Large-scale interactions in predictive processing: oscillatory versus transient dynamics.

How do the two main types of neural dynamics, aperiodic transients and oscillations, contribute to the interactions between feedforward (FF) and feedback (FB) pathways in sensory inference and predictive processing? We discuss three theoretical perspectives. First, we critically evaluate the theory that gamma and alpha/beta rhythms play a role in classic hierarchical predictive coding (HPC) by mediating FF and FB communication, respectively. Second, we outline an alternative functional model in which rapid sensory inference is mediated by aperiodic transients, whereas oscillations contribute to the stabilization of neural representations over time and plasticity processes. Third, we propose that the strong dependence of oscillations on predictability can be explained based on a biologically plausible alternative to classic HPC, namely dendritic HPC.

Minimal exposure durations reveal visual processing priorities for different stimulus attributes.

Human vision can detect a single photon, but the minimal exposure required to extract meaning from stimulation remains unknown. This requirement cannot be characterised by stimulus energy, because the system is differentially sensitive to attributes defined by configuration rather than physical amplitude. Determining minimal exposure durations required for processing various stimulus attributes can thus reveal the system's priorities. Using a tachistoscope enabling arbitrarily brief displays, we establish minimal durations for processing human faces, a stimulus category whose perception is associated with several well-characterised behavioural and neural markers. Neural and psychophysical measures show a sequence of distinct minimal exposures for stimulation detection, object-level detection, face-specific processing, and emotion-specific processing. Resolving ongoing debates, face orientation affects minimal exposure but emotional expression does not. Awareness emerges with detection, showing no evidence of subliminal perception. These findings inform theories of visual processing and awareness, elucidating the information to which the visual system is attuned.

Breathwork-induced psychedelic experiences modulate neural dynamics.

Breathwork is an understudied school of practices involving intentional respiratory modulation to induce an altered state of consciousness (ASC). We simultaneously investigate the phenomenological and neural dynamics of breathwork by combining Temporal Experience Tracing, a quantitative methodology that preserves the temporal dynamics of subjective experience, with low-density portable EEG devices. Fourteen novice participants completed a course of up to 28 breathwork sessions-of 20, 40, or 60 min-in 28 days, yielding a neurophenomenological dataset of 301 breathwork sessions. Using hypothesis-driven and data-driven approaches, we found that "psychedelic-like" subjective experiences were associated with increased neural Lempel-Ziv complexity during breathwork. Exploratory analyses showed that the aperiodic exponent of the power spectral density-but not oscillatory alpha power-yielded similar neurophenomenological associations. Non-linear neural features, like complexity and the aperiodic exponent, neurally map both a multidimensional data-driven composite of positive experiences, and hypothesis-driven aspects of psychedelic-like experience states such as high bliss.

Staying in control: Characterizing the mechanisms underlying cognitive control in high and low arousal states.

Throughout the day, humans show natural fluctuations in arousal that impact cognitive function. To study the behavioural dynamics of cognitive control during high and low arousal states, healthy participants performed an auditory conflict task during high-intensity physical exercise (N = 39) or drowsiness (N = 33). In line with the pre-registered hypotheses, conflict and conflict adaptation effects were preserved during both altered arousal states. Overall task performance was markedly poorer during low arousal, but not for high arousal. Modelling behavioural dynamics with drift diffusion analysis revealed evidence accumulation and non-decision time decelerated, and decisional boundaries became wider during low arousal, whereas high arousal was unexpectedly associated with a decrease in the interference of task-irrelevant information processing. These findings show how arousal differentially modulates cognitive control at both sides of normal alertness, and further validate drowsiness and physical exercise as key experimental models to disentangle the interaction between physiological fluctuations on cognitive dynamics.

Distributed representations of prediction error signals across the cortical hierarchy are synergistic.

