Adaptive forgetting speed in working memory.

Working memory is known to be capacity-limited and is therefore selective not only for what it encodes but also what it forgets. Explicit forgetting cues can be used effectively to free up capacity, but it is not clear how working memory adaptively forgets in the absence of explicit cues. An important implicit cue that may tune forgetting in working memory is the passage of time. When information becomes irrelevant more quickly, working memory should also forget information more quickly. In three delayed-estimation experiments, we systematically manipulated how probing probability changed as time passed on after encoding an item (i.e., the "probing hazard"). In some blocks, probing hazard decreased after encoding an item, requiring participants to only briefly retain the memory item. In other blocks, the probing hazard increased or stayed flat, as the retention interval was lengthened. In line with our hypothesis, we found that participants adapted their forgetting rate to the probing dynamics of the working memory task. When the memory item quickly became irrelevant ("decreasing" probing hazard), forgetting rate was higher than in blocks where probing hazard increased or stayed flat. The time course of these adaptations in forgetting implies a fast and flexible mechanism. Interestingly, participants could not explicitly report the order of conditions, suggesting forgetting is implicitly sped up. These findings suggest that implicit adaptations to the temporal structure of our environment tune forgetting speed in working memory, possibly contributing to the flexible allocation of limited working memory resources.

Maintenance of color memoranda in activity-quiescent working memory states: Evidence from impulse perturbation.

In the present study, we used an impulse perturbation method to probe working memory maintenance of colors in neurally active and activity-quiescent states, focusing on a set of pre-registered analyses. We analyzed the electroencephalograph (EEG) data of 30 participants who completed a delayed match-to-sample working memory task, in which one of the two items that were presented was retro-cued as task relevant. The analyses revealed that both cued and uncued colors were decodable from impulse-evoked activity, the latter in contrast to previous reports of working memory for orientation gratings. Decoding of colors from oscillations in the alpha band showed that cued items could be decoded therein whereas uncued items could not. Overall, the outcomes suggest that subtle differences exist between the representation of colors, and that of stimuli with spatial properties, but the present results also demonstrate that regardless of their specific neural state, both are accessible through visual impulse perturbation.

Dissociable event-related potential modulations of intrinsic and extrinsic factors in temporal integration.

We investigated visual temporal integration, by which multiple stimuli appearing in rapid succession are perceived as a single event. Temporal integration not only depends intrinsically on the passage of time but also, extrinsically, on the number and distribution of successive stimuli that are presented across that time interval. Here, we used a missing element task to investigate intrinsic and extrinsic factors in temporal integration, by manipulating stimulus duration and number, respectively. We found that both contributed interactively to integration performance and that varying the information rate over time did not further modulate this pattern. Intrinsic and extrinsic factors had dissociable effects on the N1, N2, N2pc, and P3 components of the event-related potential, implicating unique contributions to perceptual discrimination, spatio-temporal grouping, attention, and response decision-making. Stimulus number-induced effects on the event-related potential also generally arose later than those of stimulus duration. The latter already modulated the amplitude of the N1 and the early phase of the N2pc, while the former did not. The collective results suggest that while both intrinsic and extrinsic factors drive temporal integration, they do so in different ways. This difference during integration may eventually be reflected in the way in which we perceive longer, episodic events.

Attentional blur and blink: Effects of adaptive attentional scaling on visual awareness.

