The effects of physical activity on sleep architecture and mood in naturalistic environments.

Physical activity has been found to alter sleep architecture, but these effects have been studied predominantly in the laboratory and the generalizability of these findings to naturalistic environments and longer time intervals, as well as their psychological effects, have not been evaluated. Recent technological advancements in wearable devices have made it possible to capture detailed measures of sleep outside the lab, including timing of specific sleep stages. In the current study, we utilized photoplethysmography coupled with accelerometers and smartphone ambulatory assessment to collect daily measurements of sleep, physical activity and mood in a sample of N = 82 over multi-month data collection intervals. We found a robust inverse relationship between sedentary behavior and physical activity and sleep architecture: both low-intensity and moderate-to-vigorous physical activity were associated with increased NREM sleep and decreased REM sleep, as well as a longer REM latency, while higher levels of sedentary behavior showed the opposite pattern. A decreased REM/NREM ratio and increased REM latency were in turn associated with improved wellbeing, including increased energy, reduced stress and enhanced perceived restfulness of sleep. Our results suggest that physical activity and sleep account for unique variance in a person's mood, suggesting that these effects are at least partially independent.

Lucid dreaming occurs in activated rapid eye movement sleep, not a mixture of sleep and wakefulness.

STUDY OBJECTIVES: (1) To critically test whether a previously reported increase in frontolateral 40 Hz power in lucid REM sleep, used to justify the claim that lucid dreaming is a "hybrid state" mixing sleep and wakefulness, is attributable to the saccadic spike potential (SP) artifact as a corollary of heightened REM density. (2) To replicate the finding that lucid dreams are associated with physiological activation, including heightened eye movement density, during REM sleep. (3) To conduct an exploratory analysis of changes in EEG features during lucid REM sleep. METHODS: We analyzed 14 signal-verified lucid dreams (SVLDs) and baseline REM sleep segments from the same REM periods from six participants derived from the Stanford SVLD database. Participants marked lucidity onset with standard left-right-left-right-center (LR2c) eye-movement signals in polysomnography recordings. RESULTS: Compared to baseline REM sleep, lucid REM sleep had higher REM density (ß = 0.85, p = 0.002). Bayesian analysis supported the null hypothesis of no differences in frontolateral 40 Hz power after removal of the SP artifact (BH = 0.18) and ICA correction (BH = 0.01). Compared to the entire REM sleep period, lucid REM sleep showed small reductions in low-frequency and beta band spectral power as well as increased signal complexity (all p < 0.05), which were within the normal variance of baseline REM sleep. CONCLUSIONS: Lucid dreams are associated with higher-than-average levels of physiological activation during REM sleep, including measures of both subcortical and cortical activation. Increases in 40 Hz power in periorbital channels reflect saccadic and microsaccadic SPs as a result of higher REM density accompanying heightened activation.

Episodic thought distinguishes spontaneous cognition in waking from REM and NREM sleep.

Evidence suggests continuity between cognition in waking and sleeping states. However, one type of cognition that may differ is episodic thoughts of the past and future. The current study investigated this across waking, NREM sleep and REM sleep. We analyzed thought reports obtained from a large sample of individuals (N = 138) who underwent experience-sampling during wakefulness as well as serial awakenings in sleep. Our data suggest that while episodic thoughts are common during waking spontaneous thought, episodic thoughts of both the past and the future rarely occur in either N2 or REM sleep. Moreover, replicating previous findings, episodic thoughts during wakefulness exhibit a strong prospective bias and frequently involve autobiographical planning. Together, these results suggest that the occurrence of spontaneous episodic thoughts differs substantially across waking and dreaming sleep states. We suggest that this points to a difference in the way that human consciousness is typically experienced across the sleep-wake cycle.

Two-Way Communication in Lucid REM Sleep Dreaming.

Dreamers were long thought absolutely isolated from the outside world. Yet psychophysiological studies over the past 40 years have firmly established that lucid dreamers can use eye movements to report on their dream content in real time while in rapid eye movement (REM) sleep. We now also know that sensory input is not completely suppresssed during sleep. A recent study by Konkoly et al. illustrates how experimenters can question lucid dreamers during ongoing dreams and asks whether more extended two-way communication during lucid REM sleep dreaming is feasible.

Increased lucid dream frequency in long-term meditators but not following MBSR training.

