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Sleep duration, quality, and rest-activity pattern-a measure for inferring circadian rhythm-are influenced by multiple factors including access to electricity. Recent findings suggest that the safety and comfort afforded by technology may improve sleep but negatively impact rest-activity stability. According to the circadian entrainment hypothesis, increased access to electric lighting should lead to weaker and less uniform circadian rhythms, measured by stability of rest-activity patterns. Here, we investigate sleep in a Maya community in Guatemala who are in a transitional stage of industrialization. We predicted that (i) sleep will be shorter and less efficient in this population than in industrial settings, and that (ii) rest-activity patterns will be weaker and less stable than in contexts with greater exposure to the natural environment and stronger and more stable than in settings more buffered by technologic infrastructure. Our results were mixed. Compared to more industrialized settings, in our study population sleep was 4.87% less efficient (78.39% vs 83.26%). We found no significant difference in sleep duration. Rest-activity patterns were more uniform and less variable than in industrial settings (interdaily stability = 0.58 vs 0.43; intradaily variability = 0.53 vs 0.60). Our results suggest that industrialization does not inherently reduce characteristics of sleep quality; instead, the safety and comfort afforded by technological development may improve sleep, and an intermediate degree of environmental exposure and technological buffering may support circadian rhythm strength and stability., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2022 McKinnon et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Characteristics of the sleep-site are thought to influence the quality and duration of primate sleep, yet only a handful of studies have investigated these links experimentally. Using actigraphy and infrared videography, we quantified sleep in four lemur species (Eulemur coronatus, Lemur catta, Propithecus coquereli, and Varecia rubra) under two different experimental conditions at the Duke Lemur Center (DLC) in Durham, NC, USA. Individuals from each species underwent three weeks of simultaneous testing to investigate the hypothesis that comfort level of the sleep-site influences sleep. We obtained baseline data on normal sleep, and then, in a pair-wise study design, we compared the daily sleep times, inter-daily activity stability, and intra-daily activity variability of individuals in simultaneous experiments of sleep-site enrichment and sleep-site impoverishment. Over 164 24-hour periods from 8 individuals (2 of each species), we found evidence that enriched sleep-sites increased daily sleep times of lemurs, with an average increase of thirty-two minutes. The effect of sleep-site impoverishment was small and not statistically significant. Though our experimental manipulations altered inter-daily stability and intra-daily variability in activity patterns relative to baseline, the changes did not differ significantly between enriched and impoverished conditions. We conclude that properties of a sleep-site enhancing softness or insulation, more than the factors of surface area or stability, influence lemur sleep, with implications regarding the importance of nest building in primate evolution and the welfare and management of captive lemurs., Competing Interests: The authors have declared that no competing interests exist.

Primates spend almost half their lives asleep, yet little is known about how sleep influences their waking cognition. We hypothesized that diurnal and cathemeral lemurs differ in their need for consistent, non-segmented sleep for next-day cognitive function-including long-term memory consolidation, self-control, foraging efficiency, and sociality. Specifically, we expected that strictly diurnal Propithecus is more reliant on uninterrupted sleep for cognitive performance, as compared to four other lemur species that are more flexibly active (i.e., cathemeral). We experimentally inhibited sleep and tested next-day performance in 30 individuals of 5 lemur species over 960 total nights at the Duke Lemur Center in Durham, North Carolina. Each set of pair-housed lemurs experienced a sleep restriction and/or deprivation protocol and was subsequently tested in a variety of fitness-relevant cognitive tasks. Within-subject comparisons of performance on these tasks were made by switching the pair from the experimental sleep inhibited condition to a normal sleep environment, thus ensuring cognitive equivalency among individuals. We validated effectiveness of the protocol via actigraphy and infrared videography. Our results suggest that 'normal' non-disrupted sleep improved memory consolidation for all lemurs. Additionally, on nights of normal sleep, diurnal lemurs performed better in foraging efficiency tasks than cathemeral lemurs. Social behaviors changed in species-specific ways after exposure to experimental conditions, and self-control was not significantly linked with sleep condition. Based on these findings, the links between sleep, learning, and memory consolidation appear to be evolutionarily conserved in primates.