Attentional scaling is a crucial mechanism that enables us to flexibly allocate our attention to larger or smaller regions in the visual field. Although previous studies have demonstrated the critical role of attentional scaling in visual processing, its impact on modulating visual awareness is not yet fully understood. This study investigates the adaptive control of attentional scaling and its influence on visual awareness in an attentional blink paradigm. Participants were required to attend to the first target's location, which was manipulated either session-wise, trial-wise, or such that it could be learned across a block of trials. Discrete, all-or-none, awareness was expected when attention was allocated to a narrow area, while gradual awareness was expected when attention was allocated to a larger area. We used mixture modeling to assess second target awareness across these different attentional scales. The results revealed that participants could adaptively control their attentional scale both across stable sessions, and through (implicit) statistical learning in blocks of successive trials. This produced gradual perceptual awareness when the participants adopted a broad attentional scale, causing an attentional "blur". However, trial-wise cues did not allow for attentional scaling, resulting in more discrete target perception overall, and an attentional "blink". We conclude that the attentional scale is to some extent under adaptive control during the attentional blink/blur, where it can produce qualitatively different modes of perceptual awareness.

Adaptive Encoding Speed in Working Memory.

Humans can adapt when complex patterns unfold at a faster or slower pace, for instance when remembering a grocery list that is dictated at an increasingly fast rate. Integrating information over such timescales crucially depends on working memory, but although recent findings have shown that working memory capacity can be flexibly adapted, such adaptations have not yet been demonstrated for encoding speed. In a series of experiments, we found that young adults encoded at a faster rate when they were adapted to overall and recent stimulus duration. Interestingly, our participants were unable to use explicit cues to speed up encoding, even though these cues were objectively more informative than statistical information. Our findings suggest that adaptive tuning of encoding speed in working memory is a fundamental but largely implicit mechanism underlying our ability to keep up with the pace of our surroundings.

Impulse perturbation reveals cross-modal access to sensory working memory through learned associations.

We investigated if learned associations between visual and auditory stimuli can afford full cross-modal access to working memory. Previous research using the impulse perturbation technique has shown that cross-modal access to working memory is one-sided; visual impulses reveal both auditory and visual memoranda, but auditory impulses do not seem to reveal visual memoranda (Wolff et al., 2020b). Our participants first learned to associate six auditory pure tones with six visual orientation gratings. Next, a delayed match-to-sample task for the orientations was completed, while EEG was recorded. Orientation memories were recalled either via their learned auditory counterpart, or were visually presented. We then decoded the orientation memories from the EEG responses to both auditory and visual impulses presented during the memory delay. Working memory content could always be decoded from visual impulses. Importantly, through recall of the learned associations, the auditory impulse also evoked a decodable response from the visual WM network, providing evidence for full cross-modal access. We also observed that after a brief initial dynamic period, the representational codes of the memory items generalized across time, as well as between perceptual maintenance and long-term recall conditions. Our results thus demonstrate that accessing learned associations in long-term memory provides a cross-modal pathway to working memory that seems to be based on a common coding scheme.

Effects of gamma-aminobutyric acid on working memory and attention: A randomized, double-blinded, placebo-controlled, crossover trial.

BACKGROUND: γ-Aminobutyric acid (GABA) is a primary inhibitory neurotransmitter that plays a significant role in the central nervous system. Studies on both animals and humans show that GABA has the pharmacological potential for reducing the impact of cognitive disorders, as well as enhancing cognitive functions and mood. However, its specific effects on human attention and working memory have not yet been extensively studied. AIMS: In this randomized, double-blind, placebo-controlled, and crossover trial, we aimed to test whether the administration of 800 mg GABA, dissolved in a drink, acutely affected visual working memory (VWM) maintenance, as well as temporal and spatial attention in healthy adults. METHODS: The participants were 32 young adults (16 females and 16 males). Working memory recall precision, spatial attention and temporal attention were measured by a delayed match-to-sample task, a visual search (VS) task and a speeded rapid serial visual presentation task, respectively. Participants completed two experimental sessions (GABA and Placebo) in randomized and counterbalanced order. In each session, 45 min after administration of the drink, they completed all three aforementioned cognitive tasks. RESULTS: Linear mixed model analysis results showed that GABA increased VS time, compared to the placebo, but did not affect VS accuracy, temporal attention, nor VWM precision. CONCLUSIONS: The results suggest that GABA increases VS time but does not affect temporal attention and memory, and that previously reported effects on cognition might rely on other functions.

