Reward-priming impacts visual working memory maintenance: Evidence from human electrophysiology

Reward can benefit visual processing of reward-associated objects in a non-strategic way. Recent studies have suggested that such influence extends also to visual working memory (VWM) representations of reward-associated stimuli. However, it is not clear yet which mechanisms underlie the behavioural effects in VWM tasks: reward could directly impact on our ability to maintain representations in VWM or it could influence memory indirectly via priming of attentional selection. To distinguish between these alternatives we measured event-related potential indices of selective attention – the N2pc – and VWM maintenance – the CDA (contralateral delay activity) – while participants completed a VWM task. Results show that reward outcome in one trial caused similarly coloured targets to be strongly represented in VWM in subsequent trials, as expressed in a larger-amplitude CDA This was not preceded by a corresponding effect on attentional selection, in so far as our key manipulation had no impact on the N2pc. In a second experiment, we found that reward priming produced a behavioural benefit that emerged over time, suggesting that the representations of reward-associated items stored in VWM are more resistant to interference and decay. We conclude that when the task stresses VWM maintenance, it is at this representational level that reward will have impact.     

“Wanted!” The effects of reward on face recognition: electrophysiological correlates

The aim of the present study was to uncover the temporal dynamics of face recognition as a function of reward. Event-related potentials (ERPs) were recorded during the encoding and the subsequent old/new memory test in response to faces that could be associated with a monetary reward. The behavioral results showed that faces associated with reward at both encoding and retrieval were recognized better than the unrewarded ones. ERP responses highlighted that successful encoding predictive of subsequent memory was greater for faces associated with reward than for faces without reward-driven motivational learning. At retrieval, an early positive-going component was elicited for potentially rewarded faces on frontal regions, while the occipito-temporal N170 component showed priming effects as a function of reward. Later on, larger centro-parietal ERP components, related to recognition memory, were found selectively for reward-associated faces. Remarkably, electrophysiological responses varied in a graded manner, with the largest amplitude yielded by faces with double reward, followed by faces associated with reward only at encoding. Taken together, the present data show that the processing of outcome expectations affects face structural encoding and increases memory efficiency, yielding a robust and sustained modulation over frontal and temporal areas where reward and memory mechanisms operate in conjunction.     

颜色凸显条件下基于工作记忆内容的视觉注意

采用记忆任务与视觉搜索任务相结合的实验范式,通过操纵背景颜色与刺激颜色,考察了工作记忆内容对视觉注意的引导是否存在强自动性。实验一通过操作背景颜色以突显记忆项目,结果发现即使工作记忆内容在匹配分心物的情况下仍然自动地捕获视觉注意。实验二通过操作背景颜色以突显目标项目,结果发现工作记忆内容对视觉注意的自动引导作用消失。说明了与工作记忆内容匹配的刺激能够自动地引导视觉注意,但是这种引导并非完全自动化,具有一定的条件性。     

Do working memory-driven attention shifts speed up visual awareness?

Previous research has shown that content representations in working memory (WM) can bias attention in favor of matching stimuli in the scene. Using a visual prior-entry procedure, we here investigate whether such WM-driven attention shifts can speed up the conscious awareness of memory-matching relative to memory-mismatching stimuli. Participants were asked to hold a color cue in WM and to subsequently perform a temporal order judgment (TOJ) task by reporting either of two different-colored circles (presented to the left and right of fixation with a variable temporal interval) as having the first onset. One of the two TOJ circles could match the memory cue in color. We found that awareness of the temporal order of the circle onsets was not affected by the contents of WM, even when participants were explicitly informed that one of the TOJ circles would always match the WM contents. The null effect of WM on TOJs was not due to an inability of the memory-matching item to capture attention, since response times to the target in a follow-up experiment were improved when it appeared at the location of the memory-matching item. The present findings suggest that WM-driven attention shifts cannot accelerate phenomenal awareness of matching stimuli in the visual field.     