Sleep is a critically important dimension of primate behavior, ecology, and evolution, yet primate sleep is under-studied because current methods of analyzing sleep are expensive, invasive, and time-consuming. In contrast to electroencephalography (EEG) and actigraphy, videography is a cost-effective and non-invasive method to study sleep architecture in animals. With video data, however, it is challenging to score subtle changes that occur in different sleep states, and technology has lagged behind innovations in EEG and actigraphy. Here, we applied Eulerian videography to magnify pixels relevant to scoring sleep from video, and then compared these results to analyses based on actigraphy and standard infrared videography. We studied four species of lemurs (Eulemur coronatus, Lemur catta, Propithecus coquereli, Varecia rubra) for 12-h periods per night, resulting in 6480 1-min epochs for analysis. Cramer's V correlation between actigraphy-classified sleep and infrared videography-classified sleep revealed consistent results in eight of the nine 12-h videos scored. A sample of the infrared videography was then processed by Eulerian videography for movement magnification and re-coded. A second Cramer's V correlation analysis, between two independent scorers coding the same Eulerian-processed video, found that interobserver agreement among Eulerian videography increased sleep vs. awake, NREM, and REM classifications by 7.1%, 46.7%, and 34.3%, respectively. Furthermore, Eulerian videography was more strongly correlated with actigraphy data when compared to results from standard infrared videography. The increase in agreement between the two scorers indicates that Eulerian videography has the potential to improve the identification of sleep states in lemurs and other primates, and thus to expand our understanding of sleep architecture without the need for EEG.

Objectives: The lunar cycle is expected to influence sleep-wake patterns in human populations that have greater exposure to the environment, as might be found in forager populations that experience few environmental buffers. We investigated this "moonlight" hypothesis in two African populations: one composed of hunter-gatherers (with minimal environmental buffering) and the other rural agriculturalists (with low-to-moderate environmental buffering)., Setting: Research was conducted on Hadza hunter-gatherers from the Sengele community near Lake Eyasi in northern Tanzania and in Mandena, Madagascar, in a rural community of approximately 4000 farmers., Participants: Thirty-one adult Hadza and 21 Malagasy adults were recruited., Measurements: We used the CamNtech Motionwatch 8 actigraph and generated data on an epoch-by-epoch, 1-minute basis., Results: In general support of the moonlight hypothesis, we uncovered an association between sleep-wake patterns and lunar cycle (ie., moonlight) for Hadza hunter-gatherers. However, the direction of the effect was opposite to what we predicted: as the potential for exposure to moonlight increased, activity generally shifted to a pattern of less nighttime activity and greater daytime activity. No significant effects were found in the Malagasy agriculturalists., Conclusions: The proposal that human behaviors are linked with moon phase is a popular belief that persists despite the absence of consistent evidence. We provide the first direct evidence that lunar cycle is linked to sleep-wake pattern in a hunter-gatherer society, suggesting that moonlight does not inhibit sleep-wake patterns in the ways that electric lighting does., (Copyright © 2018 National Sleep Foundation. Published by Elsevier Inc. All rights reserved.)

Objectives: Primates vary in their sleep durations and, remarkably, humans sleep the least per 24-hr period of the 30 primates that have been studied. Using phylogenetic methods that quantitatively situate human phenotypes within a broader primate comparative context, we investigated the evolution of human sleep architecture, focusing on: total sleep duration, rapid eye movement (REM) sleep duration, non-rapid eye movement (NREM) sleep duration, and proportion of sleep in REM., Materials and Methods: We used two different Bayesian methods: phylogenetic prediction based on phylogenetic generalized least squares and a multistate Onrstein-Uhlenbeck (OU) evolutionary model of random drift and stabilizing selection., Results: Phylogenetic prediction confirmed that humans sleep less than predicted for a primate of our body mass, predation risk, brain size, foraging needs, sexual selection, and diet. These analyses further revealed that humans pack an unexpectedly higher proportion of REM sleep within a shorter overall sleep duration, and do so by reducing NREM sleep (rather than increasing REM). The OU model generally confirmed these findings, with shifts along the human lineage inferred for TST, NREM, and proportion of REM, but not for REM., Discussion: We propose that the risks and opportunity costs of sleep are responsible for shorter sleep durations in humans, with risks arising from terrestrial sleep involving threats from predators and conspecifics, and opportunity costs because time spent sleeping could be used for learning, creating material objects, and socializing., (© 2018 Wiley Periodicals, Inc.)

