended locations. We present evidence supporting the view that this process of visual selection is a key component of rehearsal in spatial working memory. Thus, although working memory has sometimes been depicted as a storage system that emerges 'downstream' of early sensory processing, current evidence suggests that spatial rehearsal recruits top-down processes that modulate the earliest stages of visual analysis.     

不同注意对视觉工作记忆外部特征绑定的影响

采用变化觉察范式,在工作记忆保持阶段插入客体特征辨别或视觉搜索次任务,探讨基于客体的注意和基于空间的注意对视觉工作记忆外部特征绑定表征的影响。结果发现,与单特征相比,两类次任务均对外部特征绑定的记忆成绩损耗更大。这表明视觉工作记忆中外部特征绑定的表征比单特征消耗更多基于客体的注意和更多基于空间的注意。     

Persistent activity in the prefrontal cortex during working memory

The dorsolateral prefrontal cortex (DLPFC) plays a crucial role in working memory. Notably, persistent activity in the DLPFC is often observed during the retention interval of delayed response tasks. The code carried by the persistent activity remains unclear, however. We critically evaluate how well recent findings from functional magnetic resonance imaging studies are compatible with current models of the role of the DLFPC in working memory. These new findings suggest that the DLPFC aids in the maintenance of information by directing attention to internal representations of sensory stimuli and motor plans that are stored in more posterior regions.     

A test of the reward-value hypothesis

Rats retain source memory (memory for the origin of information) over a retention interval of at least 1 week, whereas their spatial working memory (radial maze locations) decays within approximately 1 day. We have argued that different forgetting functions dissociate memory systems. However, the two tasks, in our previous work, used different reward values. The source memory task used multiple pellets of a preferred food flavor (chocolate), whereas the spatial working memory task provided access to a single pellet of standard chow-flavored food at each location. Thus, according to the reward-value hypothesis, enhanced performance in the source memory task stems from enhanced encoding/memory of a preferred reward. We tested the reward-value hypothesis by using a standard 8-arm radial maze task to compare spatial working memory accuracy of rats rewarded with either multiple chocolate or chow pellets at each location using a between-subjects design. The reward-value hypothesis predicts superior accuracy for high-valued rewards. We documented equivalent spatial memory accuracy for high- and low-value rewards. Importantly, a 24-h retention interval produced equivalent spatial working memory accuracy for both flavors. These data are inconsistent with the reward-value hypothesis and suggest that reward value does not explain our earlier findings that source memory survives unusually long retention intervals.     

Binding across space and time in visual working memory

Recent studies of visual short-term memory have suggested that the binding of features such as color and shape into remembered objects is relatively automatic. A series of seven experiments broadened this investigation by comparing the immediate retention of colored shapes with performance when color and shape were separated either spatially or temporally, with participants required actively to form the bound object. Attentional load was manipulated with a demanding concurrent task, and retention in working memory was then tested using a single recognition probe. Both spatial and temporal separation of features tended to impair performance, as did the concurrent task. There was, however, no evidence for greater attentional disruption of performance as a result of either spatial or temporal separation of features. Implications for the process of binding in visual working memory are discussed, and an interpretation is offered in terms of the episodic buffer component of working memory, which is assumed to be a passive store capable of holding bound objects, but not of performing the binding.     

Interactions between working memory, attention and eye movements

This paper reviews the recent findings on working memory, attention and eye movements. We discuss the research that shows that many phenomena related to visual attention taking place when selecting relevant information from the environment are similar to processes needed to keep information active in working memory. We discuss new data that show that when retrieving information from working memory, people may allocate visual spatial attention to the empty location in space that used to contain the information that has to be retrieved. Moreover, we show that maintaining a location in working memory not only may involve attention rehearsal, but might also recruit the oculomotor system. Recent findings seem to suggest that remembering a location may involve attention-based rehearsal in higher brain areas, while at the same time there is inhibition of specific motor programs at lower brain areas. We discuss the possibility that working memory functions do not reside at a special area in the brain, but emerge from the selective recruitment of brain areas that are typically involved in spatial attention and motor control.     

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.