Working memory flips the direction of serial bias through memory-based decision.

Reported perception of a new stimulus is either attracted toward or repelled away from task-irrelevant prior stimuli. While prevailing theories propose that the opposing serial biases may stem from distinct stages of information processing, the exact role of working memory (WM) in the serial bias remains unclear despite its consistent involvement in nearly all pertinent studies. Additionally, it is not well understood whether this bias is primarily driven by the biased representation itself or by the decision-making process for the new stimulus. In the present study, we used an orientation delayed estimation paradigm with an attention-demanding intervening task, designed to disrupt the maintenance of stimulus information to investigate the role of WM in serial bias. In the analysis, we scrutinized the trajectory of mouse reports and response time to investigate how the response unfolds over time. Our findings indicate that the serial bias went from repulsive to attractive when WM maintenance was interrupted by the intervening task, and that the associated response trajectories and response time exhibited patterns that cannot be explained by the biased representation alone. These results demonstrate that the task-irrelevant prior stimulus influences the decision for the new stimulus, with the direction of the bias being determined by attentional demand during WM maintenance, thereby placing significant constraints on existing theories on the serial bias effect.

Cardinal bias interacts with the stimulus history bias in orientation working memory.

Reports in a visual working memory(WM) task exhibit biases related to the categorical structure of the stimulus space (e.g., cardinal bias) as well as biases related to previously seen stumuli (e.g., serial bias). While these biases are common and can occur simultaneously, the extent to which they interact in WM remains unknown. In the present study, I used orientation delayed estimation tasks known to produce both cardinal and serial biases and found that the serial bias systematically varied based on the relative positions of the cardinal axis and the preceding stimulus in orientation space. When they were positioned in a way that generated cardinal and serial biases in the same direction (i.e., on the same side of the target orientation), reports for the target orientation exhibited a regular repulsive serial bias. However, when their positions resulted in the biases in the opposite directions (i.e., on the opposite side of the target orientation), no serial bias occurred. This absence of serial bias was replicated in a follow-up experiment where the locations of the stimulus orientation and the response probe were completely randomized, suggesting that the interaction occurs independently from location-based response preparation processes. Together, these results demonstrate that the prior stimulus and the cardinal axis impose interactive impact on the processing of new stimulus, producing differential patterns of serial bias depending on the specific stimulus being processed. These findings place significant implications on computational models addressing the nature of the stimulus history effect and its underlying mechanisms.

Judgments during perceptual comparisons predict distinct forms of memory updating.

Comparing a visual memory with new visual stimuli can bias memory content, especially when the new stimuli are perceived as similar. Perceptual comparisons of this kind may play a mechanistic role in memory updating and can explain how memories can become erroneous in daily life. To test this possibility, we investigated whether comparisons can produce other types of memory distortion beyond memory bias that are commonly implicated in erroneous memories (e.g., memory misattribution). We hypothesized that the type of memory distortion induced during a comparison depends on the perceived overlap between the memory and incoming stimulus-when the input is perceived as similar, it biases memory content; when perceived as the same, it replaces memory content. Participants completed a delayed estimation task in which they compared their memories of color (Experiment 1) and shape stimuli (Experiment 2) to probe stimuli before reporting memory content. We found systematic errors in participants' memory reports following perceived similarity and sameness that were toward the probes and larger following perceived sameness. Simulations confirmed that these errors were not explained by noisy encoding processes that occurred before comparisons. Instead, computational modeling suggested that these errors were likely explained by the probabilistic replacement of the memory by the probe following perceived sameness and integration between the memory and the probe following perceived similarity. Together, these findings suggest that perceptual comparisons can prompt distinct forms of memory updating that have been described previously and may explain how memories become erroneous during their use in everyday behavior. (PsycInfo Database Record (c) 2024 APA, all rights reserved).

Image or Information? Examining the Nature and Impact of Visualization Perceptual Classification.

