Anna Heuer

Feature-based and spatial attentional selection in visual working memory.

The contents of visual working memory (VWM) can be modulated by spatial cues presented during the maintenance interval (“retrocues”). Here, we examined whether attentional selection of representations in VWM can also be based on features. In addition, we investigated whether the mechanisms of feature-based and spatial attention in VWM differ with respect to parallel access to noncontiguous locations. In two experiments, we tested the efficacy of valid retrocues relying on different kinds of information. Specifically, participants were presented with a typical spatial retrocue pointing to two locations, a symbolic spatial retrocue (numbers mapping onto two locations), and two feature-based retrocues: a color retrocue (a blob of the same color as two of the items) and a shape retrocue (an outline of the shape of two of the items). The two cued items were presented at either contiguous or noncontiguous locations. Overall retrocueing benefits, as compared to a neutral condition, were observed for all retrocue types. Whereas feature-based retrocues yielded benefits for cued items presented at both contiguous and noncontiguous locations, spatial retrocues were only effective when the cued items had been presented at contiguous locations. These findings demonstrate that attentional selection and updating in VWM can operate on different kinds of information, allowing for a flexible and efficient use of this limited system. The observation that the representations of items presented at noncontiguous locations could only be reliably selected with feature-based retrocues suggests that feature-based and spatial attentional selection in VWM rely on different mechanisms, as has been shown for attentional orienting in the external world.

The Focus of Attention in Visual Working Memory: Protection of Focused Representations and Its Individual Variation

Visual working memory can be modulated according to changes in the cued task relevance of maintained items. Here, we investigated the mechanisms underlying this modulation. In particular, we studied the consequences of attentional selection for selected and unselected items, and the role of individual differences in the efficiency with which attention is deployed. To this end, performance in a visual working memory task as well as the CDA/SPCN and the N2pc, ERP components associated with visual working memory and attentional processes, were analysed. Selection during the maintenance stage was manipulated by means of two successively presented retrocues providing spatial information as to which items were most likely to be tested. Results show that attentional selection serves to robustly protect relevant representations in the focus of attention while unselected representations which may become relevant again still remain available. Individuals with larger retrocueing benefits showed higher efficiency of attentional selection, as indicated by the N2pc, and showed stronger maintenance-associated activity (CDA/SPCN). The findings add to converging evidence that focused representations are protected, and highlight the flexibility of visual working memory, in which information can be weighted according its relevance.

Action relevance induces an attentional weighting of representations in visual working memory

Information maintained in visual working memory (VWM) can be strategically weighted according to its task-relevance. This is typically studied by presenting cues during the maintenance interval, but under natural conditions, the importance of certain aspects of our visual environment is mostly determined by intended actions. We investigated whether representations in VWM are also weighted with respect to their potential action relevance. In a combined memory and movement task, participants memorized a number of items and performed a pointing movement during the maintenance interval. The test item in the memory task was subsequently presented either at the movement goal or at another location. We found that performance was better for test items presented at a location that corresponded to the movement goal than for test items presented at action-irrelevant locations. This effect was sensitive to the number of maintained items, suggesting that preferential maintenance of action relevant information becomes particularly important when the demand on VWM is high. We argue that weighting according to action relevance is mediated by the deployment of spatial attention to action goals, with representations spatially corresponding to the action goal benefitting from this attentional engagement. Performance was also better at locations next to the action goal than at locations farther away, indicating an attentional gradient spreading out from the action goal. We conclude that our actions continue to influence visual processing at the mnemonic level, ensuring preferential maintenance of information that is relevant for current behavioral goals.

Different Cortical Mechanisms for Spatial vs. Feature-Based Attentional Selection in Visual Working Memory

The limited capacity of visual working memory (VWM) necessitates attentional mechanisms that selectively update and maintain only the most task-relevant content. Psychophysical experiments have shown that the retroactive selection of memory content can be based on visual properties such as location or shape, but the neural basis for such differential selection is unknown. For example, it is not known if there are different cortical modules specialized for spatial vs. feature-based mnemonic attention, in the same way that has been demonstrated for attention to perceptual input. Here, we used transcranial magnetic stimulation (TMS) to identify areas in human parietal and occipital cortex involved in the selection of objects from memory based on cues to their location (spatial information) or their shape (featural information). We found that TMS over the supramarginal gyrus (SMG) selectively facilitated spatial selection, whereas TMS over the lateral occipital cortex (LO) selectively enhanced feature-based selection for remembered objects in the contralateral visual field. Thus, different cortical regions are responsible for spatial vs. feature-based selection of working memory representations. Since the same regions are involved in terms of attention to external events, these new findings indicate overlapping mechanisms for attentional control over perceptual input and mnemonic representations.

