Decoding of Working Memory Contents in Auditory Cortex Is Not Distractor-Resistant.

Working memory enables the temporary storage of relevant information in the service of behavior. Neuroimaging studies have suggested that sensory cortex is involved in maintaining contents in working memory. This raised the question of how sensory regions maintain memory representations during the exposure to distracting stimuli. Multivariate pattern analysis of fMRI signals in visual cortex has shown that the contents of visual working memory could be decoded concurrently with passively viewed distractors. The present fMRI study tested whether this finding extends to auditory working memory and to active distractor processing. We asked participants to memorize the pitch of a target sound and to compare it with a probe sound presented after a 13 s delay period. In separate conditions, we compared a blank delay phase (no distraction) with either passive listening to, or active processing of, an auditory distractor presented throughout the memory delay. Consistent with previous reports, pitch-specific memory information could be decoded in auditory cortex during the delay in trials without distraction. In contrast, decoding of target sounds in early auditory cortex dropped to chance level during both passive and active distraction. This was paralleled by memory performance decrements under distraction. Extending the analyses beyond sensory cortex yielded some evidence for memory content-specific activity in inferior frontal and superior parietal cortex during active distraction. In summary, while our findings question the involvement of early auditory cortex in the maintenance of distractor-resistant working memory contents, further research should elucidate the role of hierarchically higher regions.SIGNIFICANCE STATEMENT Information about sensory features held in working memory can be read out from hemodynamic activity recorded in human sensory cortices. Moreover, visual cortex can in parallel store visual content and process newly incoming, task-irrelevant visual input. The present study investigated the role of auditory cortex for working memory maintenance under distraction. While memorized sound frequencies could be decoded in auditory cortex in the absence of distraction, auditory distraction during the delay phase impaired memory performance and prevented decoding of information stored in working memory. Apparently, early auditory cortex is not sufficient to represent working memory contents under distraction that impairs performance. However, exploratory analyses indicated that, under distraction, higher-order frontal and parietal regions might contribute to content-specific working memory storage.

Interference between items stored for distinct tasks in visual working memory.

The action perspective on working memory suggests that memory representations are coded according to their specific temporal and behavioral task demands. This stands in contrast to theories that assume representations are stored in a task-agnostic format within a "common workspace". Here, we tested whether visual items that are memorized for different tasks are stored separately from one another or show evidence of inter-item interference during concurrent maintenance, indicating a common storage. In two experiments, we combined a framing memory task (memorize a motion direction for continuous direction report) with an embedded memory task (memorize a motion direction for a binary direction discrimination) that was placed within the retention period of the framing task. Even though the temporal and action demands were item specific, we observed two types of interference effects between the items: The embedded motion direction was (1) repulsed away and (2) degraded in precision by the motion direction of the item in the framing task. Repulsion and precision degradation increased with item similarity when both items were concurrently held in working memory. In contrast, perceptual and iconic memory control conditions revealed weaker repulsion overall and no interference effect on precision during the stimulus processing stages prior to working memory consolidation. Thus, additional inter-item interference arose uniquely within working memory. Together, our results present evidence that items that are stored for distinct tasks to be performed at distinct points in time, reside in a common workspace in working memory.

Decoding Concurrent Representations of Pitch and Location in Auditory Working Memory.

