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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.

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 STATEMENTRecent 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.

We developed a new experimental approach to compare directly how attentional orienting facilitates retrieval in human working memory and long-term memory, and how selective attention within these different memory timescales impacts incoming sensory information processing. In two experiments with healthy young adults (Ns = 30 and 44), retrospective attention cues prioritised an item encoded in working memory or long-term memory. Participants then retrieved a memory item or performed a perceptual discrimination task. The cue was informative for the retrieval task but not for the perceptual task. Attentional orienting improved memory retrieval for both memory types and also enhanced discrimination for visual stimuli at the location matching the prioritised working memory or long-term memory item. Eye-tracking data revealed a striking dissociation in gaze biases related to orienting in working memory vs. long-term memory. The findings suggest potent and at least partly dissociable attention-orienting processes for different memory timescales.

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

IntroductionMood instability in bipolar disorder (BD) is poorly understood. Here we examined cognitive and neural mechanisms related to these fluctuations and how they are changed with the mood stabilizer lithium.MethodsWe recruited volunteers with low (n=37) or high (n=40) risk of BD (using the Mood Disorder Questionnaire, MDQ). We also recruited patients with BD who were assigned (randomized, double-blind) to six weeks of lithium (n=19) or placebo (n=16) after a two-week baseline period. Participants completed mood ratings daily over 50 (healthy) or 42 (BD) days, as well as a risky decision-making task and one functional magnetic resonance imaging session. The task measured adaptation of risk taking to past outcomes (increased risk aversion after a previous win, ‘outcome history’).ResultsWhile the low MDQ group was risk averse after a win, this was less evident in the high MDQ group and least so in the patients with BD. Neurally, ‘outcome history’ was linked to medial frontal pole activation at the time of the decision. Corresponding to the behavioural effect, this activation was reduced in the high MDQ vs. the low MDQ group. While lithium did not reverse the pattern of BD in the task, it changed reward processing in the dorsolateral prefrontal cortex.DiscussionHealthy participants’ modulation of risk-taking in response to reward outcomes was reduced by risk of BD and BD. These results provide a model for how reward may prime escalation of risk-related behaviours in bipolar disorder and how mood stabilising treatments may work.

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.

Spectrum estimators that make use of averaging across time segments are ubiquitous across neuroscience. The core of this approach has not changed substantially since the 1960s, though many advances in the field of regression modelling and statistics have been made during this time. Here, we propose a new approach, the General Linear Model (GLM) Spectrum, which reframes time averaged spectral estimation as multiple regression. This brings several benefits, including the ability to do confound modelling, hierarchical modelling and significance testing via non-parametric statistics.We apply the approach to a first-level EEG resting-state dataset alternating between eyes open and eyes closed resting-state. The GLM-Spectrum can model both conditions, quantify their differences, and perform denoising through confound regression in a single step. This application is scaled up from a single channel to a whole-head recording and, finally, applied to quantify age differences across a large group-level dataset. We show that the GLM-Spectrum lends itself to rigorous modelling of within- and between-subject contrasts as well as their interactions, and that the use of model-projected spectra provides an intuitive visualisation. The GLM-Spectrum is a flexible framework for robust multi-level analysis of power spectra, with adaptive covariance and confound modelling.

International audience; Despite increased awareness of the lack of gender equity in academia and a growing number of initiatives to address issues of diversity, change is slow, and inequalities remain. A major source of inequity is gender bias, which has a substantial negative impact on the careers, work-life balance, and mental health of underrepresented groups in science. Here, we argue that gender bias is not a single problem but manifests as a collection of distinct issues that impact researchers’ lives. We disentangle these facets and propose concrete solutions that can be adopted by individuals, academic institutions, and society

Working memory enables us to retain past sensations in service of anticipated task demands. How we prepare for anticipated task demands during working memory retention remains poorly understood. Here, we focused on the role of time—asking how temporal expectations help prepare for ensuing memory-guided behavior. We manipulated the expected probe time in a delayed change-detection task and report that temporal expectation can have a profound influence on memory-guided behavioral performance. EEG measurements corroborated the utilization of temporal expectations: demonstrating the involvement of a classic EEG signature of temporal expectation—the contingent negative variation—in the context of working memory. We also report the influence of temporal expectations on 2 EEG signatures associated with visual working memory—the lateralization of 8- to 12-Hz alpha activity, and the contralateral delay activity. We observed a dissociation between these signatures, whereby alpha lateralization (but not the contralateral delay activity) adapted to the time of expected memory utilization. These data show how temporal expectations prepare visual working memory for behavior and shed new light on the electrophysiological markers of both temporal expectation and working memory.