Object-based visual working memory: an object benefit for equidistant memory items presented within simple contours.

Previous research has shown that more information can be stored in visual working memory (VWM) when multiple items belong to the same object. Here, in four experiments, we investigated the object effect on memory for spatially equidistant features by manipulating simple, task-irrelevant contours that combined these features. In Experiments 1, 3, and, 4, three grating orientations, and in Experiment 2, one color and two orientations, were presented simultaneously to be memorized. Mixture modeling was applied to estimate both the precision and the guess rates of recall errors. Overall results showed that two target features were remembered more accurately when both were part of the same object. Further analysis showed that the probability of recall increased in particular when both features were extracted from the same object. In Experiment 2, we found that the object effect was greater for features from orthogonal dimensions, but this came at the cost of lower memory precision. In Experiment 3, when we kept the locations of the features perfectly consistent over trials so that the participants could attend to these locations rather than the contour, we still found object benefits. Finally, in Experiment 4 when we manipulated the temporal order of the object and the memory features presentations, it was confirmed that the object benefit is unlikely to stem from the strategical usage of object information. These results suggested that the object benefit arises automatically, likely at an early perceptual level.

Concealed identity information detection with pupillometry in rapid serial visual presentation.

The concealed information test (CIT) relies on bodily reactions to stimuli that are hidden in mind. However, people can use countermeasures, such as purposely focusing on irrelevant things, to confound the CIT. A new method designed to prevent countermeasures uses rapid serial visual presentation (RSVP) to present stimuli on the fringe of awareness. Previous studies that used RSVP in combination with electroencephalography (EEG) showed that participants exhibit a clear reaction to their real first name, even when they try to prevent such a reaction (i.e., when their name is concealed information). Because EEG is not easily applicable outside the laboratory, we investigated here whether pupil size, which is easier to measure, can also be used to detect concealed identity information. In our first study, participants adopted a fake name, and searched for this name in an RSVP task, while their pupil sizes were recorded. Apart from this fake name, their real name and a control name also appeared in the task. We found pupil dilation in response to the task-irrelevant real name, as compared to control names. However, while most participants showed this effect qualitatively, it was not statistically significant for most participants individually. In a second study, we preregistered the proof-of-concept methodology and replicated the original findings. Taken together, our results show that the current RSVP task with pupillometry can detect concealed identity information at a group level. Further development of the method is needed to create a valid and reliable concealed identity information detector at the individual level.

Acute effects of cocoa flavanols on visual working memory: maintenance and updating.

BACKGROUND: Consumption of cocoa flavanols may have acute physiological effects on the brain due to their ability to activate nitric oxide synthesis. Nitric oxide mediates vasodilation, which increases cerebral blood flow, and can also act as a neurotransmitter. OBJECTIVES: This study aimed to examine whether cocoa flavanols have an acute influence on visual working memory (WM). METHODS: Two separate randomised, double-blind, placebo-controlled, counterbalanced crossover experiments were conducted on normal healthy young adult volunteers (NExp1 = 48 and NExp2 = 32, gender-balanced). In these experiments, 415 mg of cocoa flavanols were administered to test their acute effects on visual working memory. In the first experiment, memory recall precision was measured in a task that required only passive maintenance of grating orientations in WM. In the second experiment, recall was measured after active updating (mental rotation) of WM contents. Habitual daily flavanols intake, body mass index, and gender were also considered in the analysis. RESULTS: The results suggested that neither passive maintenance in visual WM nor active updating of WM were acutely enhanced by consumption of cocoa flavanols. Exploratory analyses with covariates (body mass index and daily flavanols intake), and the between-subjects factor of gender also showed no evidence for effects of cocoa flavanols, neither in terms of reaction time, nor accuracy. CONCLUSIONS: Overall, cocoa flavanols did not improve visual working memory recall performance during maintenance, nor did it improve recall accuracy after memory updating.