工作记忆表征精度加工需求对注意引导的影响

采用注意捕获范式, 通过行为和事件相关脑电位(ERP)实验, 探讨工作记忆表征精度加工需求对注意引导的影响, 行为结果发现, 在低精度加工需求条件下, 只有一个工作记忆表征引导注意, 且处于高激活状态的工作记忆表征产生的注意捕获大于低激活状态; 而在高精度加工需求条件下, 有两个工作记忆表征引导注意, 且处于高、低激活状态的工作记忆表征产生的注意捕获没有差异。ERP结果显示, 高精度加工需求条件下诱发的NSW和LPC大于低精度加工需求条件; 在高精度加工需求条件下, 干扰项与记忆项匹配比不匹配时, 诱发更大的N2和更小的N2pc, 而在低精度加工需求条件下, 干扰项与记忆项匹配和不匹配时诱发的N2、N2pc没有差异。研究表明, 工作记忆表征精度加工需求影响注意引导的机制可能是高精度加工需求下, 工作记忆表征消耗的认知资源增加, 搜索目标获得的资源减少, 干扰项捕获的注意增加。     

Monetary rewards influence retrieval orientations

Reward anticipation during learning is known to support memory formation, but its role in retrieval processes is so far unclear. Retrieval orientations, as a reflection of controlled retrieval processing, are one aspect of retrieval that might be modulated by reward. These processes can be measured using the event-related potentials (ERPs) elicited by retrieval cues from tasks with different retrieval requirements, such as via changes in the class of targeted memory information. To determine whether retrieval orientations of this kind are modulated by reward during learning, we investigated the effects of high and low reward expectancy on the ERP correlates of retrieval orientation in two separate experiments. The reward manipulation at study in Experiment 1 was associated with later memory performance, whereas in Experiment 2, reward was directly linked to accuracy in the study task. In both studies, the participants encoded mixed lists of pictures and words preceded by high- or low-reward cues. After 24?h, they performed a recognition memory exclusion task, with words as the test items. In addition to a previously reported material-specific effect of retrieval orientation, a frontally distributed, reward-associated retrieval orientation effect was found in both experiments. These findings suggest that reward motivation during learning leads to the adoption of a reward-associated retrieval orientation to support the retrieval of highly motivational information. Thus, ERP retrieval orientation effects not only reflect retrieval processes related to the sought-for materials, but also relate to the reward conditions with which items were combined during encoding.     

Working memory, perceptual priming, and the perception of hierarchical forms: Opposite effects of priming and working memory without memory refreshing

Previous research has shown that stimuli held in working memory (WM) can influence spatial attention. Using Navon stimuli, we explored whether and how items in WM affect the perception of visual targets at local and global levels in compound letters. Participants looked for a target letter presented at a local or global level while holding a regular block letter as a memory item. An effect of holding the target’s identity in WM was found. When memory items and targets were the same, performance was better than in a neutral condition when the memory item did not appear in the hierarchical letter (a benefit from valid cuing). When the memory item matched the distractor in the hierarchical stimulus, performance was worse than in the neutral baseline (a cost on invalid trials). These effects were greatest when the WM cue matched the global level of the hierarchical stimulus, suggesting that WM biases attention to the global level of form. Interestingly, in a no-memory priming condition, target perception was faster in the invalid condition than in the neutral baseline, reversing the effect in the WM condition. A further control experiment ruled out the effects of WM being due to participants’ refreshing their memory from the hierarchical stimulus display. The data show that information in WM biases the selection of hierarchical forms, whereas priming does not. Priming alters the perceptual processing of repeated stimuli without biasing attention.     

Value-driven attentional and oculomotor capture during goal-directed, unconstrained viewing

Covert shifts of attention precede and direct overt eye movements to stimuli that are task relevant or physically salient. A growing body of evidence suggests that the learned value of perceptual stimuli strongly influences their attentional priority. For example, previously rewarded but otherwise irrelevant and inconspicuous stimuli capture covert attention involuntarily. It is unknown, however, whether stimuli also draw eye movements involuntarily as a consequence of their reward history. Here, we show that previously rewarded but currently task-irrelevant stimuli capture both attention and the eyes. Value-driven oculomotor capture was observed during unconstrained viewing, when neither eye movements nor fixations were required, and was strongly related to individual differences in visual working memory capacity. The appearance of a reward-associated stimulus came to evoke pupil dilation over the course of training, which provides physiological evidence that the stimuli that elicit value-driven capture come to serve as reward-predictive cues. These findings reveal a close coupling of value-driven attentional capture and eye movements that has broad implications for theories of attention and reward learning.     