Multivariate analyses of hemodynamic signals serve to identify the storage of specific stimulus contents in working memory (WM). Representations of visual stimuli have been demonstrated both in sensory regions and in higher cortical areas. While previous research has typically focused on the WM maintenance of a single content feature, it remains unclear whether two separate features of a single object can be decoded concurrently. Also, much less evidence exists for representations of auditory compared with visual stimulus features. To address these issues, human participants had to memorize both pitch and perceived location of one of two sample sounds. After a delay phase, they were asked to reproduce either pitch or location. At recall, both features showed comparable levels of discriminability. Region of interest (ROI)-based decoding of functional magnetic resonance imaging (fMRI) data during the delay phase revealed feature-selective activity for both pitch and location of a memorized sound in auditory cortex and superior parietal lobule. The latter region showed higher decoding accuracy for location than pitch. In addition, location could be decoded from angular and supramarginal gyrus and both superior and inferior frontal gyrus. The latter region also showed a trend for decoding of pitch. We found no region exclusively coding pitch memory information. In summary, the present study yielded evidence for concurrent representations of pitch and location of a single object both in sensory cortex and in hierarchically higher regions, pointing toward representation formats that enable feature integration within the same anatomic brain regions.SIGNIFICANCE STATEMENT Decoding of hemodynamic signals serves to identify brain regions involved in the storage of stimulus-specific information in working memory (WM). While to-be-remembered information typically consists of several features, most previous investigations have focused on the maintenance of one memorized feature belonging to one visual object. The present study assessed the concurrent storage of two features of the same object in auditory WM. We found that both pitch and location of memorized sounds were decodable both in early sensory areas, in higher-level superior parietal cortex and, to a lesser extent, in inferior frontal cortex. While auditory cortex is known to process different features in parallel, their concurrent representation in parietal regions may support the integration of object features in WM.

Decoding Spatial Versus Non-spatial Processing in Auditory Working Memory.

OBJECTIVE: Research on visual working memory has shown that individual stimulus features are processed in both specialized sensory regions and higher cortical areas. Much less evidence exists for auditory working memory. Here, a main distinction has been proposed between the processing of spatial and non-spatial sound features. Our aim was to examine feature-specific activation patterns in auditory working memory. METHODS: We collected fMRI data while 28 healthy adults performed an auditory delayed match-to-sample task. Stimuli were abstract sounds characterized by both spatial and non-spatial information, i.e., interaural time delay and central frequency, respectively. In separate recording blocks, subjects had to memorize either the spatial or non-spatial feature, which had to be compared with a probe sound presented after a short delay. We performed both univariate and multivariate comparisons between spatial and non-spatial task blocks. RESULTS: Processing of spatial sound features elicited a higher activity in a small cluster in the superior parietal lobe than did sound pattern processing, whereas there was no significant activation difference for the opposite contrast. The multivariate analysis was applied using a whole-brain searchlight approach to identify feature-selective processing. The task-relevant auditory feature could be decoded from multiple brain regions including the auditory cortex, posterior temporal cortex, middle occipital gyrus, and extended parietal and frontal regions. CONCLUSION: In summary, the lack of large univariate activation differences between spatial and non-spatial processing could be attributable to the identical stimulation in both tasks. In contrast, the whole-brain multivariate analysis identified feature-specific activation patterns in widespread cortical regions. This suggests that areas beyond the auditory dorsal and ventral streams contribute to working memory processing of auditory stimulus features.

Variability in the analysis of a single neuroimaging dataset by many teams.

Data analysis workflows in many scientific domains have become increasingly complex and flexible. Here we assess the effect of this flexibility on the results of functional magnetic resonance imaging by asking 70 independent teams to analyse the same dataset, testing the same 9 ex-ante hypotheses1. The flexibility of analytical approaches is exemplified by the fact that no two teams chose identical workflows to analyse the data. This flexibility resulted in sizeable variation in the results of hypothesis tests, even for teams whose statistical maps were highly correlated at intermediate stages of the analysis pipeline. Variation in reported results was related to several aspects of analysis methodology. Notably, a meta-analytical approach that aggregated information across teams yielded a significant consensus in activated regions. Furthermore, prediction markets of researchers in the field revealed an overestimation of the likelihood of significant findings, even by researchers with direct knowledge of the dataset2-5. Our findings show that analytical flexibility can have substantial effects on scientific conclusions, and identify factors that may be related to variability in the analysis of functional magnetic resonance imaging. The results emphasize the importance of validating and sharing complex analysis workflows, and demonstrate the need for performing and reporting multiple analyses of the same data. Potential approaches that could be used to mitigate issues related to analytical variability are discussed.

Context information supports serial dependence of multiple visual objects across memory episodes.