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.

This study tested whether observers can benefit from temporal regularities occurring within dynamic and distraction-filled contexts to identify target stimuli. Across two online experiments, participants searched for multiple targets amongst distractors in dynamic visual- search tasks in which coloured stimuli faded in and out of the display at different times and locations. In Experiment 1, half of the targets across trials were spatiotemporally predictable, occurring at consistent time points and spatial quadrants. The other targets and the many competing distracting stimuli were completely unpredictable. Colour was unpredictable for all stimuli. In Experiment 2, half of the targets across trials were feature-temporally predictable. They appeared at consistent time points and with consistent colours. The other targets and competing distractors had unpredictable timings and colours, and all stimuli had unpredictable spatial locations. In both experiments, temporal predictability of targets was separate from just temporal order. The pattern of results was consistent across experiments, showing that participants benefitted significantly from spatiotemporal (Experiment 1) and feature-temporal (Experiment 2) predictions for identifying targets. The ability to utilise temporal regularities that occur across unpredictable intervening stimuli, and the ability to utilise such temporal regularities independently of spatial predictions, carry important theoretical implications for understanding the nature of temporal expectations. An exploratory questionnaire after each experiment indicated that participants learned spatiotemporal and feature-temporal predictions incidentally and used them implicitly and without awareness. Our study highlights the importance of the temporal dimension for guiding behaviour within dynamic and competitive contexts mirroring ecological situations and opens new questions for investigation.

Significance Everyday behavior relies on continuously selecting relevant information from the external environment as well as from internal representations in working memory. In describing the selection of external objects, a longstanding distinction is proposed between goal-directed (voluntary) and stimulus-driven (involuntary) sources of attentional selection. We voluntarily attend to things that are relevant to our goals, but our attention may also be captured involuntarily by salient events in the external world. Yet despite decades of research on how these two sources guide our perception (external selection), to date no study has examined whether and how these two sources jointly influence the selection of internal memory representations. With innovative experimental manipulations and behavioral markers of internal selection, we fill this important gap., Adaptive behavior relies on the selection of relevant sensory information from both the external environment and internal memory representations. In understanding external selection, a classic distinction is made between voluntary (goal-directed) and involuntary (stimulus-driven) guidance of attention. We have developed a task—the anti-retrocue task—to separate and examine voluntary and involuntary guidance of attention to internal representations in visual working memory. We show that both voluntary and involuntary factors influence memory performance but do so in distinct ways. Moreover, by tracking gaze biases linked to attentional focusing in memory, we provide direct evidence for an involuntary “retro-capture” effect whereby external stimuli involuntarily trigger the selection of feature-matching internal representations. We show that stimulus-driven and goal-directed influences compete for selection in memory, and that the balance of this competition—as reflected in oculomotor signatures of internal attention—predicts the quality of ensuing memory-guided behavior. Thus, goal-directed and stimulus-driven factors together determine the fate not only of perception, but also of internal representations in working memory.

At any given moment, we receive multiple signals from our different senses. Prior research has shown that signals in one sensory modality can influence neural activity and behavioural performance associated with another sensory modality. Recent human and nonhuman primate studies suggest that such cross-modal influences in sensory cortices are mediated by the synchronisation of ongoing neural oscillations. In this review, we consider two mechanisms proposed to facilitate cross-modal influences on sensory processing, namely cross-modal phase resetting and neural entrainment. We consider how top-down processes may further influence cross-modal processing in a flexible manner, and we highlight fruitful directions for further research., Highlights Electrophysiological studies suggest that cross-modal influences in sensory cortices are mediated by the synchronisation of neural oscillations through phase-resetting and neural entrainment mechanisms. Low-frequency neural oscillations in the delta, theta, and lower alpha ranges are suggested to provide temporal windows for cross-modal influences. Top-down factors, such as task goals and expectations, may modulate multisensory processing as well as neural oscillations. Bayesian computational modelling provides a new approach to probe cross-modal influences.