The modulation of reward priority by top-down knowledge

Reward-associated features capture attention automatically and continue to do so even when the reward contingencies are removed. This profile has led to the hypothesis that rewards belong to a separate class of attentional biases that is neither typically top-down nor bottom-up. The goal of these experiments was to understand the degree to which top-down knowledge can modulate value-driven attentional capture within (a) the timecourse of a single trial and (b) when the reward contingencies change explicitly over trials. The results suggested that top-down knowledge does not affect the size of value-driven attentional capture within a single trial. There were clear top-down modulations in the magnitude of value-driven capture when reward contingencies explicitly changed, but the original reward associations continued to have a persistent bias on attention. These results contribute to a growing body of evidence that reward associations bias attention through mechanisms separate from other top-down and bottom-up attentional biases.     

Cognitive control over working memory biases of selection

Across many studies, researchers have found that representations in working memory (WM) can guide visual attention toward items that match the features of the WM contents. While some researchers have contended that this occurs involuntarily, others have suggested that the impact of WM contents on attention can be strategically controlled. Here, we varied the probability that WM items would coincide with either targets or distractors in a visual search task to examine (1) whether participants could intentionally enhance or inhibit the influence of WM items on attention and (2) whether cognitive control over WM biases would also affect access to the memory contents in a surprise recognition test. We found visual search to be faster when the WM item coincided with the search target, and this effect was enhanced when the memory item reliably predicted the location of the target. Conversely, visual search was slowed when the memory item coincided with a search distractor, and this effect was diminished, but not abolished, when the memory item was reliably associated with distractors. This strategic dampening of the influence of WM items on attention came at a price to memory, however, as participants were slowest to perform WM recognition tests on blocks in which the WM contents were consistently invalid. These results document that attentional capture by WM contents is partly, but not fully, malleable by top-down control, which appears to adjust the state of the WM contents to optimize search behavior. These data illustrate the role of cognitive control in modulating the strength of WM biases of selection, and they support a tight coupling between WM and attention.     

Reduced Value-Driven Attentional Capture Among Children with ADHD Compared to Typically Developing Controls

The current study examined whether children with ADHD were more distracted by a stimulus previously associated with reward, but currently goal-irrelevant, than their typically-developing peers. In addition, we also probed the associated cognitive and motivational mechanisms by examining correlations with other behavioral tasks. Participants included 8–12 year-old children with ADHD (n = 30) and typically developing controls (n = 26). Children were instructed to visually search for color-defined targets and received monetary rewards for accurate responses. In a subsequent search task in which color was explicitly irrelevant, we manipulated whether a distractor item appeared in a previously reward-associated color. We examined whether children responded more slowly on trials with the previously-rewarded distractor present compared to trials without this distractor, a phenomenon referred to as value-driven attentional capture (VDAC), and whether children with and without ADHD differed in the extent to which they displayed VDAC. Correlations among working memory performance, immediate reward preference (delay discounting) and attentional capture were also examined. Children with ADHD were significantly less affected by the presence of the previously rewarded distractor than were control participants. Within the ADHD group, greater value-driven attentional capture was associated with poorer working memory. Although both ADHD and control participants were initially distracted by previously reward-associated stimuli, the magnitude of distraction was larger and persisted longer among control participants.     

非目标工作记忆表征下颜色关联和抽象关联的注意捕获和抑制

采用工作记忆—视觉搜索双任务范式,探讨在非目标工作记忆表征下,颜色关联和抽象关联干扰子的注意捕获和抑制现象。实验一结果发现,颜色关联干扰子在快速反应阶段捕获注意,随后被成功抑制;抽象关联干扰子仍能捕获注意。实验二消除知觉凸显后,结果发现颜色关联条件的抑制消失,干扰子重新捕获注意。结果表明,有效抑制策略受不同关联干扰子加工先后顺序的影响,选择性抑制颜色关联干扰子,而颜色特征的凸显有助于抑制策略形成。     