Sleep is essential for survival, yet it also represents a time of extreme vulnerability to predation, hostile conspecifics and environmental dangers. To reduce the risks of sleeping, the sentinel hypothesis proposes that group-living animals share the task of vigilance during sleep, with some individuals sleeping while others are awake. To investigate sentinel-like behaviour in sleeping humans, we investigated activity patterns at night among Hadza hunter-gatherers of Tanzania. Using actigraphy, we discovered that all subjects were simultaneously scored as asleep for only 18 min in total over 20 days of observation, with a median of eight individuals awake throughout the night-time period; thus, one or more individuals was awake (or in light stages of sleep) during 99.8% of sampled epochs between when the first person went to sleep and the last person awoke. We show that this asynchrony in activity levels is produced by chronotype variation, and that chronotype covaries with age. Thus, asynchronous periods of wakefulness provide an opportunity for vigilance when sleeping in groups. We propose that throughout human evolution, sleeping groups composed of mixed age classes provided a form of vigilance. Chronotype variation and human sleep architecture (including nocturnal awakenings) in modern populations may therefore represent a legacy of natural selection acting in the past to reduce the dangers of sleep., (© 2017 The Author(s).)

Objectives: We studied sleep in a rural population in Madagascar to (i) characterize sleep in an equatorial small-scale agricultural population without electricity, (ii) assess whether sleep is linked to noise levels in a dense population, and (iii) examine the effects of experimentally introduced artificial light on sleep timing., Methods: Using actigraphy, sleep-wake patterns were analyzed for both daytime napping and nighttime wakefulness in 21 participants for a sum total of 292 days. Functional linear modeling was used to characterize 24-h time-averaged circadian patterns and to investigate the effect of experimentally introduced mobile field lights on sleep timing. We also obtained the first polysomnography (PSG) recordings of sleep in a traditional population., Results: In every measure of sleep duration and quality, the Malagasy population experienced shorter and lower quality sleep when compared to similarly measured postindustrial values. The population slept for a total of 6.5 h per night and napped during 89% of recorded days. We observed a peak in activity after midnight for both sexes on 49% of nights, consistent with segmented sleep. Access to mobile field lights had no statistical effect on nighttime sleep timing. From PSG, we documented relatively short rapid eye movement (14%), poor sleep efficiency (66%), and high wake after sleep onset (162 min)., Conclusions: Sleep in this population is segmented, similar to the "first" sleep and "second" sleep reported in the historical record. Moreover, although average sleep duration and quality were lower than documented in Western populations, circadian rhythms were more stable across days., (© 2017 Wiley Periodicals, Inc.)

Objectives: Cross-cultural sleep research is critical to deciphering whether modern sleep expression is the product of recent selective pressures, or an example of evolutionary mismatch to ancestral sleep ecology. We worked with the Hadza, an equatorial, hunter-gatherer community in Tanzania, to better understand ancestral sleep patterns and to test hypotheses related to sleep segmentation., Methods: We used actigraphy to analyze sleep-wake patterns in thirty-three volunteers for a total of 393 days. Linear mixed effects modeling was performed to assess ecological predictors of sleep duration and quality. Additionally, functional linear modeling (FLM) was used to characterize 24-hr time averaged circadian patterns., Results: Compared with post-industrialized western populations, the Hadza were characterized by shorter (6.25 hr), poorer quality sleep (sleep efficiency = 68.9%), yet had stronger circadian rhythms. Sleep duration time was negatively influenced by greater activity, age, light (lux) exposure, and moon phase, and positively influenced by increased day length and mean nighttime temperature. The average daily nap ratio (i.e., the proportion of days where a nap was present) was 0.54 (SE = 0.05), with an average nap duration of 47.5 min (SE = 2.71; n = 139)., Discussion: This study showed that circadian rhythms in small-scale foraging populations are more entrained to their ecological environments than Western populations. Additionally, Hadza sleep is characterized as flexible, with a consistent early morning sleep period yet reliance upon opportunistic daytime napping. We propose that plasticity in sleep-wake patterns has been a target of natural selection in human evolution., (© 2017 Wiley Periodicals, Inc.)