Protecting working-memory content from distracting external sensory inputs and intervening tasks is an ubiquitous demand in daily life. Here, we ask whether and how temporal expectations about external events can help mitigate effects of such interference during working-memory retention. We manipulated the temporal predictability of interfering items that occurred during the retention period of a visual working-memory task and report that temporal expectations reduce the detrimental influence of external interference on subsequent memory performance. Moreover, to determine if the protective effects of temporal expectations rely on distractor suppression or involve shielding of internal representations, we compared effects after irrelevant distractors that could be ignored vs. interrupters that required a response. Whereas distractor suppression may be sufficient to confer protection from predictable distractors, any benefits after interruption are likely to involve memory shielding. We found similar benefits of temporal expectations after both types of interference. We conclude that temporal expectations may play an important role in safeguarding behaviour based on working memory – acting through mechanisms that include the shielding of internal content from external interference.

Covert spatial attention is associated with spatially specific modulation of neural activity as well as with directional biases in fixational eye-movements known as microsaccades. Recently, this link has been suggested to be obligatory, such that modulation of neural activity by covert spatial attention occurs only when paired with microsaccades toward the attended location. Here we revisited this link between microsaccades and neural modulation by covert spatial attention in humans. We investigated spatial modulation of 8-12 Hz EEG alpha activity and microsaccades in a context with no incentive for overt gaze behaviour: when attention is directed internally within the spatial layout of visual working memory. In line with a common attentional origin, we show that spatial modulations of alpha activity and microsaccades co-vary: alpha lateralisation is stronger in trials with microsaccades toward compared to 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 showed clear spatial modulation of alpha activity – comparable to the neural modulation observed in trials with attention-driven microsaccades. Thus, directional biases in microsaccades are correlated with neural signatures of covert spatial attention, but they are not a prerequisite for neural modulation by covert spatial attention to be manifest.

Behavioural 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 towards 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 post-perceptual processes such as maintenance or decision-making. To address this, we analysed behavioural and magnetoencephalographic 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. Behavioural 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 towards 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- or between-trial prior orientation – despite opposite effects on behaviour. These results suggest that repulsive biases occur at the level of sensory processing and can be overridden at post-perceptual stages to result in attractive biases in behaviour.

Older adults exposed to enriched environments (EE) 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. 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 ageing, but can be observed in markers of neurophysiology. Our results suggest that EEG metrics of evidence accumulation may index neurocognitive vulnerability of the ageing brain.SIGNIFICANCE STATEMENTResponse speed in older adults is closely linked with trajectories of cognitive ageing. Here, by recording brain activity while individuals perform a simple computer task, we identify a neural metric which 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 ageing.

Recent advances have made it possible to decode various aspects of visually presented stimuli from patterns of scalp EEG measurements. As of recently, such multivariate methods have been commonly used to decode visual-spatial features such as location, orientation, or spatial frequency. In the current study, we show that it is also possible to track visual colour processing by using Linear Discriminant Analysis on patterns of EEG activity. Building on other recent demonstrations, we show that colour decoding: (1) reflects sensory qualities (as opposed to, for example, verbal labelling) with a prominent contribution from posterior electrodes contralateral to the stimulus, (2) conforms to a parametric coding space, (3) is possible in multi-item displays, and (4) is comparable in magnitude to the decoding of visual stimulus orientation. Through subsampling our data, we also provide an estimate of the approximate number of trials and participants required for robust decoding. Finally, we show that while colour decoding can be sensitive to subtle differences in luminance, our colour decoding results are primarily driven by measured colour differences between stimuli. Colour decoding opens a relevant new dimension in which to track visual processing using scalp EEG measurements, while bypassing potential confounds associated with decoding approaches that focus on spatial features., Graphical abstract Image, graphical abstract

At any given moment our sensory systems receive multiple, often rhythmic, inputs from the environment. Processing of temporally structured events in one sensory modality can guide both behavioural and neural processing of events in other sensory modalities, but whether this occurs remains unclear. Here, we used human electroencephalography (EEG) to test the cross-modal influences of a continuous auditory frequency-modulated (FM) sound on visual perception and visual cortical activity. We report systematic fluctuations in perceptual discrimination of brief visual stimuli in line with the phase of the FM sound. We further show that this rhythmic modulation in visual perception is related to an accompanying rhythmic modulation of neural activity recorded over visual areas. Importantly, in our task, perceptual and neural visual modulations occurred without any abrupt and salient onsets in the energy of the auditory stimulation and without any rhythmic structure in the visual stimulus. As such, the results provide a critical validation for the existence and functional role of cross-modal entrainment and demonstrates its utility for organising the perception of multisensory stimulation in the natural environment.Significance StatementOur sensory environment is filled with rhythmic structures that are often multi-sensory in nature. Here we show that the alignment of neural activity to the phase of an auditory frequency-modulated sound has cross-modal consequences for vision: yielding systematic fluctuations in perceptual discrimination of brief visual stimuli that is mediated by accompanying rhythmic modulation of neural activity recorded over visual areas. These cross-modal effects on visual neural activity and perception occurred without any abrupt and salient onsets in the energy of the auditory stimulation and without any rhythmic structure in the visual stimulus. The current work shows that continuous auditory fluctuations in the natural environment can provide a pacing signal for neural activity and perception across the senses.