Strong conceptual and theoretical connections have been made between meditation practice, mindfulness and lucid dreaming. However, only a handful of empirical studies have evaluated the relationship between lucid dreaming and meditation, and conclusions remain tempered by methodological limitations. Here we evaluate the relationship between meditation, mindfulness and lucid dream frequency using several complementary methods. First, using a cross-sectional design, we evaluate differences in lucid dream frequency between long-term meditators and meditation naïve individuals. Second, we evaluate the relationship between lucid dream frequency and specific facets of trait mindfulness in both meditators and non-meditators. Third, using a blinded randomized-controlled design, we evaluate the impact of an 8-week mindfulness course on lucid dreaming frequency. Our results show that lucid dreaming is more frequent in long-term meditators compared to meditation naïve individuals. Additionally, lucid dream frequency in meditation-naïve individuals was associated with a capacity to verbalize experience, while lucid dream frequency in long-term meditators was associated with observational and decentering facets of trait mindfulness. However, an 8-week mindfulness course did not increase the frequency of lucid dreams. Together these results support a continuity between increased awareness of waking and sleeping states, provide a novel form of evidence linking meditation training to meta-awareness, and support an association between meditation practice and lucid dreaming, but leave open the specific nature of this connection.

The cognitive neuroscience of lucid dreaming.

Lucid dreaming refers to the phenomenon of becoming aware of the fact that one is dreaming during ongoing sleep. Despite having been physiologically validated for decades, the neurobiology of lucid dreaming is still incompletely characterized. Here we review the neuroscientific literature on lucid dreaming, including electroencephalographic, neuroimaging, brain lesion, pharmacological and brain stimulation studies. Electroencephalographic studies of lucid dreaming are mostly underpowered and show mixed results. Neuroimaging data is scant but preliminary results suggest that prefrontal and parietal regions are involved in lucid dreaming. A focus of research is also to develop methods to induce lucid dreams. Combining training in mental set with cholinergic stimulation has shown promising results, while it remains unclear whether electrical brain stimulation could be used to induce lucid dreams. Finally, we discuss strategies to measure lucid dreaming, including best-practice procedures for the sleep laboratory. Lucid dreaming has clinical and scientific applications, and shows emerging potential as a methodology in the cognitive neuroscience of consciousness. Further research with larger sample sizes and refined methodology is needed.

Connectivity differences between consciousness and unconsciousness in non-rapid eye movement sleep: a TMS-EEG study.

The neuronal connectivity patterns that differentiate consciousness from unconsciousness remain unclear. Previous studies have demonstrated that effective connectivity, as assessed by transcranial magnetic stimulation combined with electroencephalography (TMS-EEG), breaks down during the loss of consciousness. This study investigated changes in EEG connectivity associated with consciousness during non-rapid eye movement (NREM) sleep following parietal TMS. Compared with unconsciousness, conscious experiences during NREM sleep were associated with reduced phase-locking at low frequencies (<4 Hz). Transitivity and clustering coefficient in the delta and theta bands were also significantly lower during consciousness compared to unconsciousness, with differences in the clustering coefficient observed in scalp electrodes over parietal-occipital regions. There were no significant differences in Granger-causality patterns in frontal-to-parietal or parietal-to-frontal connectivity between reported unconsciousness and reported consciousness. Together these results suggest that alterations in spectral and spatial characteristics of network properties in posterior brain areas, in particular decreased local (segregated) connectivity at low frequencies, is a potential indicator of consciousness during sleep.

Graph Theoretical Analysis of Cortical Networks based on Conscious Experience.

The aim of the study was to investigate differences in cortical networks based on the state of consciousness. Five subjects performed a serial-awakening paradigm with electroencephalography (EEG) recordings. We considered four states of consciousness: (1) non-rapid eye movement (NREM) sleep with no conscious experience, (2) NREM sleep with conscious experience, (3) rapid eye movement (REM) sleep with conscious experience, and (4) wakefulness. We applied graph theoretical analysis to explore the cortical connectivity and network properties in five frequency bands. Connectivity between EEG channels was evaluated with the weighted phase lag index (wPLI). The characteristic path length, transitivity, and clustering coefficient were computed to evaluate functional integration and segregation of the associated brain network. There were no significant differences in wPLI among the four states of consciousness. In the beta band, functional integration in wakefulness was higher than in NREM sleep. Regarding functional segregation, in the theta band, transitivity and clustering coefficient in NREM sleep with no conscious experience were stronger than in wakefulness or REM sleep, but clustering in the beta band showed an opposite effect. The observed differences may be related to cortical bistability and add to previously observed neural correlates of consciousness.

Frequent lucid dreaming associated with increased functional connectivity between frontopolar cortex and temporoparietal association areas.

