Goal-Dependent Use of Temporal Regularities to Orient Attention under Spatial and Action Uncertainty.

The temporal regularities in our environments support the proactive dynamic anticipation of relevant events. In visual attention, one important outstanding question is whether temporal predictions must be linked to predictions about spatial locations or motor plans to facilitate behaviour. To test this, we developed a task for manipulating temporal expectations and task relevance of visual stimuli appearing within rapidly presented streams, while stimulus location and responding hand remained uncertain. Differently coloured stimuli appeared in one of two concurrent (left and right) streams with distinct temporal probability structures. Targets were defined by colour on a trial-by-trial basis and appeared equiprobably in either stream, requiring a localisation response. Across two experiments, participants were faster and more accurate at detecting temporally predictable targets compared to temporally unpredictable targets. We conclude that temporal expectations learned incidentally from temporal regularities can be called upon flexibly in a goal-driven manner to guide behaviour. Moreover, we show that visual temporal attention can facilitate performance in the absence of concomitant spatial or motor expectations in dynamically unfolding contexts.

A Neural Decision Signal during Internal Sampling from Working Memory in Humans.

How humans transform sensory information into decisions that steer purposeful behavior is a central question in psychology and neuroscience that is traditionally investigated during the sampling of external environmental signals. The decision-making framework of gradual information sampling toward a decision has also been proposed to apply when sampling internal sensory evidence from working memory. However, neural evidence for this proposal remains scarce. Here we show (using scalp EEG in male and female human volunteers) that sampling internal visual representations from working memory elicits a scalp EEG potential associated with gradual evidence accumulation-the central parietal positivity. Consistent with an evolving decision process, we show how this signal (1) scales with the time participants require to reach a decision about the cued memory content and (2) is amplified when having to decide among multiple contents in working memory. These results bring the electrophysiology of decision-making into the domain of working memory and suggest that variability in memory-guided behavior may be driven (at least in part) by variations in the sampling of our inner mental contents.

Shifting attention between perception and working memory.

Most everyday tasks require shifting the focus of attention between sensory signals in the external environment and internal contents in working memory. To date, shifts of attention have been investigated within each domain, but shifts between the external and internal domain remain poorly understood. We developed a combined perception and working-memory task to investigate and compare the consequences of shifting spatial attention within and between domains in the service of a common orientation-reproduction task. Participants were sequentially cued to attend to items either in working memory or to an upcoming sensory stimulation. Stay trials provided a baseline condition, while shift trials required participants to shift their attention to another item within the same or different domain. Validating our experimental approach, we found evidence that participants shifted attention effectively in either domain (Experiment 1). In addition, we observed greater costs when transitioning attention between as compared to within domains (Experiments 1, 2). Strikingly, these costs persisted even when participants were given more time to complete the attentional shift (Experiment 2). Biases in fixational gaze behaviour tracked attentional orienting in both domains, but revealed no latency or magnitude difference for within- versus between-domain shifts (Experiment 1). Collectively, the results from Experiments 1 and 2 suggest that shifting between attentional domains might be regulated by a unique control function. Our results break new ground for exploring the ubiquitous act of shifting attention between perception and working memory to guide adaptive behaviour in everyday cognition.

Time for What? Dissociating Explicit Timing Tasks through Electrophysiological Signatures.

Estimating durations between hundreds of milliseconds and seconds is essential for several daily tasks. Explicit timing tasks, which require participants to estimate durations to make a comparison (time for perception) or to reproduce them (time for action), are often used to investigate psychological and neural timing mechanisms. Recent studies have proposed that mechanisms may depend on specific task requirements. In this study, we conducted electroencephalogram (EEG) recordings on human participants as they estimated intervals in different task contexts to investigate the extent to which timing mechanisms depend on the nature of the task. We compared the neural processing of identical visual reference stimuli in two different tasks, in which stimulus durations were either perceptually compared or motorically reproduced in separate experimental blocks. Using multivariate pattern analyses, we could successfully decode the duration and the task of reference stimuli. We found evidence for both overlapping timing mechanisms across tasks as well as recruitment of task-dependent processes for comparing intervals for different purposes. Our findings suggest both core and specialized timing functions are recruited to support explicit timing tasks.