Two faces of perceptual awareness during the attentional blink: Gradual and discrete.

In a series of experiments, the nature of perceptual awareness during the attentional blink was investigated. Previous work has considered the attentional blink as a discrete, all-or-none phenomenon, indicative of general access to conscious awareness. Using continuous report measures in combination with mixture modeling, the outcomes showed that perceptual awareness during the attentional blink can be a gradual phenomenon. Awareness was not exclusively discrete, but also exhibited a gradual characteristic whenever the spatial extent of attention induced by the first target spanned more than a single location. Under these circumstances, mental representations of blinked targets were impoverished, but did approach the actual identities of the targets. Conversely, when the focus of attention covered only a single location, there was no evidence for any partial knowledge of blinked targets. These two different faces of awareness during the attentional blink challenge current theories of both awareness and temporal attention, which cannot explain the existence of gradual awareness of targets during the attentional blink. To account for the current outcomes, an adaptive gating model is proposed that casts awareness on a continuum between gradual and discrete, rather than as being of either single kind. (PsycInfo Database Record (c) 2022 APA, all rights reserved).

A common dynamic prior for time in duration discrimination.

Estimation of time depends heavily on both global and local statistical context. Durations that are short relative to the global distribution are systematically overestimated; durations that are locally preceded by long durations are also overestimated. Context effects are prominent in duration discrimination tasks, where a standard duration and a comparison duration are presented on each trial. In this study, we compare and test two models that posit a dynamically updating internal reference that biases time estimation on global and local scales in duration discrimination tasks. The internal reference model suggests that the internal reference operates during postperceptual stages and only interacts with the first presented duration. In contrast, a Bayesian account of time estimation implies that any perceived duration updates the internal reference and therefore interacts with both the first and second presented duration. We implemented both models and tested their predictions in a duration discrimination task where the standard duration varied from trial to trial. Our results are in line with a Bayesian perspective on time estimation. First, the standard systematically biased estimation of the comparison, such that shorter standards increased the likelihood of reporting that the comparison was shorter. Second, both the previous standard and comparison systematically biased time estimation of subsequent trials in the same direction. Third, more precise observers showed smaller biases. In sum, our findings suggest a common dynamic prior for time that is updated by each perceived duration and where the relative weighting of old and new observations is determined by their relative precision.

A functional spiking-neuron model of activity-silent working memory in humans based on calcium-mediated short-term synaptic plasticity.

In this paper, we present a functional spiking-neuron model of human working memory (WM). This model combines neural firing for encoding of information with activity-silent maintenance. While it used to be widely assumed that information in WM is maintained through persistent recurrent activity, recent studies have shown that information can be maintained without persistent firing; instead, information can be stored in activity-silent states. A candidate mechanism underlying this type of storage is short-term synaptic plasticity (STSP), by which the strength of connections between neurons rapidly changes to encode new information. To demonstrate that STSP can lead to functional behavior, we integrated STSP by means of calcium-mediated synaptic facilitation in a large-scale spiking-neuron model and added a decision mechanism. The model was used to simulate a recent study that measured behavior and EEG activity of participants in three delayed-response tasks. In these tasks, one or two visual gratings had to be maintained in WM, and compared to subsequent probes. The original study demonstrated that WM contents and its priority status could be decoded from neural activity elicited by a task-irrelevant stimulus displayed during the activity-silent maintenance period. In support of our model, we show that it can perform these tasks, and that both its behavior as well as its neural representations are in agreement with the human data. We conclude that information in WM can be effectively maintained in activity-silent states by means of calcium-mediated STSP.

Adaptive event integration in the missing element task.