How do people internalize visualizations: as images or information? In this study, we investigate the nature of internalization for visualizations (i.e., how the mind encodes visualizations in memory) and how memory encoding affects its retrieval. This exploratory work examines the influence of various design elements on a user's perception of a chart. Specifically, which design elements lead to perceptions of visualization as an image (aims to provide visual references, evoke emotions, express creativity, and inspire philosophic thought) or as information (aims to present complex data, information, or ideas concisely and promote analytical thinking)? Understanding how design elements contribute to viewers perceiving a visualization more as an image or information will help designers decide which elements to include to achieve their communication goals. For this study, we annotated 500 visualizations and analyzed the responses of 250 online participants, who rated the visualizations on a bilinear scale as 'image' or 'information.' We then conducted an in-person study ( n = 101) using a free recall task to examine how the image/information ratings and design elements impacted memory. The results revealed several interesting findings: Image-rated visualizations were perceived as more aesthetically 'appealing,' 'enjoyable,' and 'pleasing.' Information-rated visualizations were perceived as less 'difficult to understand' and more aesthetically 'likable' and 'nice,' though participants expressed higher 'positive' sentiment when viewing image-rated visualizations and felt less 'guided to a conclusion.' The presence of axes and text annotations heavily influenced the likelihood of participants rating the visualization as 'information.' We also found different patterns among participants that were older. Importantly, we show that visualizations internalized as 'images' are less effective in conveying trends and messages, though they elicit a more positive emotional judgment, while 'informative' visualizations exhibit annotation focused recall and elicit a more positive design judgment. We discuss the implications of this dissociation between aesthetic pleasure and perceived ease of use in visualization design.

Qualitatively Different Delay-Dependent Working Memory Distortions in People With Schizophrenia and Healthy Control Participants.

BACKGROUND: Impairments in working memory (WM) have been well documented in people with schizophrenia (PSZ). However, these quantitative WM impairments can often be explained by nonspecific factors, such as impaired goal maintenance. Here, we used a spatial orientation delayed response task to explore a qualitative difference in WM dynamics between PSZ and healthy control participants (HCs). More specifically, we took advantage of the discovery that WM representations may drift either toward or away from previous trial targets (serial dependence). We tested the hypothesis that WM representations would drift toward the previous trial target in HCs but away from the previous trial target in PSZ. METHODS: We assessed serial dependence in PSZ (n = 31) and HCs (n = 25) using orientation as the to-be-remembered feature and memory delays lasting from 0 to 8 seconds. Participants were asked to remember the orientation of a teardrop-shaped object and reproduce the orientation after a delay period of varying length. RESULTS: Consistent with prior studies, we found that current trial memory representations were less precise in PSZ than in HCs. We also found that WM for the current trial orientation drifted toward the previous trial orientation in HCs (representational attraction) but drifted away from the previous trial orientation in PSZ (representational repulsion). CONCLUSIONS: These results demonstrate a qualitative difference in WM dynamics between PSZ and HCs that cannot be easily explained by nuisance factors such as reduced effort. Most computational neuroscience models also fail to explain these results because they maintain information solely by means of sustained neural firing, which does not extend across trials. The results suggest a fundamental difference between PSZ and HCs in longer-term memory mechanisms that persist across trials, such as short-term potentiation and neuronal adaptation.

Qualitatively different delay-dependent working memory distortions in people with schizophrenia and healthy control subjects.

Background: Impairments in working memory(WM) have been well-documented in people with schizophrenia(PSZ). However, these quantitative WM impairments can often be explained by nonspecific factors, such as impaired goal maintenance. Here, we used a spatial orientation delayed-response task to explore a qualitative difference in WM dynamics between PSZ and healthy control subjects(HCS). Specifically, we took advantage of the discovery that WM representations may drift either toward or away from previous-trial targets(serial dependence). We tested the hypothesis that WM representations drift toward the previous-trial target in HCS but away from the previous-trial target in PSZ. Methods: We assessed serial dependence in PSZ(N=31) and HCS(N=25), using orientation as the to-be-remembered feature and memory delays from 0 to 8s. Participants were asked to remember the orientation of a teardrop-shaped object and reproduce the orientation after a varying delay period. Results: Consistent with prior studies, we found that current-trial memory representations were less precise in PSZ than in HCS. We also found that WM for the current-trial orientation drifted toward the previous-trial orientation in HCS(representational attraction) but drifted away from the previous-trial orientation in PSZ(representational repulsion). Conclusions: These results demonstrate a qualitative difference in WM dynamics between PSZ and HCS that cannot easily be explained by nuisance factors such as reduced effort. Most computational neuroscience models also fail to explain these results, because they maintain information solely by means of sustained neural firing, which does not extend across trials. The results suggest a fundamental difference between PSZ and HCS in longer-term memory mechanisms that persist across trials, such as short-term potentiation and neuronal adaptation.

Perception of opposite-direction motion in random dot kinematograms.