Humans typically lack awareness that they are dreaming while dreaming. However, at times a remarkable exception occurs and reflective consciousness can be regained while dreaming, referred to as lucid dreaming. While most individuals experience lucid dreams rarely there is substantial variance in lucid dream frequency. The neurobiological basis of lucid dreaming is unknown, but evidence points to involvement of anterior prefrontal cortex (aPFC) and parietal cortex. This study evaluated the neuroanatomical/neurofunctional correlates of frequent lucid dreams and specifically whether functional connectivity of aPFC is associated with frequent lucid dreams. We analyzed structural and functional magnetic resonance imaging from an exceptional sample of fourteen individuals who reported ≥3 lucid dreams/week and a control group matched on age, gender and dream recall that reported ≤1 lucid dream/year. Compared to controls, the frequent lucid dream group showed significantly increased resting-state functional connectivity between left aPFC and bilateral angular gyrus, bilateral middle temporal gyrus and right inferior frontal gyrus, and higher node degree and strength in left aPFC. In contrast, no significant differences in brain structure were observed. Our results suggest that frequent lucid dreaming is associated with increased functional connectivity between aPFC and temporoparietal association areas, regions normally deactivated during sleep.

Human Rapid Eye Movement Sleep Shows Local Increases in Low-Frequency Oscillations and Global Decreases in High-Frequency Oscillations Compared to Resting Wakefulness.

It is often assumed that during rapid eye movement (REM) sleep the cerebral cortex homogenously shows electroencephalogram (EEG) activity highly similar to wakefulness. However, to date no studies have compared neural oscillatory activity in human REM sleep to resting wakefulness with high spatial sampling. In the current study, we evaluated high-resolution topographical changes in neural oscillatory power between both early and late naturalistic REM sleep and resting wakefulness in adult humans. All-night recordings with 256-channel high-density EEG (hd-EEG) were collected in healthy volunteers (N = 12). Topographic analysis revealed that, compared to wake, both the first and last cycle of REM sleep were associated with increased low-frequency oscillations in local central and occipital regions. In contrast, high-frequency activity in both α and ß bands (8-20 Hz) was globally decreased during both early and late REM sleep cycles compared to wakefulness. No significant differences in topographic power in any frequency band were observed between REM sleep cycles occurring early and late in the night. We replicated these findings in an independent dataset (N = 33). Together, our findings show that human REM sleep shows consistent topographical changes in oscillatory power across both early and late sleep cycles compared to resting wakefulness.

Smooth tracking of visual targets distinguishes lucid REM sleep dreaming and waking perception from imagination.

Humans are typically unable to engage in sustained smooth pursuit for imagined objects. However, it is unknown to what extent smooth tracking occurs for visual imagery during REM sleep dreaming. Here we examine smooth pursuit eye movements during tracking of a slow-moving visual target during lucid dreams in REM sleep. Highly similar smooth pursuit tracking was observed during both waking perception and lucid REM sleep dreaming, in contrast to the characteristically saccadic tracking observed during visuomotor imagination. Our findings suggest that, in this respect, the visual imagery that occurs during REM sleep is more similar to perception than imagination. The data also show that the neural circuitry of smooth pursuit can be driven by a visual percept in the absence of retinal stimulation and that specific voluntary shifts in the direction of experienced gaze within REM sleep dreams are accompanied by corresponding rotations of the physical eyes.

Pre-sleep treatment with galantamine stimulates lucid dreaming: A double-blind, placebo-controlled, crossover study.

Lucid dreaming is a remarkable state of consciousness in which one is aware of the fact that one is dreaming while continuing to dream. Based on the strong relationship between physiological activation during rapid eye-movement sleep and lucid dreaming, our pilot research investigated whether enhancing cortical activation via acetylcholinesterease inhibition (AChEI) would increase the frequency of lucid dreams and found AChEI to be a promising method for lucid dream induction. In the current study we sought to quantify the size and reliability of the effect of AChEI on lucid dreaming, dream recall and dream content as well as to test the effectiveness of an integrated lucid dream induction protocol which combined cholinergic stimulation with other methods for lucid dream induction. Participants (N = 121) with high dream recall and an interest in lucid dreaming were randomly assigned counterbalanced orders of 3 doses of galantamine (0, 4 and 8 mg). On 3 consecutive nights, they awoke approximately 4.5 hours after lights out, recalled a dream, ingested the capsules and stayed out of bed for at least 30 minutes. Participants then returned to bed and practiced the Mnemonic Induction of Lucid Dreams technique while returning to sleep. The percentage of participants who reported a lucid dream was significantly increased for both 4 mg (27%, odds ratio = 2.29) and 8 mg doses (42%, odds ratio = 4.46) compared to the active placebo procedure (14%). Galantamine also significantly increased dream recall, sensory vividness and complexity (p<0.05). Dream recall, cognitive clarity, control, positive emotion, vividness and self-reflection were increased during lucid compared to non-lucid dreams (p<0.0001). These results show that galantamine increases the frequency of lucid dreams in a dose-related manner. Furthermore, the integrated method of taking galantamine in the last third of the night with at least 30 minutes of sleep interruption and with an appropriately focused mental set is one of the most effective methods for inducing lucid dreams available today.