A relevant question concerning inter-areal communication in the cortex is whether these interactions are synergistic. Synergy refers to the complementary effect of multiple brain signals conveying more information than the sum of each isolated signal. Redundancy, on the other hand, refers to the common information shared between brain signals. Here, we dissociated cortical interactions encoding complementary information (synergy) from those sharing common information (redundancy) during prediction error (PE) processing. We analyzed auditory and frontal electrocorticography (ECoG) signals in five common awake marmosets performing two distinct auditory oddball tasks and investigated to what extent event-related potentials (ERP) and broadband (BB) dynamics encoded synergistic and redundant information about PE processing. The information conveyed by ERPs and BB signals was synergistic even at lower stages of the hierarchy in the auditory cortex and between auditory and frontal regions. Using a brain-constrained neural network, we simulated the synergy and redundancy observed in the experimental results and demonstrated that the emergence of synergy between auditory and frontal regions requires the presence of strong, long-distance, feedback, and feedforward connections. These results indicate that distributed representations of PE signals across the cortical hierarchy can be highly synergistic.

Slow-wave brain connectivity predicts executive functioning and group belonging in socially vulnerable individuals.

Important efforts have been made to describe the neural and cognitive features of healthy and clinical populations. However, the neural and cognitive features of socially vulnerable individuals remain largely unexplored, despite their proneness to developing neurocognitive disorders. Socially vulnerable individuals can be characterised as socially deprived, having a low socioeconomic status, suffering from chronic social stress, and exhibiting poor social adaptation. While it is known that such individuals are likely to perform worse than their peers on executive function tasks, studies on healthy but socially vulnerable groups are lacking. In the current study, we explore whether neural power and connectivity signatures can characterise executive function performance in healthy but socially vulnerable individuals, shedding light on the impairing effects that chronic stress and social disadvantages have on cognition. We measured resting-state electroencephalography and executive functioning in 38 socially vulnerable participants and 38 matched control participants. Our findings indicate that while neural power was uninformative, lower delta and theta phase synchrony are associated with worse executive function performance in all participants, whereas delta phase synchrony is higher in the socially vulnerable group compared to the control group. Finally, we found that delta phase synchrony and years of schooling are the best predictors for belonging to the socially vulnerable group. Overall, these findings suggest that exposure to chronic stress due to socioeconomic factors and a lack of education are associated with changes in slow-wave neural connectivity and executive functioning.

Feedback information transfer in the human brain reflects bistable perception in the absence of report.

In the search for the neural basis of conscious experience, perception and the cognitive processes associated with reporting perception are typically confounded as neural activity is recorded while participants explicitly report what they experience. Here, we present a novel way to disentangle perception from report using eye movement analysis techniques based on convolutional neural networks and neurodynamical analyses based on information theory. We use a bistable visual stimulus that instantiates two well-known properties of conscious perception: integration and differentiation. At any given moment, observers either perceive the stimulus as one integrated unitary object or as two differentiated objects that are clearly distinct from each other. Using electroencephalography, we show that measures of integration and differentiation based on information theory closely follow participants' perceptual experience of those contents when switches were reported. We observed increased information integration between anterior to posterior electrodes (front to back) prior to a switch to the integrated percept, and higher information differentiation of anterior signals leading up to reporting the differentiated percept. Crucially, information integration was closely linked to perception and even observed in a no-report condition when perceptual transitions were inferred from eye movements alone. In contrast, the link between neural differentiation and perception was observed solely in the active report condition. Our results, therefore, suggest that perception and the processes associated with report require distinct amounts of anterior-posterior network communication and anterior information differentiation. While front-to-back directed information is associated with changes in the content of perception when viewing bistable visual stimuli, regardless of report, frontal information differentiation was absent in the no-report condition and therefore is not directly linked to perception per se.

Principles of large-scale neural interactions.