Dynamic states in working memory modulate guidance of visual attention: Evidence from an n-back paradigm

Items held in working memory (WM) can automatically bias attention when they reappear in visual displays. Recent evidence, however, suggests that WM biases of attention may be reduced under certain conditions, for example with increasing memory load. We employed a dual task paradigm to investigate how WM biases are affected by dynamic updating of memory contents. 1-back and 2-back versions of a memory task with colour stimuli were interrupted at intervals by an unrelated visual search task. Reappearance in the search display of the item that was currently active in WM guided attention, while suppressed or inactive items did not. We conclude that the rapid updating of memory contents facilitates the shifting of memory representations into different activity states on a moment-to-moment basis. The finding is consistent with models that propose that only one item can be “active” in WM at any one time to guide attention.     

The role of attention in the associative binding of emotionally arousing words

In the present study, we examined the role of attention in modulating the memory benefit of emotional arousal for same-valence word pair associations. To assess the role of attention either at encoding or at retrieval, participants studied lists of positive, neutral, and negative words pairs under full attention, divided attention at encoding, or divided attention at retrieval, and then were tested on the single words and on the associations between words. Consistent with past studies, memory accuracy was higher for emotional items than for neutral items, and no memory difference was observed across emotional arousal conditions for associations when encoding occurred under full attention. In contrast, memory accuracy was higher for emotionally arousing items and associations relative to neutral items when encoding occurred under divided attention. Finally, dividing attention at retrieval revealed similar effects across emotion conditions, suggesting that retrieval of emotional stimuli relative to neutral stimuli, unlike encoding, does not benefit from automatic processing. The discussion emphasizes the role of automatic processing during encoding in producing the benefit of emotionally enhanced memory, as well as the extent to which controlled attention is responsible for eliminating or reversing (relative to neutral materials) emotionally enhanced memory for associations. Additionally, the implications of the divided-attention-at-retrieval manipulation include consideration of the way in which emotional items may be consciously processed during encoding.     

Individual differences in autistic trait load in the general population predict visual working memory performance

Prior studies have reported instances of both intact and impaired working memory (WM) performance in people with autism spectrum disorder (ASD). In order to investigate the relation between autistic traits that extend into the normal population and WM, 104 normal college-aged students who varied in their levels of autistic traits were tested. The loading of ASD-associated traits in the normal population leads to differing predictions about WM performance. ASD traits related to a local processing style (or “attention to detail”) might enhance WM while ASD-associated traits related to difficulty switching attention and reorienting focus (or “social interaction”) might impair WM performance. To assess these predictions, participants filled out the Autism Spectrum Quotient (AQ) and performed a working memory task with both visual and verbal variants. AQ scores were then broken into “attention to detail” and “social interaction” factors, as proposed by Hoekstra and colleagues. The results showed that AQ scores did not predict verbal WM performance but they did predict visual WM performance. The social interaction and attention to detail factors of the AQ had opposing relationships with visual WM performance: A higher level of social difficulty was associated with significantly poorer visual WM performance while a higher level of attention to detail was associated with enhanced visual WM performance. Further investigation of the relation between AQ and WM using the original five-factor model proposed by Baron-Cohen and colleagues (2001) revealed an association between impoverished imagination and visual WM overall.     

非目标工作记忆表征下颜色关联和抽象关联的注意捕获和抑制

采用工作记忆—视觉搜索双任务范式,探讨在非目标工作记忆表征下,颜色关联和抽象关联干扰子的注意捕获和抑制现象。实验1结果发现,颜色关联干扰子在快速反应阶段捕获注意,随后被成功抑制;抽象关联干扰子仍能捕获注意。实验2消除知觉凸显后,结果发现颜色关联条件的抑制消失,干扰子重新捕获注意。结果表明,有效抑制策略受不同关联干扰子加工先后顺序的影响,选择性抑制颜色关联干扰子,而颜色特征的凸显有助于抑制策略形成。     

Mixed signals: The effect of conflicting reward- and goal-driven biases on selective attention