The Energy Landscape of Neurophysiological Activity Implicit in Brain Network Structure.

A critical mystery in neuroscience lies in determining how anatomical structure impacts the complex functional dynamics of the brain. How does large-scale brain circuitry constrain states of neuronal activity and transitions between those states? We address these questions using a maximum entropy model of brain dynamics informed by white matter tractography. We demonstrate that the most probable brain states - characterized by minimal energy - display common activation profiles across brain areas: local spatially-contiguous sets of brain regions reminiscent of cognitive systems are co-activated frequently. The predicted activation rate of these systems is highly correlated with the observed activation rate measured in a separate resting state fMRI data set, validating the utility of the maximum entropy model in describing neurophysiological dynamics. This approach also offers a formal notion of the energy of activity within a system, and the energy of activity shared between systems. We observe that within- and between-system energies cleanly separate cognitive systems into distinct categories, optimized for differential contributions to integrated versus segregated function. These results support the notion that energetic and structural constraints circumscribe brain dynamics, offering insights into the roles that cognitive systems play in driving whole-brain activation patterns.

The Phenomenal Contents and Neural Correlates of Spontaneous Thoughts across Wakefulness, NREM Sleep, and REM Sleep.

Thoughts occur during wake as well as during dreaming sleep. Using experience sampling combined with high-density EEG, we investigated the phenomenal qualities and neural correlates of spontaneously occurring thoughts across wakefulness, non-rapid eye movement (NREM) sleep, and REM sleep. Across all states, thoughts were associated with activation of a region of the midcingulate cortex. Thoughts during wakefulness additionally involved a medial prefrontal region, which was associated with metacognitive thoughts during wake. Phenomenologically, waking thoughts had more metacognitive content than thoughts during both NREM and REM sleep, whereas thoughts during REM sleep had a more social content. Together, these results point to a core neural substrate for thoughts, regardless of behavioral state, within the midcingulate cortex, and suggest that medial prefrontal regions may contribute to metacognitive content in waking thoughts.

The neural correlates of dreaming.

Consciousness never fades during waking. However, when awakened from sleep, we sometimes recall dreams and sometimes recall no experiences. Traditionally, dreaming has been identified with rapid eye-movement (REM) sleep, characterized by wake-like, globally 'activated', high-frequency electroencephalographic activity. However, dreaming also occurs in non-REM (NREM) sleep, characterized by prominent low-frequency activity. This challenges our understanding of the neural correlates of conscious experiences in sleep. Using high-density electroencephalography, we contrasted the presence and absence of dreaming in NREM and REM sleep. In both NREM and REM sleep, reports of dream experience were associated with local decreases in low-frequency activity in posterior cortical regions. High-frequency activity in these regions correlated with specific dream contents. Monitoring this posterior 'hot zone' in real time predicted whether an individual reported dreaming or the absence of dream experiences during NREM sleep, suggesting that it may constitute a core correlate of conscious experiences in sleep.

Consciousness isn't all-or-none: Evidence for partial awareness during the attentional blink.

Alternative views of the nature of consciousness posit that awareness of an object is either an all-or-none phenomenon or that awareness can be partial, occurring independently for different levels of representation. The all-or-none hypothesis predicts that when one feature of an object is identified, all other features should be consciously accessible. The partial awareness hypothesis predicts that one feature may reach consciousness while others do not. These competing predictions were tested in two experiments that presented two targets within a central stream of letters. We used the attentional blink evoked by the first target to assess consciousness for two different features of the second target. The results provide evidence that there can be a severe impairment in conscious access to one feature even when another feature is accurately reported. This behavioral evidence supports the partial awareness hypothesis, showing that consciousness of different features of the same object can be dissociated.