Serial dependence is thought to promote perceptual stability by compensating for small changes of an object's appearance across memory episodes. So far, it has been studied in situations that comprised only a single object. The question of how we selectively create temporal stability of several objects remains unsolved. In a memory task, objects can be differentiated by their to-be-memorized feature (content) as well as accompanying discriminative features (context). We test whether congruent context features, in addition to content similarity, support serial dependence. In four experiments, we observe a stronger serial dependence between objects that share the same context features across trials. Apparently, the binding of content and context features is not erased but rather carried over to the subsequent memory episode. As this reflects temporal dependencies in natural settings, our findings reveal a mechanism that integrates corresponding content and context features to support stable representations of individualized objects over time.

Visual objects interact differently during encoding and memory maintenance.

The storage mechanisms of working memory are the matter of an ongoing debate. The sensory recruitment hypothesis states that memory maintenance and perceptual encoding rely on the same neural substrate. This suggests that the same cortical mechanisms that shape object perception also apply to maintained memory content. We tested this prediction using the Direction Illusion, i.e., the mutual repulsion of two concurrently visible motion directions. Participants memorized the directions of two random dot patterns for later recall. In Experiments 1 and 2, we varied the temporal separation of spatially distinct stimuli to manipulate perceptual concurrency, while keeping concurrency within working memory constant. We observed mutual motion repulsion only under simultaneous stimulus presentation, but proactive repulsion and retroactive attraction under immediate stimulus succession. At inter-stimulus intervals of 0.5 and 2 s, however, proactive repulsion vanished, while the retroactive attraction remained. In Experiment 3, we presented both stimuli at the same spatial position and observed a reappearance of the repulsion effect. Our results indicate that the repulsive mechanisms that shape object perception across space fade during the transition from a perceptual representation to a consolidated memory content. This suggests differences in the underlying structure of perceptual and mnemonic representations. The persistence of local interactions, however, indicates different mechanisms of spatially global and local feature interactions.

Gaze-based and attention-based rehearsal in spatial working memory.

How do we maintain information about spatial configurations in mind? Many working memory (WM) models assume that rehearsal processes are used to counteract forgetting in WM. Here, we investigated the contributions of gaze-based and attention-based rehearsal for protecting spatial representations from time-based forgetting. Participants memorized 6 locations selected from a grid of 30 scattered dots. Memory was tested after 1.5 or 4.5 s, and this interval was either blank or the grid remained onscreen (which is assumed to provide rehearsal support). In 2 experiments, we monitored eye movements during the retention phase, or asked participants to fixate the screen center. In 3 subsequent experiments, we tested spatial WM under dual-task conditions inhibiting shifts of visuospatial attention or central attention to the memoranda. Memory was better and more resistant to time-based forgetting in the grid than blank condition. Recording of fixations showed more frequent and efficient gaze-based rehearsal in the presence of the grid. Fixations toward distractor locations occurred at a similar frequency in the blank and grid conditions, and it did not predict incorrect recalls. Inhibition of eye-movements or shifts of visuospatial attention impaired memory overall, but it did not change the grid benefit nor the rate of time-based forgetting. In contrast, distracting central attention increased time-based forgetting regardless of grid presence. These results indicate that (a) the grid benefit is only partially explained by rehearsal; (b) gaze-errors (i.e., distractor fixations) do not lead to more forgetting; and (c) the maintenance of spatial representations over time depends on central processing. (PsycInfo Database Record (c) 2020 APA, all rights reserved).

On-item fixations during serial encoding do not affect spatial working memory.

