Under real-life conditions, increased light exposure during wakefulness seems associated with imp... more Under real-life conditions, increased light exposure during wakefulness seems associated with improved sleep quality, quantified as reduced time awake during bed time, increased time spent in non-rapid eye movement (NREM) sleep, or increased power of the electroencephalogram delta band (0.5-4 Hz). The causality of these important relationships and their dependency on circadian phase and/or time awake has not been studied in depth. To disentangle possible circadian and homeostatic interactions, we employed a forced desynchrony protocol under dim light (6 lux) and under bright light (1300 lux) during wakefulness. Our protocol consisted of a fast cycling sleep-wake schedule (13 h wakefulness—5 h sleep; 4 cycles), followed by 3 h recovery sleep in a within-subject cross-over design. Individuals (8 men) were equipped with 10 polysomnography electrodes. Subjective sleep quality was measured immediately after wakening with a questionnaire. Results indicated that circadian variation in delt...
The circadian system affects physiological, psychological, and molecular mechanisms in the body, ... more The circadian system affects physiological, psychological, and molecular mechanisms in the body, resulting in varying physical performance over the day. The timing and relative size of these effects are important for optimizing sport performance. In this study, Olympic swim times (from 2004 to 2016) were used to determine time-of-day and circadian effects under maximal motivational conditions. Data of athletes who made it to the finals (N = 144, 72 female) were included and normalized on individual levels based on the average swim times over race types (heat, semifinal, and final) per individual for each stroke, distance and Olympic venue. Normalized swim times were analyzed with a linear mixed model and a sine fitted model. Swim performance was better during finals as compared to semi-finals and heats. Performance was strongly affected by time-of-day, showing fastest swim times in the late afternoon around 17:12 h, indicating 0.32% improved performance relative to 08:00 h. This stu...
Human thermoregulation is strictly regulated by the preoptic area of the hypothalamus, which is d... more Human thermoregulation is strictly regulated by the preoptic area of the hypothalamus, which is directly influenced by the suprachiasmatic nucleus (SCN). The main input pathway of the SCN is light. Here, thermoregulatory effects of light were assessed in humans in a forced desynchrony (FD) design. The FD experiment was performed in dim light (DL, 6 lux) and bright white light (BL, 1300 lux) in 8 men in a semi-randomized within-subject design. A 4 × 18 h FD protocol (5 h sleep, 13 h wake) was applied, with continuous core body temperature (CBT) and skin temperature measurements at the forehead, clavicles, navel, palms, foot soles and toes. Skin temperature parameters indicated sleep-wake modulations as well as internal clock variations. All distal skin temperature parameters increased during sleep, when CBT decreased. Light significantly affected temperature levels during the wake phase, with decreased temperature measured at the forehead and toes and increased navel and clavicular s...
International journal of legal medicine, Jan 5, 2017
Trace deposition timing reflects a novel concept in forensic molecular biology involving the use ... more Trace deposition timing reflects a novel concept in forensic molecular biology involving the use of rhythmic biomarkers for estimating the time within a 24-h day/night cycle a human biological sample was left at the crime scene, which in principle allows verifying a sample donor's alibi. Previously, we introduced two circadian hormones for trace deposition timing and recently demonstrated that messenger RNA (mRNA) biomarkers significantly improve time prediction accuracy. Here, we investigate the suitability of metabolites measured using a targeted metabolomics approach, for trace deposition timing. Analysis of 171 plasma metabolites collected around the clock at 2-h intervals for 36 h from 12 male participants under controlled laboratory conditions identified 56 metabolites showing statistically significant oscillations, with peak times falling into three day/night time categories: morning/noon, afternoon/evening and night/early morning. Time prediction modelling identified 10 ...
In species that initiate multiple broods in a single season there is usually a trade-off between ... more In species that initiate multiple broods in a single season there is usually a trade-off between the number of young in the first brood, and the timing and occurrence of subsequent broods in the same season. The hypothesis that experimental reduction of clutch size reduces the duration of post-hedging care was tested in great tits, Parus major, and the effect
Proceedings of the Royal Society B: Biological Sciences, 2013
Seasonal recurrence of biological processes (phenology) and its relationship to environmental cha... more Seasonal recurrence of biological processes (phenology) and its relationship to environmental change is recognized as being of key scientific and public concern, but its current study largely overlooks the extent to which phenology is based on biological time-keeping mechanisms. We highlight the relevance of physiological and neurobiological regulation for organisms’ responsiveness to environmental conditions. Focusing on avian and mammalian examples, we describe circannual rhythmicity of reproduction, migration and hibernation, and address responses of animals to photic and thermal conditions. Climate change and urbanization are used as urgent examples of anthropogenic influences that put biological timing systems under pressure. We furthermore propose that consideration ofHomo sapiensas principally a ‘seasonal animal’ can inspire new perspectives for understanding medical and psychological problems.