Evidence for adaptive event integration has previously been provided using the Rapid Serial Visual Presentation (RSVP) task. However, it is not straightforward to generalize this finding to other types of tasks that measure temporal integration, because integration in such tasks is known to vary, depending on the method that is used. This variability has been seen as an indication that integration may result from more than a single type of perceptual persistence, and that different integration tasks may not tap into same type of persistence. Therefore, we investigated whether adaptive control of integration in the RSVP task can be replicated using another technique for measuring temporal integration, which may rely more on low-level mechanisms, namely the dot-array integration or Missing Element Task (MET). As in the RSVP studies, stimulus speed expectancy was presently manipulated. The results indicated that integration performance in the MET was not subject to adaptive control. We argue that this discrepancy with previous RSVP studies can most likely be attributed to a specific difference in the type of persistence underlying task performance. Temporal integration in the MET might rely mostly on visible persistence, while for the RSVP task integration relies more on informational persistence. The present findings suggest that, contrary to informational persistence, visible persistence may not be susceptible to adaptive control.

Drifting codes within a stable coding scheme for working memory.

Working memory (WM) is important to maintain information over short time periods to provide some stability in a constantly changing environment. However, brain activity is inherently dynamic, raising a challenge for maintaining stable mental states. To investigate the relationship between WM stability and neural dynamics, we used electroencephalography to measure the neural response to impulse stimuli during a WM delay. Multivariate pattern analysis revealed representations were both stable and dynamic: there was a clear difference in neural states between time-specific impulse responses, reflecting dynamic changes, yet the coding scheme for memorised orientations was stable. This suggests that a stable subcomponent in WM enables stable maintenance within a dynamic system. A stable coding scheme simplifies readout for WM-guided behaviour, whereas the low-dimensional dynamic component could provide additional temporal information. Despite having a stable subspace, WM is clearly not perfect-memory performance still degrades over time. Indeed, we find that even within the stable coding scheme, memories drift during maintenance. When averaged across trials, such drift contributes to the width of the error distribution.

Unimodal and Bimodal Access to Sensory Working Memories by Auditory and Visual Impulses.

It is unclear to what extent sensory processing areas are involved in the maintenance of sensory information in working memory (WM). Previous studies have thus far relied on finding neural activity in the corresponding sensory cortices, neglecting potential activity-silent mechanisms, such as connectivity-dependent encoding. It has recently been found that visual stimulation during visual WM maintenance reveals WM-dependent changes through a bottom-up neural response. Here, we test whether this impulse response is uniquely visual and sensory-specific. Human participants (both sexes) completed visual and auditory WM tasks while electroencephalography was recorded. During the maintenance period, the WM network was perturbed serially with fixed and task-neutral auditory and visual stimuli. We show that a neutral auditory impulse-stimulus presented during the maintenance of a pure tone resulted in a WM-dependent neural response, providing evidence for the auditory counterpart to the visual WM findings reported previously. Interestingly, visual stimulation also resulted in an auditory WM-dependent impulse response, implicating the visual cortex in the maintenance of auditory information, either directly or indirectly, as a pathway to the neural auditory WM representations elsewhere. In contrast, during visual WM maintenance, only the impulse response to visual stimulation was content-specific, suggesting that visual information is maintained in a sensory-specific neural network, separated from auditory processing areas.SIGNIFICANCE STATEMENT Working memory is a crucial component of intelligent, adaptive behavior. Our understanding of the neural mechanisms that support it has recently shifted: rather than being dependent on an unbroken chain of neural activity, working memory may rely on transient changes in neuronal connectivity, which can be maintained efficiently in activity-silent brain states. Previous work using a visual impulse stimulus to perturb the memory network has implicated such silent states in the retention of line orientations in visual working memory. Here, we show that auditory working memory similarly retains auditory information. We also observed a sensory-specific impulse response in visual working memory, while auditory memory responded bimodally to both visual and auditory impulses, possibly reflecting visual dominance of working memory.

Between one event and two: The locus of the effect of stimulus contrast on temporal integration.