Computational models of motion perception suggest that the perceived direction of weak motion signals may sometimes be directly opposite to the true stimulus motion direction. However, this possibility cannot be assessed by using standard 2AFC motion discrimination paradigms because two opposite directions of motion were used in most studies (e.g., leftward vs. rightward). We were able to obtain robust evidence of opposite-direction motion reports by using a random-dot-kinematogram (RDK) paradigm in which the motion direction varied over 360° and observers were asked to estimate the exact motion direction. These opposite-direction motion reports were replicable across multiple display types and feedback conditions, and observers had greater confidence in their opposite-direction responses than in true guess responses. When we fed RDKs into a computational model of motion processing, we found that the model estimated substantial motion activity in the direction opposite to the coherent stimulus direction, even though no such motion was objectively present in the stimuli, suggesting that the opposite-direction motion perception may be a consequence of the properties of motion-selective neurons in visual cortex. Together, these results demonstrate that the known properties of the visual system may lead to reports of motion that are directly opposite to the true direction.

Working Memory Content Is Distorted by Its Use in Perceptual Comparisons.

Visual information around us is rarely static. To perform a task in such a dynamic environment, we often have to compare current visual input with our working memory (WM) representation of the immediate past. However, little is known about what happens to a WM representation when it is compared with perceptual input. To test this, we asked young adults (N = 170 total in three experiments) to compare a new visual input with a WM representation prior to reporting the WM representation. We found that the perceptual comparison biased the WM report, especially when the input was subjectively similar to the WM representation. Furthermore, using computational modeling and individual-differences analyses, we found that this similarity-induced memory bias was driven by representational integration, rather than incidental confusion, between the WM representation and subjectively similar input. Together, our findings highlight a novel source of WM distortion and suggest a general mechanism that determines how WM interacts with new visual input.

Association Between Failures in Perceptual Updating and the Severity of Psychosis in Schizophrenia.

Importance: Recent accounts suggest that delusions and hallucinations may result from alterations in how prior knowledge is integrated with new information, but experimental evidence supporting this idea has been complex and inconsistent. Evidence from a simpler perceptual task would make clear whether psychotic symptoms are associated with overreliance on prior information and impaired updating. Objective: To investigate whether individuals with schizophrenia or schizoaffective disorder (PSZ) and healthy control individuals (HCs) differ in the ability to update their beliefs based on evidence in a relatively simple perceptual paradigm. Design, Setting, and Participants: This case-control study included individuals who met DSM-IV criteria for PSZ and matched HC participants in 2 independent samples. The PSZ group was recruited from the Maryland Psychiatric Research Center, Yale University, and community clinics, and the HC group was recruited from the community. To test perceptual updating, a random dot kinematogram paradigm was implemented in which dots moving coherently in a single direction were mixed with randomly moving dots. On 50% of trials, the direction of coherent motion changed by 90° midway through the trial. Participants were asked to report the direction perceived at the end of the trial. The Peters Delusions Inventory and Brief Psychiatric Rating Scale (BPRS) were used to quantify the severity of positive symptoms. Data were collected from September 2018 to March 2020 and were analyzed from approximately March 2020 to March 2021. Main Outcomes and Measures: Critical measures included the proportion of responses centered around the initial direction vs the subsequent changed direction and the overall precision of motion perception and reaction times. Results: A total of 48 participants were included in the PSZ group (31 [65%] male; mean [SD] age, 36.56 [9.76] years) and 36 in the HC group (22 [61%] male; mean [SD] age, 35.67 [10.74] years) in the original sample. An independent replication sample included 42 participants in the PSZ group (29 [69%] male; mean [SD] age, 33.98 [11.03] years) and 34 in the HC group (20 [59%] male; mean [SD] age, 34.29 [10.44] years). In line with previous research, patients with PSZ were less precise and had slower reaction times overall. The key finding was that patients with PSZ were significantly more likely (original sample: mean, 27.88 [95% CI, 24.19-31.57]; replication sample: mean, 26.70 [95% CI, 23.53-29.87]) than HC participants (original sample: mean, 18.86 [95% CI, 16.56-21.16]; replication sample: mean, 15.67 [95% CI, 12.61-18.73]) to report the initial motion direction rather than the final one. Moreover, the tendency to report the direction of initial motion correlated with the degree of conviction on the Peters Delusions Inventory (original sample: r = 0.32 [P = .05]; replication sample: r = 0.30 [P = .05]) and the Brief Psychiatric Rating Scale Reality Distortion score (original sample: r = 0.55 [P = .001]; replication sample: r = 0.35 [P = .03]) and severity of hallucinations (original sample: r = 0.39 [P = .02]; replication sample: r = 0.30 [P = .05]). Conclusions and Relevance: The findings of this case-control study suggest that the severity of psychotic symptoms is associated with a tendency to overweight initial information over incoming sensory evidence. These results are consistent with predictive coding accounts of the origins of positive symptoms and suggest that deficits in very elementary perceptual updating may be a critical mechanism in psychosis.