With time-place learning, animals link a stimulus with the location and the time of day. This abi... more With time-place learning, animals link a stimulus with the location and the time of day. This ability may optimize resource localization and predator avoidance in daily changing environments. Time-place learning is a suitable task to study the interaction of the circadian system and memory. Previously, we showed that time-place learning in mice depends on the circadian system and Cry1 and/or Cry2 clock genes. We questioned whether time-place learning is Cry specific or also depends on other core molecular clock genes. Here, we show that Per1/Per2 double mutant mice, despite their arrhythmic phenotype, acquire time-place learning similar to wild-type mice. As well as an established role in circadian rhythms, Per genes have also been implicated in the formation and storage of memory. We found no deficiencies in short-term spatial working memory in Per mutant mice compared to wild-type mice. Moreover, both Per mutant and wild-type mice showed similar long-term memory for contextual fea...
Light-induced improvements in alertness are more prominent during nighttime than during the day, ... more Light-induced improvements in alertness are more prominent during nighttime than during the day, suggesting that alerting effects of light may depend on internal clock time or wake duration. Relative contributions of both factors can be quantified using a forced desynchrony (FD) designs. FD designs have only been conducted under dim light conditions (<10 lux) since light above this amount can induce non-uniform phase progression of the circadian pacemaker (also called relative coordination). This complicates the mathematical separation of circadian clock phase from homeostatic sleep pressure effects. Here we investigate alerting effects of light in a novel 4 × 18 h FD protocol (5 h sleep, 13 h wake) under dim (6 lux) and bright light (1300 lux) conditions. Hourly saliva samples (melatonin and cortisol assessment) and 2-hourly test sessions were used to assess effects of bright light on subjective and objective alertness (electroencephalography and performance). Results reveal (1)...
Endogenous daily (circadian) rhythms allow organisms to anticipate daily changes in the environme... more Endogenous daily (circadian) rhythms allow organisms to anticipate daily changes in the environment. Most mammals are specialized to be active during the night (nocturnal) or day (diurnal). However, typically nocturnal mammals become diurnal when energetically challenged by cold or hunger. The circadian thermo-energetics (CTE) hypothesis predicts that diurnal activity patterns reduce daily energy expenditure (DEE) compared with nocturnal activity patterns. Here, we tested the CTE hypothesis by quantifying the energetic consequences of relevant environmental factors in mice. Under natural conditions, diurnality reduces DEE by 6-10% in energetically challenged mice. Combined with night-time torpor, as observed in mice under prolonged food scarcity, DEE can be reduced by ∼20%. The dominant factor determining the energetic benefit of diurnality is thermal buffering provided by a sheltered resting location. Compared with nocturnal animals, diurnal animals encounter higher ambient temperatures during both day and night, leading to reduced thermogenesis costs in temperate climates. Analysis of weather station data shows that diurnality is energetically beneficial on almost all days of the year in a temperate climate region. Furthermore, diurnality provides energetic benefits at all investigated geographical locations on European longitudinal and latitudinal transects. The reduction of DEE by diurnality provides an ultimate explanation for temporal niche switching observed in typically nocturnal small mammals under energetically challenging conditions. Diurnality allows mammals to compensate for reductions in food availability and temperature as it reduces energetic needs. The optimal circadian organization of an animal ultimately depends on the balance between energetic consequences and other fitness consequences of the selected temporal niche.
Circadian clocks drive daily rhythms in physiology and behavior which allow organisms to anticipa... more Circadian clocks drive daily rhythms in physiology and behavior which allow organisms to anticipate predictable daily changes in the environment. In most mammals, circadian rhythms result in nocturnal activity patterns although plasticity of the circadian system allows activity patterns to shift to different times of day. Such plasticity is seen when food access is restricted to a few hours during the resting (light) phase resulting in food anticipatory activity (FAA) in the hours preceding food availability. The mechanisms underlying FAA are unknown but data suggests the involvement of the reward system and homeostatic regulation of metabolism. We previously demonstrated the isolated effect of metabolism by inducing diurnality in response to energetic challenges. Here the importance of reward timing in inducing daytime activity is assessed. The daily activity distribution of mice earning palatable chocolate at their preferred time by working in a running wheel was compared with tha...