Ample evidence suggests that there is overlap between the eye-movement system and spatial working memory. Such overlapping structures or capacities may result in interference on the one hand and beneficial support on the other. We investigated eye-movement control during encoding of verbal or spatial information, keeping the display the same between tasks. Saccades to to-be-encoded items were scarce during spatial encoding in comparison with verbal encoding. However, despite replicating this difference across different tasks (serial, free recall) and presentation modalities (simultaneous, sequential presentation), we found no relation between item fixations and memory performance-that is, no costs or benefits. Inducing a change from covert to overt encoding did not affect spatial memory performance as well. In contrast, regressive fixations on prior items, that were no longer on the screen, were associated with increased spatial memory performance. Regressions occurred mainly at the end of the encoding period and were targeted at the first presented item. Our results suggest a dissociation between two types of fixations that accompany serial spatial memory: On-item fixations are epiphenomenal; regressions indicate rehearsal or output preparation.

Two types of serial dependence in visual working memory.

Stimulus representations in working memory depend on memory traces of past stimuli both from previous trials and from the current trial. However, it is unclear whether the same or different mechanisms underlie this serial dependence across and within trials. We directly contrasted estimates of bias for pairs of immediately successive stimuli across and within trials. In each trial, participants memorized two consecutive motion direction stimuli (S1 and S2) and after a short delay were cued to report one of them. We found serial dependence across trials: The current S1 was attracted towards the preceding S2 when the latter had been cued for report. In contrast, within the same trial S2 was repulsed from S1. In addition, repulsion within a trial occurred for a broader range of motion direction differences between stimuli than attraction across trials. A second experiment in which 25% of trials did not require a response demonstrated that across-trial attraction did not depend on whether the previous S2 actually had to be reported. Our findings provide evidence for two types of serial dependence operating across and within trials. They support the notion of different mechanisms integrating or segregating current from similar past memory contents depending on their task relevance.

Sequential whole report accesses different states in visual working memory.

Working memory (WM) enables a rapid access to a limited number of items that are no longer physically present. WM studies usually involve the encoding and retention of multiple items, while probing a single item only. Hence, little is known about how well multiple items can be reported from WM. Here we asked participants to successively report each of up to 8 encoded Gabor patches from WM. Recall order was externally cued, and stimulus orientations had to be reproduced on a continuous dimension. Participants were able to sequentially report items from WM with an above-chance precision even at high set sizes. It is important that we observed that precision varied systematically with report order: It dropped steeply from the first to the second report but decreased only slightly thereafter. The observed trajectory of precision decrease across reports was better captured as a discontinuous rather than an exponential function, suggesting that items were reported from different states in visual WM. The following 3 experiments replicated these findings. In particular, they showed that the observed drop could not be explained by a retro-cueing benefit of the first report, a longer delay duration for later reports or a visual interference effect of the first report. Instead, executive interference of the first report reduced precision of subsequent reports. Together, the results show that a sequential whole-report procedure allows the assessment of qualitatively different states in visual WM. (PsycINFO Database Record

Automatized smoking-related action schemata are reflected by reduced fMRI activity in sensorimotor brain regions of smokers.

In the later stages of addiction, automatized processes play a prominent role in guiding drug-seeking and drug-taking behavior. However, little is known about the neural correlates of automatized drug-taking skills and drug-related action knowledge in humans. We employed functional magnetic resonance imaging (fMRI) while smokers and non-smokers performed an orientation affordance task, where compatibility between the hand used for a behavioral response and the spatial orientation of a priming stimulus leads to shorter reaction times resulting from activation of the corresponding motor representations. While non-smokers exhibited this behavioral effect only for control objects, smokers showed the affordance effect for both control and smoking-related objects. Furthermore, smokers exhibited reduced fMRI activation for smoking-related as compared to control objects for compatible stimulus-response pairings in a sensorimotor brain network consisting of the right primary motor cortex, supplementary motor area, middle occipital gyrus, left fusiform gyrus and bilateral cingulate gyrus. In the incompatible condition, we found higher fMRI activation in smokers for smoking-related as compared to control objects in the right primary motor cortex, cingulate gyrus, and left fusiform gyrus. This suggests that the activation and performance of deeply embedded, automatized drug-taking schemata employ less brain resources. This might reduce the threshold for relapsing in individuals trying to abstain from smoking. In contrast, the interruption or modification of already triggered automatized action representations require increased neural resources.