What mechanisms underlie flexible inter-areal communication in the cortex? We consider four mechanisms for temporal coordination and their contributions to communication: (1) Oscillatory synchronization (communication-through-coherence); (2) communication-through-resonance; (3) non-linear integration; and (4) linear signal transmission (coherence-through-communication). We discuss major challenges for communication-through-coherence based on layer- and cell-type-specific analyses of spike phase-locking, heterogeneity of dynamics across networks and states, and computational models for selective communication. We argue that resonance and non-linear integration are viable alternative mechanisms that facilitate computation and selective communication in recurrent networks. Finally, we consider communication in relation to cortical hierarchy and critically examine the hypothesis that feedforward and feedback communication use fast (gamma) and slow (alpha/beta) frequencies, respectively. Instead, we propose that feedforward propagation of prediction errors relies on the non-linear amplification of aperiodic transients, whereas gamma and beta rhythms represent rhythmic equilibrium states that facilitate sustained and efficient information encoding and amplification of short-range feedback via resonance.

Cortical, muscular, and kinetic activity underpinning attentional focus strategies during visuomotor control.

Focusing internally on movement control or bodily sensations is frequently shown to disrupt the effectiveness and efficiency of motor control when compared to focusing externally on the outcome of movement. Whilst the behavioral consequences of these attentional strategies are well-documented, it is unclear how they are explained at the corticomuscular level. The aim of the present study was to investigate how attentional focus strategies affect kinetic, cortical, muscular, and corticomuscular activity during an isometric force precision task. In a repeated measures design, we measured force, EEG and EMG activity from twenty-seven participants who performed isometric contractions of the right hand whilst encouraged to adopt either an internal or external focus through a combination of instructions, secondary tasks, and self-report evaluations. Results indicated that focusing internally led to poorer force accuracy and steadiness compared to an external focus. An internal focus also increased muscle activity of the forearm flexor, increased EEG alpha activity across the parieto-occipital cortex, lowered frontal midline EEG theta activity, and lowered beta corticomuscular coherence between the forearm flexor and contralateral motor cortex. The results of this study provide a holistic understanding of how attentional focus strategies alter corticomuscular control during an isometric force precision task, paving the way for exploring how the behavioral consequences of attentional strategies can be explained at the corticomuscular levels across a wide range of motor tasks and contexts.

Ketamine and sleep modulate neural complexity dynamics in cats.

There is increasing evidence that the level of consciousness can be captured by neural informational complexity: for instance, complexity, as measured by the Lempel Ziv (LZ) compression algorithm, decreases during anaesthesia and non-rapid eye movement (NREM) sleep in humans and rats, when compared with LZ in awake and REM sleep. In contrast, LZ is higher in humans under the effect of psychedelics, including subanaesthetic doses of ketamine. However, it is both unclear how this result would be modulated by varying ketamine doses, and whether it would extend to other species. Here, we studied LZ with and without auditory stimulation during wakefulness and different sleep stages in five cats implanted with intracranial electrodes, as well as under subanaesthetic doses of ketamine (5, 10, and 15 mg/kg i.m.). In line with previous results, LZ was lowest in NREM sleep, but similar in REM and wakefulness. Furthermore, we found an inverted U-shaped curve following different levels of ketamine doses in a subset of electrodes, primarily in prefrontal cortex. However, it is worth noting that the variability in the ketamine dose-response curve across cats and cortices was larger than that in the sleep-stage data, highlighting the differential local dynamics created by two different ways of modulating conscious state. These results replicate previous findings, both in humans and other species, demonstrating that neural complexity is highly sensitive to capture state changes between wake and sleep stages while adding a local cortical description. Finally, this study describes the differential effects of ketamine doses, replicating a rise in complexity for low doses, and further fall as doses approach anaesthetic levels in a differential manner depending on the cortex.

Affective modulation of cognitive control: A systematic review of EEG studies.

In recent years, a growing corpus of research has been conducted utilizing a variety of behavioral and neurophysiological methodologies to investigate the relationship of emotion and cognition, yielding unique insights into fundamental concerns about the human mind and mental disease. Electroencephalography (EEG) has been utilized to investigate how emotional states alter neural markers of cognitive control. The current study is a systematic analysis of EEG research that looks at affective modulation (mood, emotion) of cognitive control and its many sub-processes (e.g., cognitive flexibility, inhibitory control, and working memory). The PRISMA standards were followed in this review, which looked at experimental designs and tasks, as well as methodological elements of EEG recording and analysis across research. A total of 35 articles were chosen for qualitative synthesis as a consequence of the search. The examination of event-related potentials (ERPs), which showed affective modulation of 19 different components, was the most common electrophysiological approach used across research. The majority of the investigations focused on N2 and P3, indicating that affective induction has a strong influence on attentional processes and response inhibition. Future research should look into different methodologies such as source location and connection metrics to better understand the brain's areas and dynamic response during affective induction activities. It is also suggested that the technical components of the report be more explicit in order to promote study comparability and replication.