Memories are about the past, but they serve the future. Memory research often emphasizes the former aspect: focusing on the functions that re-constitute (re-member) experience and elucidating the various types of memories and their interrelations, timescales, and neural bases. Here we highlight the prospective nature of memory in guiding selective attention, focusing on functions that use previous experience to anticipate the relevant events about to unfold—to “premember” experience. Memories of various types and timescales play a fundamental role in guiding perception and performance adaptively, proactively, and dynamically. Consonant with this perspective, memories are often recorded according to expected future demands. Using working memory as an example, we consider how mnemonic content is selected and represented for future use. This perspective moves away from the traditional representational account of memory toward a functional account in which forward-looking memory traces are informationally and computationally tuned for interacting with incoming sensory signals to guide adaptive behavior., Nobre and Stokes consider how memories guide proactive attention. Introducing the concept of premembering, this perspective advances a functional account of memory. Memory traces are informationally and computationally tuned for interacting with incoming sensory signals to guide adaptive behavior.

Brain areas that control gaze are also recruited for covert shifts of spatial attention1–9. In the external space of perception, there is a natural, ecological link between the control of gaze and of spatial attention, since information sampled at covertly attended locations can inform where to look next2,10,11. Attention can also be directed internally to representations held within the spatial layout of visual working memory12–16. In such cases, the incentive for using attention to direct gaze disappears since there are no external targets to scan. Here we investigate whether the brain’s oculomotor system also participates in attention within the internal space of memory. Paradoxically, we reveal this participation through gaze behaviour itself. We demonstrate that selecting an item from visual working memory biases gaze in the direction of the memorised location of that item – despite there being nothing to look at and even though location memory was never explicitly probed. This retrospective ‘gaze bias’ occurs only when an item is not already in the internal focus of attention, and predicts the performance benefit associated with the focusing of internal attention. We conclude that the oculomotor system also participates in the focusing of attention within memorised space, leaving traces all the way to the eyes.

In this article, we propose a method to track trial-specific neural dynamics of stimulus processing and decision making with high temporal precision. By applying this novel method to a perceptual template-matching task, we tracked representational brain states associated with the cascade of neural processing, from early sensory areas to higher order areas that are involved in integration and decision making. We address a major limitation of the traditional decoding approach: that it relies on consistent timing of these processes over trials. Using a TUDA approach, we found that the timing of the cognitive processes involved in perceptual judgments can vary considerably over trials. This revealed that the sequence of processing states was consistent for all subjects and trials, even when the timing of these states varied. Furthermore, we found that the specific timing of states on each trial was related to the quality of performance over trials. Altogether, this work not only highlights the serious pitfalls and misleading interpretations that result from assuming stimulus processing to be synchronous across trials but can also open important avenues to investigate learning and quantify plasticity.