Regional white matter variation associated with domain-specific metacognitive accuracy.

The neural mechanisms that mediate metacognitive ability (the capacity to accurately reflect on one's own cognition and experience) remain poorly understood. An important question is whether metacognitive capacity is a domain-general skill supported by a core neuroanatomical substrate or whether regionally specific neural structures underlie accurate reflection in different cognitive domains. Providing preliminary support for the latter possibility, recent findings have shown that individual differences in metacognitive ability in the domains of memory and perception are related to variation in distinct gray matter volume and resting-state functional connectivity. The current investigation sought to build on these findings by evaluating how metacognitive ability in these domains is related to variation in white matter microstructure. We quantified metacognitive ability across memory and perception domains and used diffusion spectrum imaging to examine the relation between high-resolution measurements of white matter microstructure and individual differences in metacognitive accuracy in each domain. We found that metacognitive accuracy for perceptual decisions and memory were uncorrelated across individuals and that metacognitive accuracy in each domain was related to variation in white matter microstructure in distinct brain areas. Metacognitive accuracy for perceptual decisions was associated with increased diffusion anisotropy in white matter underlying the ACC, whereas metacognitive accuracy for memory retrieval was associated with increased diffusion anisotropy in the white matter extending into the inferior parietal lobule. Together, these results extend previous findings linking metacognitive ability in the domains of perception and memory to variation in distinct brain structures and connections.

Domain-specific enhancement of metacognitive ability following meditation training.

Contemplative mental practices aim to enable individuals to develop greater awareness of their own cognitive and affective states through repeated examination of first-person experience. Recent cross-sectional studies of long-term meditation practitioners suggest that the subjective reports of such individuals are better calibrated with objective indices; however, the impact of mental training on metacognitive ability has not yet been examined in a randomized controlled investigation. The present study evaluated the impact of a 2-week meditation-training program on introspective accuracy in the domains of perception and memory. Compared with an active control group that elicited no change, we found that a 2-week meditation program significantly enhanced introspective accuracy, quantified by metacognitive judgments of cognition on a trial-by-trial basis, in a memory but not a perception domain. Together, these data suggest that, in at least some domains, the human capacity to introspect is plastic and can be enhanced through training.

The decoupled mind: mind-wandering disrupts cortical phase-locking to perceptual events.

The mind flows in a "stream of consciousness," which often neglects immediate sensory input in favor of focusing on intrinsic, self-generated thoughts or images. Although considerable research has documented the disruptive influences of task-unrelated thought for perceptual processing and task performance, the brain dynamics associated with these phenomena are not well understood. Here we investigate the possibility, suggested by several convergent lines of research, that task-unrelated thought is associated with a reduction in the trial-to-trial phase consistency of the oscillatory neural signal in response to perceptual input. Using an experience sampling paradigm coupled with continuous high-density electroencephalography, we observed that task-unrelated thought was associated with a reduction of the P1 ERP, replicating prior observations that mind-wandering is accompanied by a reduction of the brain-evoked response to sensory input. Time-frequency analysis of the oscillatory neural response revealed a decrease in theta-band cortical phase-locking, which peaked over parietal scalp regions. Furthermore, we observed that task-unrelated thought impacted the oscillatory mode of the brain during the initiation of a task-relevant action, such that more cortical processing was required to meet task demands. Together, these findings document that the attenuation of perceptual processing that occurs during task-unrelated thought is associated with a reduction in the temporal fidelity with which the brain responds to a stimulus and suggest that increased neural processing may be required to recouple attention to a task. More generally, these data provide novel confirmatory evidence for the mechanisms through which attentional states facilitate the neural processing of sensory input.

Thinking one thing, saying another: the behavioral correlates of mind-wandering while reading aloud.

Although mind-wandering during silent reading is well documented, to date no research has investigated whether similar processes occur during reading aloud. In the present study, participants read a passage either silently or aloud while periodically being probed about mind-wandering. Although their comprehension accuracies were similar for both reading conditions, participants reported more mind-wandering while they were reading aloud. These episodes of mindless reading were associated with nearly normal prosody, but were nevertheless distinguished by subtle fluctuations in volume that were predictive of both overall comprehension accuracy and individual sentence comprehension. Together, these findings reveal that previously hidden within the common activity of reading aloud lies: (1) a demonstration of the remarkable automaticity of speech, (2) a situation that is surprisingly conducive to mind-wandering, (3) subtle vocal signatures of mind-wandering and comprehension accuracy, and (4) the promise of developing useful interventions to improve reading.