Time for Memories.

The ability to store information about the past to dynamically predict and prepare for the future is among the most fundamental tasks the brain performs. To date, the problems of understanding how the brain stores and organizes information about the past (memory) and how the brain represents and processes temporal information for adaptive behavior have generally been studied as distinct cognitive functions. This Symposium explores the inherent link between memory and temporal cognition, as well as the potential shared neural mechanisms between them. We suggest that working memory and implicit timing are interconnected and may share overlapping neural mechanisms. Additionally, we explore how temporal structure is encoded in associative and episodic memory and, conversely, the influences of episodic memory on subsequent temporal anticipation and the perception of time. We suggest that neural sequences provide a general computational motif that contributes to timing and working memory, as well as the spatiotemporal coding and recall of episodes.

Using XR (Extended Reality) for Behavioral, Clinical, and Learning Sciences Requires Updates in Infrastructure and Funding.

Extended reality (XR, including augmented and virtual reality) creates a powerful intersection between information technology and cognitive, clinical, and education sciences. XR technology has long captured the public imagination, and its development is the focus of major technology companies. This article demonstrates the potential of XR to (1) deliver behavioral insights, (2) transform clinical treatments, and (3) improve learning and education. However, without appropriate policy, funding, and infrastructural investment, many research institutions will struggle to keep pace with the advances and opportunities of XR. To realize the full potential of XR for basic and translational research, funding should incentivize (1) appropriate training, (2) open software solutions, and (3) collaborations between complementary academic and industry partners. Bolstering the XR research infrastructure with the right investments and incentives is vital for delivering on the potential for transformative discoveries, innovations, and applications.

Evidence Accumulation Rate Moderates the Relationship between Enriched Environment Exposure and Age-Related Response Speed Declines.

Older adults exposed to enriched environments (EEs) maintain relatively higher levels of cognitive function, even in the face of compromised markers of brain health. Response speed (RS) is often used as a simple proxy to measure the preservation of global cognitive function in older adults. However, it is unknown which specific selection, decision, and/or motor processes provide the most specific indices of neurocognitive health. Here, using a simple decision task with electroencephalography (EEG), we found that the efficiency with which an individual accumulates sensory evidence was a critical determinant of the extent to which RS was preserved in older adults (63% female, 37% male). Moreover, the mitigating influence of EE on age-related RS declines was most pronounced when evidence accumulation rates were shallowest. These results suggest that the phenomenon of cognitive reserve, whereby high EE individuals can better tolerate suboptimal brain health to facilitate the preservation of cognitive function, is not just applicable to neuroanatomical indicators of brain aging but can be observed in markers of neurophysiology. Our results suggest that EEG metrics of evidence accumulation may index neurocognitive vulnerability of the aging brain.Significance Statement Response speed in older adults is closely linked with trajectories of cognitive aging. Here, by recording brain activity while individuals perform a simple computer task, we identify a neural metric that is a critical determinant of response speed. Older adults exposed to greater cognitive and social stimulation throughout a lifetime could maintain faster responding, even when this neural metric was impaired. This work suggests EEG is a useful technique for interrogating how a lifetime of stimulation benefits brain health in aging.

Attention in flux.

Selective attention comprises essential infrastructural functions supporting cognition-anticipating, prioritizing, selecting, routing, integrating, and preparing signals to guide adaptive behavior. Most studies have examined its consequences, systems, and mechanisms in a static way, but attention is at the confluence of multiple sources of flux. The world advances, we operate within it, our minds change, and all resulting signals progress through multiple pathways within the dynamic networks of our brains. Our aim in this review is to raise awareness of and interest in three important facets of how timing impacts our understanding of attention. These include the challenges posed to attention by the timing of neural processing and psychological functions, the opportunities conferred to attention by various temporal structures in the environment, and how tracking the time courses of neural and behavioral modulations with continuous measures yields surprising insights into the workings and principles of attention.