IntroductionWith the pressing need to develop treatments that slow or stop the progression of Alzheimers 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 AnalysisThe New Therapeutics in Alzheimers disease study (NTAD) aims to identify a biomarker set from magneto/electro-encephalographic that is sensitive to disease and progression over one year. The study will recruit 100 people with amyloid-positive mild cognitive impairment or early-stage Alzheimers disease and 30 healthy controls aged between 50 and 85 years. Repeat measurements of the clinical, cognitive and imaging data (magnetoencephalography, electroencephalography and magnetic resonance imaging) of participants with Alzheimers disease or mild cognitive impairment will be taken at baseline and at one year. To assess reliability of magneto/electro-encephalographic changes, a subset of 30 participants with mild cognitive impairment or early-stage Alzheimers disease will undergo repeat magneto/electro-encephalographic two weeks after baseline. Clinical and cognitive assessment will be repeated at 2 years. Linear mixed models of baseline and longitudinal change in neurophysiology are the primary analyses of interest, supported by Bayesian inference. Additional outputs will include relative effect sizes for physiological markers, atrophy and cognitive change and the respective numbers needed to treat each arm of simulated clinical trials of a future disease modifying therapy. Ethics and disseminationThe 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. STRENGTHS AND LIMITATIONS OF THIS STUDYO_ST_ABSStrengthsC_ST_ABSO_LINTAD is a longitudinal, multicentre study of magneto/electro-encephalographic measures of Alzheimers disease progression C_LIO_LINTAD assesses the test-retest reliability of magneto/electro-encephalographic parameters C_LIO_LIAll participants with early-stage Alzheimers disease or mild cognitive impairment will be amyloid-positive C_LI LimitationsO_LIAttrition during follow up may limit inferences C_LIO_LIRecruitment is from volunteer panels and clinical services that may not reflect national diversity of people affected by Alzheimers disease C_LI

Temporal orienting of attention can affect multiple stages of processing to guide adaptive behaviour. We tested whether temporal expectation in different task contexts is compromised in individuals with Parkinson's disease (PD). In Experiment 1 two temporal‐orienting tasks were used: a speeded task emphasizing motor preparation and a non‐speeded task emphasizing perceptual discrimination using rapid serial visual presentation. In both tasks, auditory cues indicated the likelihood of a target appearing after a short or long interval. In the speeded‐response task, participants used the cues to anticipate an easily detectable target stimulus. In the non‐speeded perceptual‐discrimination task, participants used the cues to help discriminate a target letter embedded in a stream of letters. Relative to healthy participants, participants with PD did not show altered temporal orienting effects in the speeded‐response task. However, they were impaired in using temporal cues to improve perceptual discrimination. In Experiment 2, we tested whether the temporal‐orienting deficits in the perceptual‐discrimination task depended on the requirement to ignore temporally distracting stimuli. We replicated the impaired temporal orienting for perceptual discrimination in an independent group of individuals with PD, and showed the impairment was abolished when individuals were on their dopaminergic medication. In a task without any distracting letters, however, patients off or on medication benefited normally from temporal orienting cues. Our findings suggest that deficits in temporal orienting in individuals with PD interact with specific task demands, such as the requirement to select target from temporally competing distractors., Temporal orienting of attention can affect multiple stages of processing to guide adaptive behaviour. We tested whether temporal expectation in different task contexts is compromised in individuals with Parkinson’s disease (PD). In Experiment 1 two temporal‐orienting tasks were used: a speeded task emphasizing motor preparation and a non‐speeded task emphasizing perceptual discrimination using rapid serial visual presentation (RSVP). In both tasks, auditory cues indicated the likelihood of a target appearing after a short or long interval. In the speeded‐response task, participants used the cues to anticipate an easily detectable target stimulus. In the non‐speeded perceptual‐discrimination task, participants used the cues to help discriminate a target letter embedded in a stream of letters. Relative to healthy participants, participants with PD did not show altered temporal orienting effects in the speeded‐response task. However, they were impaired in using temporal cues to improve perceptual discrimination. In Experiment 2, we tested whether the temporal‐orienting deficits in the perceptual‐discrimination task depended on the requirement to ignore temporally distracting stimuli. We replicated the impaired temporal orienting for perceptual discrimination in an independent group of individuals with PD, and showed the impairment was abolished when individuals were on their dopaminergic medication. In a task without any distracting letters, however, patients off or on medication benefited normally from temporal orienting cues. Our findings suggest that deficits in temporal orienting in individuals with PD interact with specific task demands, such as the requirement to select target from temporally competing distractors.