Selective weighting of action-related feature dimensions in visual working memory

Planning an action primes feature dimensions that are relevant for that particular action, increasing the impact of these dimensions on perceptual processing. Here, we investigated whether action planning also affects the short-term maintenance of visual information. In a combined memory and movement task, participants were to memorize items defined by size or color while preparing either a grasping or a pointing movement. Whereas size is a relevant feature dimension for grasping, color can be used to localize the goal object and guide a pointing movement. The results showed that memory for items defined by size was better during the preparation of a grasping movement than during the preparation of a pointing movement. Conversely, memory for color tended to be better when a pointing movement rather than a grasping movement was being planned. This pattern was not only observed when the memory task was embedded within the preparation period of the movement, but also when the movement to be performed was only indicated during the retention interval of the memory task. These findings reveal that a weighting of information in visual working memory according to action relevance can even be implemented at the representational level during maintenance, demonstrating that our actions continue to influence visual processing beyond the perceptual stage.

The necessity to choose causes reward-related anticipatory biasing: Parieto-occipital alpha-band oscillations reveal suppression of low-value targets

Positive outcome of actions can be maximized by choosing the option with the highest reward. For saccades, it has recently been suggested that the necessity to choose is, in fact, an important factor mediating reward effects: latencies to single low-reward targets increased with an increasing proportion of interleaved choice-trials, in which participants were free to choose between two targets to obtain either a high or low reward. Here, we replicate this finding for manual responses, demonstrating that this effect of choice is a more general, effector-independent phenomenon. Oscillatory activity in the alpha and beta band in the preparatory period preceding target onset was analysed for a parieto-occipital and a centrolateral region of interest to identify an anticipatory neural biasing mechanism related to visuospatial attention or motor preparation. When the proportion of interleaved choices was high, an increase in lateralized posterior alpha power indicated that the hemifield associated with a low reward was suppressed in preparation for reward-maximizing target selection. The larger the individual increase in lateralized alpha power, the slower the reaction times to low-reward targets. At a broader level, these findings support the notion that reward only affects responses when behaviour can be optimized to maximize positive outcome.

Separate and combined effects of action relevance and motivational value on visual working memory

Visual working memory contents can be selectively weighted according to differences in their task-relevance. In the present study, we examined the influence of two more indirect selection biases established by a concurrent task or learned reward associations: action relevance and motivational value. In three experiments, memory performance was assessed with the same color change detection task. Potential action relevance and motivational value were each determined by a specific feature of the memory items (location or shape, respectively) and manipulated orthogonally. Investigated separately (Experiments 1A and 1B), both selection biases modulated visual working memory. In combination (Experiment 2), action relevance and motivational value still each exerted an influence, but not in a fully independent and additive manner. While action relevance impacted performance irrespective of the reward associated with the items, an effect of motivational value was only observed for action-relevant items. These results support the notion that visual working memory is automatically biased as an inherent part of action planning. More generally, these findings highlight the versatile nature of visual working memory: Contents can be flexibly weighted to reflect differences in their importance, taking into account several sources of information.

The necessity to choose causes the effects of reward on saccade preparation

When humans have to choose between different options, they can maximize their payoff by choosing the option that yields the highest reward. Information about reward is not only used to optimize decisions but also for movement preparation to minimize reaction times to rewarded targets. Here, we show that this is especially true in contexts in which participants additionally have to choose between different options. We probed eye movement preparation by measuring saccade latencies to differently rewarded single targets (single-trial) appearing left or right from fixation. In choice-trials, both targets were displayed and participants were free to decide for one target to receive the corresponding reward. In blocks without choice-trials, single-trial latencies were not or only weakly affected by reward. With choice-trials present, the influence of reward increased with the proportion and difficulty of choices and decreased when a cue indicated that no choice will be necessary. Choices caused a delay in subsequent single-trial responses to the non-chosen option. Taken together, our results suggest that reward affects saccade preparation mainly when the outcome is uncertain and depends on the participants’ behavior, for instance when they have to choose between targets differing in reward.