Multiple and Dissociable Effects of Sensory History on Working-Memory Performance.

Behavioral reports of sensory information are biased by stimulus history. The nature and direction of such serial-dependence biases can differ between experimental settings; both attractive and repulsive biases toward previous stimuli have been observed. How and when these biases arise in the human brain remains largely unexplored. They could occur either via a change in sensory processing itself and/or during postperceptual processes such as maintenance or decision-making. To address this, we tested 20 participants (11 female) and analyzed behavioral and magnetoencephalographic (MEG) data from a working-memory task in which participants were sequentially presented with two randomly oriented gratings, one of which was cued for recall at the end of the trial. Behavioral responses showed evidence for two distinct biases: (1) a within-trial repulsive bias away from the previously encoded orientation on the same trial, and (2) a between-trial attractive bias toward the task-relevant orientation on the previous trial. Multivariate classification of stimulus orientation revealed that neural representations during stimulus encoding were biased away from the previous grating orientation, regardless of whether we considered the within-trial or between-trial prior orientation, despite opposite effects on behavior. These results suggest that repulsive biases occur at the level of sensory processing and can be overridden at postperceptual stages to result in attractive biases in behavior.SIGNIFICANCE STATEMENT Recent experience biases behavioral reports of sensory information, possibly capitalizing on the temporal regularity in our environment. It is still unclear at what stage of stimulus processing such serial biases arise. Here, we recorded behavior and neurophysiological [magnetoencephalographic (MEG)] data to test whether neural activity patterns during early sensory processing show the same biases seen in participants' reports. In a working-memory task that produced multiple biases in behavior, responses were biased toward previous targets, but away from more recent stimuli. Neural activity patterns were uniformly biased away from all previously relevant items. Our results contradict proposals that all serial biases arise at an early sensory processing stage. Instead, neural activity exhibited mostly adaptation-like responses to recent stimuli.

Microsaccades transiently lateralise EEG alpha activity.

The lateralisation of 8-12 Hz alpha activity is a canonical signature of human spatial cognition that is typically studied under strict fixation requirements. Yet, even during attempted fixation, the brain produces small involuntary eye movements known as microsaccades. Here we report how spontaneous microsaccades - made in the absence of incentives to look elsewhere - can themselves drive transient lateralisation of EEG alpha power according to microsaccade direction. This transient lateralisation of posterior alpha power occurs similarly following start and return microsaccades and is, at least for start microsaccades, driven by increased alpha power ipsilateral to microsaccade direction. This reveals new links between spontaneous microsaccades and human electrophysiological brain activity. It highlights how microsaccades are an important factor to consider in studies relating alpha activity - including spontaneous fluctuations in alpha activity - to spatial cognition, such as studies on visual attention, anticipation, and working memory.

Heading Direction Tracks Internally Directed Selective Attention in Visual Working Memory.

We shift our gaze even when we orient attention internally to visual representations in working memory. Here, we show the bodily orienting response associated with internal selective attention is widespread as it also includes the head. In three virtual reality experiments, participants remembered 2 visual items. After a working memory delay, a central color cue indicated which item needed to be reproduced from memory. After the cue, head movements became biased in the direction of the memorized location of the cued memory item-despite there being no items to orient toward in the external environment. The heading-direction bias had a distinct temporal profile from the gaze bias. Our findings reveal that directing attention within the spatial layout of visual working memory bears a strong relation to the overt head orienting response we engage when directing attention to sensory information in the external environment. The heading-direction bias further demonstrates common neural circuitry is engaged during external and internal orienting of attention.

The Relationship between Short- and Long-Term Memory Is Preserved across the Age Range.

Both short- and long-term memories decline with healthy ageing. The aims of the current study were twofold: firstly, to build on previous studies and investigate the presence of a relationship between short- and long-term memories and, secondly, to examine cross-sectionally whether there are changes in this relationship with age. In two experiments, participants across the age range were tested on contextual-spatial memories after short and long memory durations. Experimental control in stimulus materials and task demands enabled the analogous encoding and probing for both memory durations, allowing us to examine the relationship between the two memory systems. Across two experiments, in line with previous studies, we found both short-term memory and long-term memory declined from early to late adulthood. Additionally, there was a significant relationship between short- and long-term memory performance, which, interestingly, persisted throughout the age range. Our findings suggest a significant degree of common vulnerability to healthy ageing for short- and long-term memories sharing the same spatial-contextual associations. Furthermore, our tasks provide a sensitive and promising framework for assessing and comparing memory function at different timescales in disorders with memory deficits at their core.