Objective: To compare different scoring parameter settings in actigraphy software for inferring sleep and wake bouts for validating analytical techniques outside of laboratory environments., Design: To identify parameter settings that best identify napping during periods of wakefulness, we analyzed 137 days on which participants reported daytime napping, as compared with a random subset of 30 days when no naps were reported. To identify settings that identify periods of wakefulness during sleep, we used data from a subsample of women who reported discrete wake bouts while nursing at night., Setting: Equatorial Tanzania in January to February 2016., Participants: The Hadza-a non-industrial foraging population., Measurements: Thirty-three subjects participated in the study for 393 observation days. Using the Bland-Altman technique to determine concordance, we analyzed reported events of daytime napping and nighttime wake bouts., Results: Only 1 parameter setting could reliably detect reported naps (15-minute nap length, ≤50 counts). Moreover, of the 6 tested parameter settings to detect wake bouts, the setting where the sleep-wake algorithm was parameterized to detect 20 consecutive minutes throughout the designated sleep period did not overestimate or underestimate wake bouts, had the lowest mean difference, and did not significantly differ from reported wake-bout events., Conclusion: We propose operational definitions for multiple dimensions of segmented sleep and conclude that actigraphy is an effective method for detecting segmented sleep in future cross-site comparative research. The implications of such work are far reaching, as sleep research in preindustrial and developing societies is documenting natural sleep-wake patterns in previously inaccessible environments., (Copyright \© 2016 National Sleep Foundation. Published by Elsevier Inc. All rights reserved.)

Over the past four decades, scientists have made substantial progress in understanding the evolution of sleep patterns across the Tree of Life. Remarkably, the specifics of sleep along the human lineage have been slow to emerge. This is surprising, given our unique mental and behavioral capacity and the importance of sleep for individual cognitive performance. One view is that our species' sleep architecture is in accord with patterns documented in other mammals. We promote an alternative view, that human sleep is highly derived relative to that of other primates. Based on new and existing evidence, we specifically propose that humans are more efficient in their sleep patterns than are other primates, and that human sleep is shorter, deeper, and exhibits a higher proportion of REM than expected. Thus, we propose the sleep intensity hypothesis: Early humans experienced selective pressure to fulfill sleep needs in the shortest time possible. Several factors likely served as selective pressures for more efficient sleep, including increased predation risk in terrestrial environments, threats from intergroup conflict, and benefits arising from increased social interaction. Less sleep would enable longer active periods in which to acquire and transmit new skills and knowledge, while deeper sleep may be critical for the consolidation of those skills, leading to enhanced cognitive abilities in early humans., (© 2015 Wiley Periodicals, Inc.)

Background: Sleep is a biological enigma. Despite occupying much of an animal's life, and having been scrutinized by numerous experimental studies, there is still no consensus on its function. Similarly, no hypothesis has yet explained why species have evolved such marked variation in their sleep requirements (from 3 to 20 hours a day in mammals). One intriguing but untested idea is that sleep has evolved by playing an important role in protecting animals from parasitic infection. This theory stems, in part, from clinical observations of intimate physiological links between sleep and the immune system. Here, we test this hypothesis by conducting comparative analyses of mammalian sleep, immune system parameters, and parasitism., Results: We found that evolutionary increases in mammalian sleep durations are strongly associated with an enhancement of immune defences as measured by the number of immune cells circulating in peripheral blood. This appeared to be a generalized relationship that could be independently detected in 4 of the 5 immune cell types and in both of the main sleep phases. Importantly, no comparable relationships occur in related physiological systems that do not serve an immune function. Consistent with an influence of sleep on immune investment, mammalian species that sleep for longer periods also had substantially reduced levels of parasitic infection., Conclusion: These relationships suggest that parasite resistance has played an important role in the evolution of mammalian sleep. Species that have evolved longer sleep durations appear to be able to increase investment in their immune systems and be better protected from parasites. These results are neither predicted nor explained by conventional theories of sleep evolution, and suggest that sleep has a much wider role in disease resistance than is currently appreciated.

Sleep is a pervasive characteristic of mammalian species, yet its purpose remains obscure. It is often proposed that 'sleep is for the brain', a view that is supported by experimental studies showing that sleep improves cognitive processes such as memory consolidation. Some comparative studies have also reported that mammalian sleep durations are higher among more encephalized species. However, no study has assessed the relationship between sleep and the brain structures that are implicated in specific cognitive processes across species. The hippocampus, neocortex and amygdala are important for memory consolidation and learning and are also in a highly actived state during sleep. We therefore investigated the evolutionary relationship between mammalian sleep and the size of these brain structures using phylogenetic comparative methods. We found that evolutionary increases in the size of the amygdala are associated with corresponding increases in NREM sleep durations. These results are consistent with the hypothesis that NREM sleep is functionally linked with specializations of the amygdala, including perhaps memory processing.