Broadband Dynamics Rather than Frequency-Specific Rhythms Underlie Prediction Error in the Primate Auditory Cortex.

Detection of statistical irregularities, measured as a prediction error response, is fundamental to the perceptual monitoring of the environment. We studied whether prediction error response is associated with neural oscillations or asynchronous broadband activity. Electrocorticography was conducted in three male monkeys, who passively listened to the auditory roving oddball stimuli. Local field potentials (LFPs) recorded over the auditory cortex underwent spectral principal component analysis, which decoupled broadband and rhythmic components of the LFP signal. We found that the broadband component captured the prediction error response, whereas none of the rhythmic components were associated with statistical irregularities of sounds. The broadband component displayed more stochastic, asymmetrical multifractal properties than the rhythmic components, which revealed more self-similar dynamics. We thus conclude that the prediction error response is captured by neuronal populations generating asynchronous broadband activity, defined by irregular dynamic states, which, unlike oscillatory rhythms, appear to enable the neural representation of auditory prediction error response.SIGNIFICANCE STATEMENT This study aimed to examine the contribution of oscillatory and asynchronous components of auditory local field potentials in the generation of prediction error responses to sensory irregularities, as this has not been directly addressed in the previous studies. Here, we show that mismatch negativity-an auditory prediction error response-is driven by the asynchronous broadband component of potentials recorded in the auditory cortex. This finding highlights the importance of nonoscillatory neural processes in the predictive monitoring of the environment. At a more general level, the study demonstrates that stochastic neural processes, which are often disregarded as neural noise, do have a functional role in the processing of sensory information.

Preserved sensory processing but hampered conflict detection when stimulus input is task-irrelevant.

Conflict detection in sensory input is central to adaptive human behavior. Perhaps unsurprisingly, past research has shown that conflict may even be detected in the absence of conflict awareness, suggesting that conflict detection is an automatic process that does not require attention. To test the possibility of conflict processing in the absence of attention, we manipulated task relevance and response overlap of potentially conflicting stimulus features across six behavioral tasks. Multivariate analyses on human electroencephalographic data revealed neural signatures of conflict only when at least one feature of a conflicting stimulus was attended, regardless of whether that feature was part of the conflict, or overlaps with the response. In contrast, neural signatures of basic sensory processes were present even when a stimulus was completely unattended. These data reveal an attentional bottleneck at the level of objects, suggesting that object-based attention is a prerequisite for cognitive control operations involved in conflict detection.

Focusing your attention on one thing can leave you surprisingly unaware of what goes on around you. A classic experiment known as 'the invisible gorilla' highlights this phenomenon. Volunteers were asked to watch a clip featuring basketball players, and count how often those wearing white shirts passed the ball: around half of participants failed to spot that someone wearing a gorilla costume wandered into the game and spent nine seconds on screen. Yet, things that you are not focusing on can sometimes grab your attention anyway. Take for example, the 'cocktail party effect', the ability to hear your name among the murmur of a crowded room. So why can we react to our own names, but fail to spot the gorilla? To help answer this question, Nuiten et al. examined how paying attention affects the way the brain processes input. Healthy volunteers were asked to perform various tasks while the words 'left' or 'right' played through speakers. The content of the word was sometimes consistent with its location ('left' being played on the left speaker), and sometimes opposite ('left' being played on the right speaker). Processing either the content or the location of the word is relatively simple for the brain; however detecting a discrepancy between these two properties is challenging, requiring the information to be processed in a brain region that monitors conflict in sensory input. To manipulate whether the volunteers needed to pay attention to the words, Nuiten et al. made their content or location either relevant or irrelevant for a task. By analyzing brain activity and task performance, they were able to study the effects of attention on how the word properties were processed. The results showed that the volunteers' brains were capable of dealing with basic information, such as location or content, even when their attention was directed elsewhere. But discrepancies between content and location could only be detected when the volunteers were focusing on the words, or when their content or location was directly relevant to the task. The findings by Nuiten et al. suggest that while performing a difficult task, our brains continue to react to basic input but often fail to process more complex information. This, in turn, has implications for a range of human activities such as driving. New technology could potentially help to counteract this phenomenon, aiming to direct attention towards complex information that might otherwise be missed.