Turning Attention Inside Out: How Working Memory Serves Behavior.

Flexible behavior requires guidance not only by sensations that are available immediately but also by relevant mental contents carried forward through working memory. Therefore, selective-attention functions that modulate the contents of working memory to guide behavior (inside-out) are just as important as those operating on sensory signals to generate internal contents (outside-in). We review the burgeoning literature on selective attention in the inside-out direction and underscore its functional, flexible, and future-focused nature. We discuss in turn the purpose (why), targets (what), sources (when), and mechanisms (how) of selective attention inside working memory, using visual working memory as a model. We show how the study of internal selective attention brings new insights concerning the core cognitive processes of attention and working memory and how considering selective attention and working memory together paves the way for a rich and integrated understanding of how mind serves behavior.

New Therapeutics in Alzheimer's Disease Longitudinal Cohort study (NTAD): study protocol.

INTRODUCTION: With the pressing need to develop treatments that slow or stop the progression of Alzheimer's disease, new tools are needed to reduce clinical trial duration and validate new targets for human therapeutics. Such tools could be derived from neurophysiological measurements of disease. METHODS AND ANALYSIS: The New Therapeutics in Alzheimer's Disease study (NTAD) aims to identify a biomarker set from magneto/electroencephalography that is sensitive to disease and progression over 1 year. The study will recruit 100 people with amyloid-positive mild cognitive impairment or early-stage Alzheimer's disease and 30 healthy controls aged between 50 and 85 years. Measurements of the clinical, cognitive and imaging data (magnetoencephalography, electroencephalography and MRI) of all participants will be taken at baseline. These measurements will be repeated after approximately 1 year on participants with Alzheimer's disease or mild cognitive impairment, and clinical and cognitive assessment of these participants will be repeated again after approximately 2 years. To assess reliability of magneto/electroencephalographic changes, a subset of 30 participants with mild cognitive impairment or early-stage Alzheimer's disease will also undergo repeat magneto/electroencephalography 2 weeks after baseline. Baseline and longitudinal changes in neurophysiology are the primary analyses of interest. Additional outputs will include atrophy and cognitive change and estimated numbers needed to treat each arm of simulated clinical trials of a future disease-modifying therapy. ETHICS AND DATA STATEMENT: The study has received a favourable opinion from the East of England Cambridge Central Research Ethics Committee (REC reference 18/EE/0042). Results will be disseminated through internal reports, peer-reviewed scientific journals, conference presentations, website publication, submission to regulatory authorities and other publications. Data will be made available via the Dementias Platform UK Data Portal on completion of initial analyses by the NTAD study group.

Opening Questions in Visual Working Memory.

In this reflective piece on visual working memory, I depart from the laboriously honed skills of writing a review. Instead of integrating approaches, synthesizing evidence, and building a cohesive perspective, I scratch my head and share niggles and puzzlements. I expose where my scholarship and understanding are stumped by findings and standard views in the literature.

Functional but not obligatory link between microsaccades and neural modulation by covert spatial attention.

Covert spatial attention is associated with spatial modulation of neural activity as well as with directional biases in fixational eye movements known as microsaccades. We studied how these two 'fingerprints' of attention are interrelated in humans. We investigated spatial modulation of 8-12 Hz EEG alpha activity and microsaccades when attention is directed internally within the spatial layout of visual working memory. Consistent with a common origin, spatial modulations of alpha activity and microsaccades co-vary: alpha lateralisation is stronger in trials with microsaccades toward versus away from the memorised location of the to-be-attended item and occurs earlier in trials with earlier microsaccades toward this item. Critically, however, trials without attention-driven microsaccades nevertheless show clear spatial modulation of alpha activity - comparable to trials with attention-driven microsaccades. Thus, directional biases in microsaccades correlate with neural signatures of spatial attention, but they are not necessary for neural modulation by spatial attention to be manifest.