Attentional selection depends on the interaction between exogenous (stimulus-driven), endogenous (goal-driven), and selection history (experience-driven) factors. While endogenous and exogenous biases have been widely investigated, less is known about their interplay with value-driven attention. The present study investigated the interaction between reward-history and goal-driven biases on perceptual sensitivity (d’) and response time (RT) in a modified cueing paradigm presenting two coloured cues, followed by sinusoidal gratings. Participants responded to the orientation of one of these gratings. In Experiment 1, one cue signalled reward availability but was otherwise task irrelevant. In Experiment 2, the same cue signalled reward, and indicated the target’s most likely location at the opposite side of the display. This design introduced a conflict between reward-driven biases attracting attention and goal-driven biases directing it away. Attentional effects were examined comparing trials in which cue and target appeared at the same versus opposite locations. Two interstimulus interval (ISI) levels were used to probe the time course of attentional effects. Experiment 1 showed performance benefits at the location of the reward-signalling cue and costs at the opposite for both ISIs, indicating value-driven capture. Experiment 2 showed performance benefits only for the long ISI when the target was at the opposite to the reward-associated cue. At the short ISI, only performance costs were observed. These results reveal the time course of these biases, indicating that reward-driven effects influence attention early but can be overcome later by goal-driven control. This suggests that reward-driven biases are integrated as attentional priorities, just as exogenous and endogenous factors.     

Automatic guidance of visual attention from verbal working memory

Previous studies have shown that visual attention can be captured by stimuli matching the contents of working memory (WM). Here, the authors assessed the nature of the representation that mediates the guidance of visual attention from WM. Observers were presented with either verbal or visual primes (to hold in memory, Experiment 1; to verbalize, Experiment 2; or merely to attend, Experiment 3) and subsequently were required to search for a target among different distractors, each embedded within a colored shape. In half of the trials, an object in the search array matched the prime, but this object never contained the target. Despite this, search was impaired relative to a neutral baseline in which the prime and search displays did not match. An interesting finding is that verbal primes were effective in generating the effects, and verbalization of visual primes elicited similar effects to those elicited when primes were held in WM. However, the effects were absent when primes were only attended. The data suggest that there is automatic encoding into WM when items are verbalized and that verbal as well as visual WM can guide visual attention.     

The plateau in mnemonic resolution across large set sizes indicates discrete resource limits in visual working memory

The precision of visual working memory (WM) representations declines monotonically with increasing storage load. Two distinct models of WM capacity predict different shapes for this precision-by-set-size function. Flexible-resource models, which assert a continuous allocation of resources across an unlimited number of items, predict a monotonic decline in precision across a large range of set sizes. Conversely, discrete-resource models, which assert a relatively small item limit for WM storage, predict that precision will plateau once this item limit is exceeded. Recent work has demonstrated such a plateau in mnemonic precision. Moreover, the set size at which mnemonic precision reached asymptote has been strongly predicted by estimated item limits in WM. In the present work, we extend this evidence in three ways. First, we show that this empirical pattern generalizes beyond orientation memory to color memory. Second, we rule out encoding limits as the source of discrete limits by demonstrating equivalent performance across simultaneous and sequential presentations of the memoranda. Finally, we demonstrate that the analytic approach commonly used to estimate precision yields flawed parameter estimates when the range of stimulus space is narrowed (e.g., a 180o rather than a 360o orientation space) and typical numbers of observations are collected. Such errors in parameter estimation reconcile an apparent conflict between our findings and others based on different stimuli. These findings provide further support for discrete-resource models of WM capacity.     

Value-driven attentional capture is modulated by spatial context

When stimuli are associated with reward outcome, their visual features acquire high attentional priority such that stimuli possessing those features involuntarily capture attention. Whether a particular feature is predictive of reward, however, will vary with a number of contextual factors. One such factor is spatial location: for example, red berries are likely to be found in low-lying bushes, whereas yellow bananas are likely to be found on treetops. In the present study, I explore whether the attentional priority afforded to reward-associated features is modulated by such location-based contingencies. The results demonstrate that when a stimulus feature is associated with a reward outcome in one spatial location but not another, attentional capture by that feature is selective to when it appears in the rewarded location. This finding provides insight into how reward learning effectively modulates attention in an environment with complex stimulus–reward contingencies, thereby supporting efficient foraging.