Breaking the cardinal rule: The impact of interitem interaction and attentional priority on the cardinal biases in orientation working memory.

Although it is not typically assumed in influential models of visual working memory (WM), representations in WM are systematically biased by multiple factors. Orientation representations are biased away from the cardinal axis (i.e., cardinal bias) and they are biased away from or toward the other orientation simultaneously held in WM (i.e., interitem interaction). The present study investigated the extent to which these two bias mechanisms interact in WM. In Experiment 1, participants remembered two sequentially presented orientations and reproduced both orientations after a short delay. Cardinal biases were assessed separately for the trials where the two mechanisms produce biases in the same direction (i.e., congruent trials) and the trials where they produce biases in the opposite direction (i.e., incongruent trials). Whereas congruent trials exhibited a typical cardinal bias, incongruent trials exhibited no cardinal bias, demonstrating that the cardinal bias was canceled out by the interitem interaction. Follow-up experiments extended these results by manipulating attentional priority for the two orientations by means of precue (Experiment 2) and postcue (Experiment 3). In both experiments, attentionally prioritized items exhibited a typical cardinal bias irrespective of the congruency whereas attentionally unprioritized items exhibited a reversal of the cardinal bias in the incongruent trials, demonstrating that selective attention modulates the influence of the interitem interaction. Together, these results suggest that WM leverages information about specific stimuli and their relationship to support a given behavioral goal.

The Time Course of Face Representations during Perception and Working Memory Maintenance.

Successful social communication requires accurate perception and maintenance of invariant (face identity) and variant (facial expression) aspects of faces. While numerous studies investigated how face identity and expression information is extracted from faces during perception, less is known about the temporal aspects of the face information during perception and working memory (WM) maintenance. To investigate how face identity and expression information evolve over time, I recorded electroencephalography (EEG) while participants were performing a face WM task where they remembered a face image and reported either the identity or the expression of the face image after a short delay. Using multivariate event-related potential (ERP) decoding analyses, I found that the two types of information exhibited dissociable temporal dynamics: Although face identity was decoded better than facial expression during perception, facial expression was decoded better than face identity during WM maintenance. Follow-up analyses suggested that this temporal dissociation was driven by differential maintenance mechanisms: Face identity information was maintained in a more "activity-silent" manner compared to facial expression information, presumably because invariant face information does not need to be actively tracked in the task. Together, these results provide important insights into the temporal evolution of face information during perception and WM maintenance.

Neural evidence for categorical biases in location and orientation representations in a working memory task.

Previous research demonstrated that visual representations in working memory exhibit biases with respect to the categorical structure of the stimulus space. However, a majority of those studies used behavioral measures of working memory, and it is not clear whether the working memory representations per se are influenced by the categorical structure or whether the biases arise in decision or response processes during the report. Here, I applied a multivariate decoding technique to EEG data collected during working memory tasks to determine whether neural activity associated with the representations in working memory is categorically biased prior to the report. I found that the decoding of spatial working memory was biased away from the nearest cardinal location, consistent with the biases observed in the behavioral responses. In a follow-up experiment which was designed to prevent the use of a response preparation strategy, I found that the decoding still exhibited categorical biases. Together, these results provide neural evidence that working memory representations themselves are categorically biased, imposing important constraints on the models of working memory representations.

Increased repulsion of working memory representations in schizophrenia.

Computational neuroscience models propose that working memory (WM) involves recurrent excitatory feedback loops that maintain firing over time along with lateral inhibition that prevents the spreading of activity to other feature values. In behavioral paradigms, this lateral inhibition appears to cause a repulsion of WM representations away from each other and from other strong sources of input. Recent computational models of schizophrenia have proposed that reduction in the strength of inhibition relative to strength of excitation may underlie impaired cognition, and this leads to the prediction that repulsion effects should be reduced in people with schizophrenia spectrum disorders (PSZ) relative to healthy control subjects (HCS). We tested this hypothesis in 2 experiments measuring WM repulsion effects. In Experiment 1, 45 PSZ and 32 HCS remembered the location of a single object relative to a centrally presented visual landmark and reported this location after a short delay. The reported location was repelled away from the landmark in both groups, but this repulsion effect was increased rather than decreased in PSZ relative to HCS. In Experiment 2, 41 PSZ and 34 HCS remembered 2 sequentially presented orientations and reported each orientation after a short delay. The reported orientations were biased away from each other in both groups, and this repulsion effect was again more pronounced in PSZ than in HCS. Contrary to the widespread hypothesis of reduced inhibition in schizophrenia, we provide robust evidence from 2 experiments showing that the behavioral performance of PSZ exhibited an exaggeration rather than a reduction of competitive inhibition. (PsycInfo Database Record (c) 2020 APA, all rights reserved).