Neural oscillations are thought to play a central role in orchestrating activity states between distant neural populations. In humans, long-range neural connectivity has been particularly well characterised for 13-30 Hz beta activity which becomes phase coupled between the motor cortex and the contralateral muscle during isometric contraction. Based on this and related observations, beta activity and connectivity have been linked to sustaining stable cognitive and motor states – or the ‘status quo’ – in the brain. Recently, however, beta activity has been shown to be short-lived, as opposed to sustained – 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 an isometric-contraction (gripping) task designed to yield sustained behavioural output. If cortico-muscular beta connectivity is directly responsible for sustaining a stable motor state, then beta activity should be (or become) sustained in this context. In contrast, we found that beta activity and connectivity with the downstream muscle were transient, even when participants engaged in sustained gripping. Moreover, we found that sustained motor requirements did not prolong beta-event duration in comparison to rest. These findings suggest that long-range neural synchronisation may entail short ‘bursts’ of frequency-specific connectivity, even when task demands – and behaviour – are sustained.HighlightsTrial-average 13-30 Hz beta activity and connectivity with the muscle appear sustained during stable motor behaviourSingle-trial beta activity and connectivity are short-lived, even when motor behaviour is sustainedSustained task demands do not prolong beta-event duration in comparison to resting state

To perform a task continuously over an extended period of time, it is necessary to maintain an adequate level of arousal. In cognitive research, traditional studies have used repetitive, monotonous tasks to learn about the dynamics of arousal in tasks that require sustained vigilance, such as driving or monitoring a factory line. To date, studies have rarely considered whether observers use task-embedded regularities in such continuous contexts to anticipate events and regulate arousal accordingly. In the current study, we explored whether arousal is an adaptive process that reacts to temporal stimulus predictability. Across two experiments, we used pupillometry as a proxy measure of arousal in human observers performing continuous tasks. Within the tasks, we manipulated the temporal predictability of relevant events by presenting stimuli either in a fixed rhythm or at varied intervals. Temporal predictability led to the lowering of tonic levels of arousal. Trial-wise temporal structures led to short-scale changes in pupil size related to temporal anticipation. Accordingly, we suggest that arousal is sensitive to varying levels of temporal predictability and dynamically adapts to task demands to bring about performance benefits as well as energetic efficiencies overall.Statement of RelevancePeople often have to sustain focus and high levels of performance on extended and non-stimulating tasks (e.g., driving, sowing, monitoring data acquisition). A critical factor to the success (or failure) in sustained performance is arousal – the ‘energetic state’ of the cognitive system. Here we used pupil dilation as a proxy to study levels of arousal during sustained performance on monotonous tasks. We reveal that arousal is dynamically regulated to support performance according to an important fundamental property of any task: its temporal structure. When the timing of task-relevant events is predictable, arousal levels fluctuate accordingly, saving energy overall while also optimally guiding performance. Our study and findings add ecological validity to the study of temporal expectations, by moving investigations beyond typical trial-by-trial designs. They also carry significant implications for clinical studies relying on sustained performance tasks.

Summary Working memory can maintain multiple sensory representations to serve unfolding sequential behaviour, such as while making tea or planning a route. How the human mind juggles internal representations as they become relevant to guide sequential behaviour remains poorly understood. Specifically, while there is good evidence that we can flexibly switch priorities among representations in working memory1, 2, 3, 4, it is unclear how and when dormant memory representations are brought into focus during sequential behaviour. Capitalising on a recently established and temporally precise gaze marker of internal selection5,6, we reveal that the focus in the mind moves to the next-relevant memory representation while behaviour associated with the presently relevant memory representation is still ongoing. Thus, like visual sampling of external objects in the world7, 8, 9, internal visual sampling also ‘looks ahead’ to the next object in memory during sequential behaviour., van Ede et al. show that, when sequential behaviour is guided by internal visual representations, the focus in mind ‘looks ahead’ to the next object in memory while behaviour associated with the preceding object is still ongoing.

Long-range communication through the motor system is thought to be facilitated by phase coupling between neural activity in the 15–30 Hz beta range. During periods of sustained muscle contraction (grip), such coupling is manifest between motor cortex and the contralateral forearm muscles—measured as the cortico-muscular coherence. We examined alterations in cortico-muscular coherence in individuals with Parkinson’s disease, while equating grip strength between individuals with Parkinson’s disease (off their medication) and healthy control participants. We show a marked reduction in beta cortico-muscular coherence in the Parkinson’s disease group, even though the grip strength was comparable between the two groups. Moreover, the reduced cortico-muscular coherence was related to motor symptoms, so that individuals with lower cortico-muscular coherence also displayed worse motor symptoms. These findings highlight the cortico-muscular coherence as a simple, effective and clinically relevant neural marker of Parkinson’s disease pathology, with the potential to aid monitoring of disease progression and the efficacy of novel treatments for Parkinson’s disease., Zokaei et al. report reduced cortico-muscular coherence—coupling between cortex and muscle—in individuals with Parkinson’s disease compared to controls, during well-equated muscle contraction between the two groups. This reduction is related to motor symptoms in patients. This measure can provide a sensitive and objective measure of disease-related motor deficits., Graphical Abstract Graphical Abstract