Transient beta activity and cortico-muscular connectivity during sustained motor behaviour.

Neural oscillations are thought to play a central role in orchestrating activity states between distant neural populations. For example, during isometric contraction, 13-30 Hz beta activity becomes phase coupled between the motor cortex and the contralateral muscle. This and related observations have led to the proposal that beta activity and connectivity sustain stable cognitive and motor states - or the 'status quo' - in the brain. Recently, however, beta activity at the single-trial level has been shown to be short-lived - though so far this has been reported for regional beta activity in tasks without sustained motor demands. Here, we measured magnetoencephalography (MEG) and electromyography (EMG) in 18 human participants performing a sustained isometric contraction (gripping) task. If cortico-muscular beta connectivity is directly responsible for sustaining a stable motor state, then beta activity within single trials should be (or become) sustained in this context. In contrast, we found that motor beta activity and connectivity with the downstream muscle were transient. Moreover, we found that sustained motor requirements did not prolong beta-event duration in comparison to rest. These findings suggest that neural synchronisation between the brain and the muscle involves short 'bursts' of frequency-specific connectivity, even when task demands - and motor behaviour - are sustained.

Consequences of predictable temporal structure in multi-task situations.

In everyday life, we often anticipate the timing of an upcoming task or event while actively engaging in another. Here, we investigated the effects of predictable temporal structure within such a multi-task scenario. In a visual working-memory task, we manipulated whether the onset of a working-memory probe could be predicted in time, while also embedding a simple intervening task within the delay period. We first show that working-memory performance benefitted from temporal expectations, even though an intervening task had to be completed in the interim. Moreover, temporal predictions regarding the upcoming working-memory probe additionally affected performance on the intervening task, resulting in faster responses when the memory probe was expected early, and slower responses when the memory probe was expected late, as compared to when it was temporally unpredictable. Because the intervening task always occurred at the same time during the memory delay, differences in performance on this intervening task result from a between-task consequence of temporal expectation. Thus, we show that within multi-task settings, knowing when working-memory contents will be required for an upcoming task not only facilitates performance of the associated working-memory task, but can also influence the performance of other, intervening tasks.

Be there on time: Spatial-temporal regularities guide young children's attention in dynamic environments.

Children's ability to benefit from spatiotemporal regularities to detect goal-relevant targets was tested in a dynamic, extended context. Young adults and children (from a low-deprivation area school in the United Kingdom; N = 80; 5-6 years; 39 female; ethics approval did not permit individual-level race/ethnicity surveying) completed a dynamic visual-search task. Targets and distractors faded in and out of a display over seconds. Half of the targets appeared at predictable times and locations. Search performance in children was poorer overall. Nevertheless, they benefitted equivalently from spatiotemporal regularities, detecting more predictable than unpredictable targets. Children's benefits from predictions correlated positively with their attention. The study brings ecological validity to the study of attentional guidance in children, revealing striking behavioral benefits of dynamic experience-based predictions.

Eyes wide open: Regulation of arousal by temporal expectations.

Maintaining adequate levels of arousal is essential for sustaining performance on extended tasks. To investigate arousal in prolonged tasks such as driving studies have traditionally used monotonous task designs. Both ecological and experimental settings often contain embedded temporal regularities, but it is unknown whether these enable adaptive modulation of arousal. We explored whether temporal predictability can modulate arousal according to the timing of anticipated relevant events. In two experiments, we manipulated the temporal predictability of events to test for behavioural benefits and arousal modulation, using pupillometry as a proxy measure. High temporal predictability significantly lowered the tonic level of arousal briefly increased arousal in anticipation of upcoming stimuli, whereas low temporal predictability resulted in tonically elevated arousal. These novel findings suggest that arousal levels flexibly adapt to the temporal structures of events and bring about energy efficiencies in the context of high levels of behavioural performance.