Beyond imagination: Hypnotic visual hallucination induces greater lateralised brain activity than visual mental imagery.

Hypnotic suggestions can produce a broad range of perceptual experiences, including hallucinations. Visual hypnotic hallucinations differ in many ways from regular mental images. For example, they are usually experienced as automatic, vivid, and real images, typically compromising the sense of reality. While both hypnotic hallucination and mental imagery are believed to mainly rely on the activation of the visual cortex via top-down mechanisms, it is unknown how they differ in the neural processes they engage. Here we used an adaptation paradigm to test and compare top-down processing between hypnotic hallucination, mental imagery, and visual perception in very highly hypnotisable individuals whose ability to hallucinate was assessed. By measuring the N170/VPP event-related complex and using multivariate decoding analysis, we found that hypnotic hallucination of faces involves greater top-down activation of sensory processing through lateralised neural mechanisms in the right hemisphere compared to mental imagery. Our findings suggest that the neural signatures that distinguish hypnotically hallucinated faces from imagined faces lie in the right brain hemisphere.

In your phase: neural phase synchronisation underlies visual imagery of faces.

Mental imagery is the process through which we retrieve and recombine information from our memory to elicit the subjective impression of "seeing with the mind's eye". In the social domain, we imagine other individuals while recalling our encounters with them or modelling alternative social interactions in future. Many studies using imaging and neurophysiological techniques have shown several similarities in brain activity between visual imagery and visual perception, and have identified frontoparietal, occipital and temporal neural components of visual imagery. However, the neural connectivity between these regions during visual imagery of socially relevant stimuli has not been studied. Here we used electroencephalography to investigate neural connectivity and its dynamics between frontal, parietal, occipital and temporal electrodes during visual imagery of faces. We found that voluntary visual imagery of faces is associated with long-range phase synchronisation in the gamma frequency range between frontoparietal electrode pairs and between occipitoparietal electrode pairs. In contrast, no effect of imagery was observed in the connectivity between occipitotemporal electrode pairs. Gamma range synchronisation between occipitoparietal electrode pairs predicted subjective ratings of the contour definition of imagined faces. Furthermore, we found that visual imagery of faces is associated with an increase of short-range frontal synchronisation in the theta frequency range, which temporally preceded the long-range increase in the gamma synchronisation. We speculate that the local frontal synchrony in the theta frequency range might be associated with an effortful top-down mnemonic reactivation of faces. In contrast, the long-range connectivity in the gamma frequency range along the fronto-parieto-occipital axis might be related to the endogenous binding and subjective clarity of facial visual features.

Decreased Alertness Reconfigures Cognitive Control Networks.