Background: All animals thus far studied sleep, but little is known about the ecological factors that generate differences in sleep characteristics across species, such as total sleep duration or division of sleep into multiple bouts across the 24-hour period (i.e., monophasic or polyphasic sleep activity). Here we address these questions using an evolutionary agent-based model. The model is spatially explicit, with food and sleep sites distributed in two clusters on the landscape. Agents acquire food and sleep energy based on an internal circadian clock coded by 24 traits (one for each hour of the day) that correspond to "genes" that evolve by means of a genetic algorithm. These traits can assume three different values that specify the agents' behavior: sleep (or search for a sleep site), eat (or search for a food site), or flexibly decide action based on relative levels of sleep energy and food energy. Individuals with higher fitness scores leave more offspring in the next generation of the simulation, and the model can therefore be used to identify evolutionarily adaptive circadian clock parameters under different ecological conditions., Results: We systematically varied input parameters related to the number of food and sleep sites, the degree to which food and sleep sites overlap, and the rate at which food patches were depleted. Our results reveal that: (1) the increased costs of traveling between more spatially separated food and sleep clusters select for monophasic sleep, (2) more rapid food patch depletion reduces sleep times, and (3) agents spend more time attempting to acquire the "rarer" resource, that is, the average time spent sleeping is positively correlated with the number of food patches and negatively correlated with the number of sleep patches. "Flexible" genes, in general, do not appear to be advantageous, though their arrangements in the agents' genome show characteristic patterns that suggest that selection acts on their distribution., Conclusion: Collectively, the output suggests that ecological factors can have striking effects on sleep patterns. Moreover, our results demonstrate that a simple model can produce clear and sensible patterns, thus allowing it to be used to investigate a wide range of questions concerning the ecology of sleep. Quantitative data presently are unavailable to test the model predictions directly, but patterns are consistent with comparative evidence from different species, and the model can be used to target ecological factors to investigate in future research.

Sleep is essential for survival, yet it also represents a time of extreme vulnerability to predation, hostile conspecifics and environmental dangers. To reduce the risks of sleeping, the sentinel hypothesis proposes that group-living animals share the task of vigilance during sleep, with some individuals sleeping while others are awake. To investigate sentinel-like behaviour in sleeping humans, we investigated activity patterns at night among Hadza hunter-gatherers of Tanzania. Using actigraphy, we discovered that all subjects were simultaneously scored as asleep for only 18 min in total over 20 days of observation, with a median of eight individuals awake throughout the nighttime period; thus, one or more individuals was awake (or in light stages of sleep) during 99.8% of sampled epochs between when the first person went to sleep and the last person awoke. We show that this asynchrony in activity levels is produced by chronotype variation, and that chronotype covaries with age. Thus, asynchronous periods of wakefulness provide an opportunity for vigilance when sleeping in groups. We propose that throughout human evolution, sleeping groups composed of mixed age classes provided a form of vigilance. Chronotype variation and human sleep architecture (including nocturnal awakenings) in modern populations may therefore represent a legacy of natural selection acting in the past to reduce the dangers of sleep. [ABSTRACT FROM AUTHOR]

All mammals thus far studied sleep, yet important questions remain concerning the ecological factors that influence sleep patterns. Here, we developed an evolutionary individual-based model to investigate the effect of predation pressure on prey sleep. We investigated three ecological conditions, including one that assumed a dynamic interaction between predator and prey behaviour. In condition 1, we found that monophasic predators (i.e. with one sleep bout per 24h) select for monophasic prey that sleep perfectly out of phase with predators. In condition 2, predators were monophasic but the safety of prey varied as a function of their activity (sleeping versus awake). In this condition, the prey adjusted their sleeping behaviour to lower the risk of predation. Finally, in condition 3, we modelled a more dynamic interaction between predator and prey, with predator activity dependent on prey activity in the previous hour. In this scenario, the prey adjusted their behaviour relative to one another, resulting in either greater or lesser synchrony in prey as a function of predator searching behaviour. Collectively, our model demonstrates that predator behaviour can have a strong influence on prey sleep patterns, including whether prey are monophasic or polyphasic (i.e. with many sleep bouts per 24h). The model further suggests that the timing of sleep relative to predator behaviour may depend strongly on how other potential prey partition the activity period. [ABSTRACT FROM AUTHOR]