The effects of relative stimulus contrast on temporal integration were investigated in a missing element task. Integration frequency was strongly modulated when the contrast of either the first or the second stimulus display was reduced. When the contrast of the first display was low, integration was enhanced, while it was reduced when the contrast of the second display was low. To reveal the processing phases implicated in these modulations of integration, the amplitude of ERP components was examined. N1 component amplitude was increased when the second display was low contrast, matching a full contrast condition. At the N2pc component, amplitude was strongly suppressed only in the former condition. P3 amplitude was also lowest when contrast on the second display was low but with successively increasing amplitudes observed for the other conditions, largely matching the pattern observed in behavioral performance. Taken together, contrast effects on temporal integration seem to originate from increased discriminative processing of the first stimulus display in particular (N1), which is consequently followed by an impairment in attentional processing (N2pc) and working memory consolidation (P3) of the missing element location.

Temporal integration and attentional selection of color and contrast target pairs in rapid serial visual presentation.

Performance in a dual target rapid serial visual presentation task was investigated, dependent on whether the color or the contrast of the targets was the same or different. Both identification accuracy on the second target, as a measure of temporal attention, and the frequency of temporal integration were measured. When targets had a different color (red or blue), overall identification accuracy of the second target and identification accuracy of the second target at Lag 1 were both higher than when targets had the same color. At the same time, increased temporal integration of the targets at Lag 1 was observed in the different color condition, even though actual (non-integrated) single targets never consisted of multiple colors. When the color pairs were made more similar, so that they all fell within the range of a single nominal hue (blue), these effects were not observed. Different findings were obtained when contrast was manipulated. Identification accuracy of the second target was higher in the same contrast condition than in the different contrast condition. Higher identification accuracy of both targets was furthermore observed when they were presented with high contrast, while target contrast did not influence temporal integration at all. Temporal attention and integration were thus influenced differently by target contrast pairing than by (categorical) color pairing. Categorically different color pairs, or more generally, categorical feature pairs, may thus afford a reduction in temporal competition between successive targets that eventually enhances attention and integration.

Visual and auditory temporal integration in healthy younger and older adults.

As people age, they tend to integrate successive visual stimuli over longer intervals than younger adults. It may be expected that temporal integration is affected similarly in other modalities, possibly due to general, age-related cognitive slowing of the brain. However, the previous literature does not provide convincing evidence that this is the case in audition. One hypothesis is that the primacy of time in audition attenuates the degree to which temporal integration in that modality extends over time as a function of age. We sought to settle this issue by comparing visual and auditory temporal integration in younger and older adults directly, achieved by minimizing task differences between modalities. Participants were presented with a visual or an auditory rapid serial presentation task, at 40-100 ms/item. In both tasks, two subsequent targets were to be identified. Critically, these could be perceptually integrated and reported by the participants as such, providing a direct measure of temporal integration. In both tasks, older participants integrated more than younger adults, especially when stimuli were presented across longer time intervals. This difference was more pronounced in vision and only marginally significant in audition. We conclude that temporal integration increases with age in both modalities, but that this change might be slightly less pronounced in audition.

The acute effects of cocoa flavanols on temporal and spatial attention.

In this study, we investigated how the acute physiological effects of cocoa flavanols might result in specific cognitive changes, in particular in temporal and spatial attention. To this end, we pre-registered and implemented a randomized, double-blind, placebo- and baseline-controlled crossover design. A sample of 48 university students participated in the study and each of them completed the experimental tasks in four conditions (baseline, placebo, low dose, and high-dose flavanol), administered in separate sessions with a 1-week washout interval. A rapid serial visual presentation task was used to test flavanol effects on temporal attention and integration, and a visual search task was similarly employed to investigate spatial attention. Results indicated that cocoa flavanols improved visual search efficiency, reflected by reduced reaction time. However, cocoa flavanols did not facilitate temporal attention nor integration, suggesting that flavanols may affect some aspects of attention, but not others. Potential underlying mechanisms are discussed.