Humans' remarkable capacity to flexibly adapt their behavior based on rapid situational changes is termed cognitive control. Intuitively, cognitive control is thought to be affected by the state of alertness; for example, when drowsy, we feel less capable of adequately implementing effortful cognitive tasks. Although scientific investigations have focused on the effects of sleep deprivation and circadian time, little is known about how natural daily fluctuations in alertness in the regular awake state affect cognitive control. Here we combined a conflict task in the auditory domain with EEG neurodynamics to test how neural and behavioral markers of conflict processing are affected by fluctuations in alertness. Using a novel computational method, we segregated alert and drowsy trials from two testing sessions and observed that, although participants (both sexes) were generally sluggish, the typical conflict effect reflected in slower responses to conflicting information compared with nonconflicting information, as well as the moderating effect of previous conflict (conflict adaptation), were still intact. However, the typical neural markers of cognitive control-local midfrontal theta-band power changes-that participants show during full alertness were no longer noticeable when alertness decreased. Instead, when drowsy, we found an increase in long-range information sharing (connectivity) between brain regions in the same frequency band. These results show the resilience of the human cognitive control system when affected by internal fluctuations of alertness and suggest that there are neural compensatory mechanisms at play in response to physiological pressure during diminished alertness.SIGNIFICANCE STATEMENT The normal variability in alertness we experience in daily tasks is rarely taken into account in cognitive neuroscience. Here we studied neurobehavioral dynamics of cognitive control with decreasing alertness. We used the classic Simon task where participants hear the word "left" or "right" in the right or left ear, eliciting slower responses when the word and the side are incongruent-the conflict effect. Participants performed the task both while fully awake and while getting drowsy, allowing for the characterization of alertness modulating cognitive control. The changes in the neural signatures of conflict from local theta oscillations to a long-distance distributed theta network suggest a reconfiguration of the underlying neural processes subserving cognitive control when affected by alertness fluctuations.

Alertness fluctuations when performing a task modulate cortical evoked responses to transcranial magnetic stimulation.

Transcranial magnetic stimulation (TMS) has been widely used in human cognitive neuroscience to examine the causal role of distinct cortical areas in perceptual, cognitive and motor functions. However, it is widely acknowledged that the effects of focal cortical stimulation can vary substantially between participants and even from trial to trial within individuals. Recent work from resting state functional magnetic resonance imaging (fMRI) studies has suggested that spontaneous fluctuations in alertness over a testing session can modulate the neural dynamics of cortical processing, even when participants remain awake and responsive to the task at hand. Here we investigated the extent to which spontaneous fluctuations in alertness during wake-to-sleep transition can account for the variability in neurophysiological responses to TMS. We combined single-pulse TMS with neural recording via electroencephalography (EEG) to quantify changes in motor and cortical reactivity with fluctuating levels of alertness defined objectively on the basis of ongoing brain activity. We observed rapid, non-linear changes in TMS-evoked responses with decreasing levels of alertness, even while participants remained responsive in the behavioural task. Specifically, we found that the amplitude of motor evoked potentials peaked during periods of EEG flattening, whereas TMS-evoked potentials increased and remained stable during EEG flattening and the subsequent occurrence of theta ripples that indicate the onset of NREM stage 1 sleep. Our findings suggest a rapid and complex reorganization of active neural networks in response to spontaneous fluctuations of alertness over relatively short periods of behavioural testing during wake-to-sleep transition.

Dissociable Neural Information Dynamics of Perceptual Integration and Differentiation during Bistable Perception.

At any given moment, we experience a perceptual scene as a single whole and yet we may distinguish a variety of objects within it. This phenomenon instantiates two properties of conscious perception: integration and differentiation. Integration is the property of experiencing a collection of objects as a unitary percept and differentiation is the property of experiencing these objects as distinct from each other. Here, we evaluated the neural information dynamics underlying integration and differentiation of perceptual contents during bistable perception. Participants listened to a sequence of tones (auditory bistable stimuli) experienced either as a single stream (perceptual integration) or as two parallel streams (perceptual differentiation) of sounds. We computed neurophysiological indices of information integration and information differentiation with electroencephalographic and intracranial recordings. When perceptual alternations were endogenously driven, the integrated percept was associated with an increase in neural information integration and a decrease in neural differentiation across frontoparietal regions, whereas the opposite pattern was observed for the differentiated percept. However, when perception was exogenously driven by a change in the sound stream (no bistability), neural oscillatory power distinguished between percepts but information measures did not. We demonstrate that perceptual integration and differentiation can be mapped to theoretically motivated neural information signatures, suggesting a direct relationship between phenomenology and neurophysiology.