Cortical hyperactivation at low working memory load: A primary processing abnormality in people with schizophrenia?

A frequent finding when studying substrates of working memory (WM) deficits in people with schizophrenia (PSZ) is task-induced hyperactivation relative to healthy control subjects (HCS) when WM load is low. Hyperactivation accompanying similar performance is commonly attributed to cognitive deficits rendering relatively easy operations more resource-consuming. To test if hyperactivation at low load really is secondary to cognitive impairment in PSZ, we re-analyzed functional MRI data showing left posterior parietal cortex (PPC) hyperactivation in PSZ when holding a single color-item in WM. In subgroups matched for the number of items successfully stored in WM (K) by excluding the highest-performing HCS and lowest-performing PSZ, performance was almost identical across all set sizes (1-7). While BOLD activation at the larger set sizes did not differ between groups, PSZ still robustly hyperactivated left PPC when a single item had to be maintained. The same pattern was observed in subgroups matched for model-based estimates of WM capacity or attentional lapse rate. Given that in the K-matched subsamples PSZ performed as well as HCS even in the most challenging load conditions and that no BOLD signal difference was seen at high loads, it is implausible that PSZ over-recruited WM-related neural structures because they were more challenged by maintaining a single item in WM. Instead, the findings are consistent with a primary schizophrenia-related processing abnormality as proposed by the hyperfocusing hypothesis, which suggests that an abnormally narrow but intense focusing of processing resources is central to many aspects of impaired cognition in PSZ.

Assessing the information content of ERP signals in schizophrenia using multivariate decoding methods.

Multivariate pattern classification (decoding) methods are commonly employed to study mechanisms of neurocognitive processing in typical individuals, where they can be used to quantify the information that is present in single-participant neural signals. These decoding methods are also potentially valuable in determining how the representation of information differs between psychiatric and non-psychiatric populations. Here, we examined ERPs from people with schizophrenia (PSZ) and healthy control subjects (HCS) in a working memory task that involved remembering 1, 3, or 5 items from one side of the display and ignoring the other side. We used the spatial pattern of ERPs to decode which side of the display was being held in working memory. One might expect that decoding accuracy would be inevitably lower in PSZ as a result of increased noise (i.e., greater trial-to-trial variability). However, we found that decoding accuracy was greater in PSZ than in HCS at memory load 1, consistent with previous research in which memory-related ERP signals were larger in PSZ than in HCS at memory load 1. We also observed that decoding accuracy was strongly related to the ratio of the memory-related ERP activity and the noise level. In addition, we found similar noise levels in PSZ and HCS, counter to the expectation that PSZ would exhibit greater trial-to-trial variability. Together, these results demonstrate that multivariate decoding methods can be validly applied at the individual-participant level to understand the nature of impaired cognitive function in a psychiatric population.

Serial dependence in vision: Merely encoding the previous-trial target is not enough.

The reported perception of a visual stimulus on one trial can be biased by the stimulus that was presented on the previous trial. In the present study we asked whether encoding the previous-trial stimulus is sufficient to produce this serial dependence effect, or whether the effect also depends on postencoding processes. To distinguish between these possibilities, we designed a task in which participants reported either the color or the direction of a set of colored moving dots on each trial. The to-be-reported dimension was indicated by a postcue after stimulus offset, so participants were required to encode both features of every stimulus. We assessed serial dependence for motion perception as a function of which feature dimension had been reported on the previous trial. In Experiment 1, we found a serial dependence effect for motion only when participants had reported the direction of motion on the previous trial, and not when they had encoded the direction of motion but reported the color of the stimulus. Experiment 2 confirmed that this pattern of results was not driven by the difficulty of the color task. When we used the same response modality for both motion and color reports in Experiment 3, we found significant serial dependence effects following both color-report and motion-report trials, but the effect was significantly weaker following color-report trials. Together, these findings indicate that postperceptual processes play a critical role in serial dependence and that the mere encoding of the previous-trial target is not sufficient to produce the serial dependence effect.