The human brain immediately plans for prospective action when storing sensations into working memory., Working memory serves as the buffer between past sensations and future behavior, making it vital to understand not only how we encode and retain sensory information in memory but also how we plan for its upcoming use. We ask when prospective action goals emerge alongside the encoding and retention of visual information in working memory. We show that prospective action plans do not emerge gradually during memory delays but are brought into memory early, in tandem with sensory encoding. This action encoding (i) precedes a second stage of action preparation that adapts to the time of expected memory utilization, (ii) occurs even ahead of an intervening motor task, and (iii) predicts visual memory–guided behavior several seconds later. By bringing prospective action plans into working memory at an early stage, the brain creates a dual (visual-motor) memory code that can make memories more effective and robust for serving ensuing behavior.

It is believed that children have difficulties in guiding attention while facing distraction. However, developmental accounts of spatial attention rely on traditional search designs using static displays. In real life, dynamic environments can embed regularities that afford anticipation and benefit performance. We developed a dynamic visual-search task to test the ability of children to benefit from spatio-temporal regularities to detect goal-relevant targets appearing within an extended dynamic context amidst irrelevant distracting stimuli. We compared children and adults in detecting predictable vs. unpredictable targets fading in and out among competing distracting stimuli. While overall search performance was poorer in children, both groups detected more predictable targets. This effect was confined to task-relevant information. Additionally, we report how predictions are related to individual differences in attention. Altogether, our results indicate a striking capacity of prediction-led guidance towards task-relevant information in dynamic environments, refining traditional views about poor goal-driven attention in childhood.

In everyday life, attentional templates-which facilitate the perception of task-relevant sensory inputs-are often based on associations in long-term memory. We ask whether templates retrieved from memory are necessarily faithful reproductions of the encoded information or if associative-memory templates can be functionally adapted after retrieval in service of current task demands. Participants learned associations between four shapes and four colored gratings, each with a characteristic combination of color (green or pink) and orientation (left or right tilt). On each trial, observers saw one shape followed by a grating and indicated whether the pair matched the learned shape-grating association. Across experimental blocks, we manipulated the types of nonmatch (lure) gratings most often presented. In some blocks the lures were most likely to differ in color but not tilt, whereas in other blocks this was reversed. If participants functionally adapt the retrieved template such that the distinguishing information between lures and targets is prioritized, then they should overemphasize the most commonly diagnostic feature dimension within the template. We found evidence for this in the behavioral responses to the lures: participants were more accurate and faster when responding to common versus rare lures, as predicted by the functional-but not the strictly veridical-template hypothesis. This shows that templates retrieved from memory can be functionally biased to optimize task performance in a flexible, context-dependent, manner.

Temporal expectations enable anticipatory brain states that prepare us for upcoming perception and action. We investigated the purpose-dependent nature and consequences of cued temporal expectations on brain and behavior in male and female human volunteers, using two matched visual-motor tasks that stressed either response speed or visual accuracy. We show that the consequences of temporal expectations are fundamentally purpose dependent. Temporal expectations predominantly affected response times when visual demands were low and speed was more important, but perceptual accuracy when visual demands were more challenging. Using magnetoencephalography, we further show how temporal expectations latch onto anticipatory neural states associated with concurrent spatial expectations—modulating task-specific anticipatory neural lateralization of oscillatory brain activity in a modality- and frequency-specific manner. By relating these brain states to behavior, we finally reveal how the behavioral relevance of such anticipatory brain states is similarly purpose dependent.SIGNIFICANCE STATEMENTKnowing when events may occur helps to prepare neural activity for upcoming perception and action. It is becoming increasingly clear that distinct sources of temporal expectations may facilitate performance via distinct mechanisms. Another relevant dimension to consider regards the distinct purposes that temporal expectations may serve. Here, we demonstrate that the consequences of temporal expectations on neurophysiological brain activity and behavior are fundamentally purpose dependent, and show how temporal expectations interact with task-relevant neural states in a modality- and frequency-specific manner. This brings the important insight that the ways in which temporal expectations influence brain and behavior, and how brain activity is related to behavior, are not fixed properties but rather depend on the task at hand.