Reactivation of Previous Experiences in a Working Memory Task.

Recent experiences influence the processing of new information even when those experiences are irrelevant to the current task. Does this reflect the indirect effects of a passively maintained representation of the previous experience, or is this representation reactivated when a new event occurs? To answer this question, we attempted to decode the orientation of the stimulus on the previous trial from the electroencephalogram on the current trial in a working memory task. Behavioral data confirmed that the previous-trial stimulus orientation influenced the reported orientation on the current trial, even though the previous-trial orientation was now task irrelevant. In two independent experiments, we found that the previous-trial orientation could be decoded from the current-trial electroencephalogram, indicating that the current-trial stimulus reactivated or boosted the representation of the previous-trial orientation. These results suggest that the effects of recent experiences on behavior are driven, in part, by a reactivation of those experiences and not solely by the indirect effects of passive memory traces.

Decoding motion direction using the topography of sustained ERPs and alpha oscillations.

The present study sought to determine whether scalp electroencephalogram (EEG) signals contain decodable information about the direction of motion in random dot kinematograms (RDKs), in which the motion information is spatially distributed and mixed with random noise. Any direction of motion from 0 to 360° was possible, and observers reported the precise direction of motion at the end of a 1500-ms stimulus display. We decoded the direction of motion separately during the motion period (during which motion information was being accumulated) and the report period (during which a shift of attention was necessary to make a fine-tuned direction report). Machine learning was used to decode the precise direction of motion (within ±11.25°) from the scalp distribution of either alpha-band EEG activity or sustained event-related potentials (ERPs). We found that ERP-based decoding was above chance (1/16) during both the stimulus and the report periods, whereas alpha-based decoding was above chance only during the report period. Thus, sustained ERPs contain information about spatially distributed direction-of-motion, providing a new method for observing the accumulation of sensory information with high temporal resolution. By contrast, the scalp topography of alpha-band EEG activity appeared to mainly reflect spatially focused attentional processes rather than sensory information.

What happens to an individual visual working memory representation when it is interrupted?

This study tested the hypothesis that even the simplest cognitive tasks require the storage of information in working memory (WM), distorting any information that was previously stored in WM. Experiment 1 tested this hypothesis by requiring observers to perform a simple letter discrimination task while they were holding a single orientation in WM. We predicted that performing the task on the interposed letter stimulus would cause the orientation memory to become less precise and more categorical compared to when the letter was absent or when it was present but could be ignored. This prediction was confirmed. Experiment 2 tested the modality specificity of this effect by replacing the visual letter discrimination task with an auditory pitch discrimination task. Unlike the interposed visual stimulus, the interposed auditory stimulus produced little or no disruption of WM, consistent with the use of modality-specific representations. Thus, performing a simple visual discrimination task, but not a simple auditory discrimination task, distorts information about a single feature being maintained in visual WM. We suggest that the interposed task eliminates information stored within the focus of attention, leaving behind a WM representation outside the focus of attention that is relatively imprecise and categorical.

Dissociable Decoding of Spatial Attention and Working Memory from EEG Oscillations and Sustained Potentials.

In human scalp EEG recordings, both sustained potentials and alpha-band oscillations are present during the delay period of working memory tasks and may therefore reflect the representation of information in working memory. However, these signals may instead reflect support mechanisms rather than the actual contents of memory. In particular, alpha-band oscillations have been tightly tied to spatial attention and may not reflect location-independent memory representations per se. To determine how sustained and oscillating EEG signals are related to attention and working memory, we attempted to decode which of 16 orientations was being held in working memory by human observers (both women and men). We found that sustained EEG activity could be used to decode the remembered orientation of a stimulus, even when the orientation of the stimulus varied independently of its location. Alpha-band oscillations also carried clear information about the location of the stimulus, but they provided little or no information about orientation independently of location. Thus, sustained potentials contain information about the object properties being maintained in working memory, consistent with previous evidence of a tight link between these potentials and working memory capacity. In contrast, alpha-band oscillations primarily carry location information, consistent with their link to spatial attention.SIGNIFICANCE STATEMENT Working memory plays a key role in cognition, and working memory is impaired in several neurological and psychiatric disorders. Previous research has suggested that human scalp EEG recordings contain signals that reflect the neural representation of information in working memory. However, to conclude that a neural signal actually represents the object being remembered, it is necessary to show that the signal contains fine-grained information about that object. Here, we show that sustained voltages in human EEG recordings contain fine-grained information about the orientation of an object being held in memory, consistent with a memory storage signal.