Proceedings of the National Academy of Sciences of the United States of America, Jan 21, 2014
The mammalian circadian system synchronizes daily timing of activity and rest with the environmen... more The mammalian circadian system synchronizes daily timing of activity and rest with the environmental light-dark cycle. Although the underlying molecular oscillatory mechanism is well studied, factors that influence phenotypic plasticity in daily activity patterns (temporal niche switching, chronotype) are presently unknown. Molecular evidence suggests that metabolism may influence the circadian molecular clock, but evidence at the level of the organism is lacking. Here we show that a metabolic challenge by cold and hunger induces diurnality in otherwise nocturnal mice. Lowering ambient temperature changes the phase of circadian light-dark entrainment in mice by increasing daytime and decreasing nighttime activity. This effect is further enhanced by simulated food shortage, which identifies metabolic balance as the underlying common factor influencing circadian organization. Clock gene expression analysis shows that the underlying neuronal mechanism is downstream from or parallel to ...
Deep hibernators that spend winter in a hypothermic coma below ground can still emerge and reprod... more Deep hibernators that spend winter in a hypothermic coma below ground can still emerge and reproduce in spring at the right moment. A recent study shows that specific cells of the pituitary may harbor the internal calendar responsible for this.
Time as an ecological niche variable or &amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;... more Time as an ecological niche variable or &amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;quot;temporal niche&amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;quot; can be defined in the context of the most prominent environmental cycles, including the tidal cycle, the lunar day and month, the solar day, and the earth year. For the current review, we focus on the 24-h domain generated through the earth&amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;#39;s rotation around its axis (solar day). The daily environmental cycles of light and temperature are a dominant ecological factor generating a variety of adaptations among animals. In this review, we describe these adaptations with a special focus on the visual system and on the adaptive plasticity of activity patterns. Our goals are: (1) Underscore the importance of the 24-h time axis as critical variable in the ecological niche. (2) Highlight cases of temporal niche switches at the evolutionary timescale (phylogenetic level). (3) Review temporal niche switching within an individual&amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;#39;s lifespan. (4) Evaluate possible underlying mechanisms for temporal niche switching. (5) Describe a new hypothesis of circadian thermoenergetics which may explain several cases of temporal niche switching in mammals. With this, we hope to inspire experiments under natural conditions or more complex laboratory environments, aimed to reveal environmental factors and mechanisms underlying specific temporal programs.
Circadian clocks of adolescents typically run late-including sleep times-yet adolescents generall... more Circadian clocks of adolescents typically run late-including sleep times-yet adolescents generally are expected at school early in the morning. Due to this mismatch between internal (circadian) and external (social) times, adolescents suffer from chronic sleep deficiency, which, in turn, affects academic performance negatively. This constellation affects students' future career prospects. Our study correlates chronotype and examination performance. In total, 4734 grades were collected from 741 Dutch high school students (ages 11-18 years) who had completed the Munich ChronoType Questionnaire to estimate their internal time. Overall, the lowest grades were obtained by students who were very late chronotypes (MSFsc > 5.31 h) or slept very short on schooldays (SDw < 7.03 h). The effect of chronotype on examination performance depended on the time of day that examinations were taken. Opposed to late types, early chronotypes obtained significantly higher grades during the early...
Under real-life conditions, increased light exposure during wakefulness seems associated with imp... more Under real-life conditions, increased light exposure during wakefulness seems associated with improved sleep quality, quantified as reduced time awake during bed time, increased time spent in non-rapid eye movement (NREM) sleep, or increased power of the electroencephalogram delta band (0.5-4 Hz). The causality of these important relationships and their dependency on circadian phase and/or time awake has not been studied in depth. To disentangle possible circadian and homeostatic interactions, we employed a forced desynchrony protocol under dim light (6 lux) and under bright light (1300 lux) during wakefulness. Our protocol consisted of a fast cycling sleep-wake schedule (13 h wakefulness—5 h sleep; 4 cycles), followed by 3 h recovery sleep in a within-subject cross-over design. Individuals (8 men) were equipped with 10 polysomnography electrodes. Subjective sleep quality was measured immediately after wakening with a questionnaire. Results indicated that circadian variation in delt...
The circadian system affects physiological, psychological, and molecular mechanisms in the body, ... more The circadian system affects physiological, psychological, and molecular mechanisms in the body, resulting in varying physical performance over the day. The timing and relative size of these effects are important for optimizing sport performance. In this study, Olympic swim times (from 2004 to 2016) were used to determine time-of-day and circadian effects under maximal motivational conditions. Data of athletes who made it to the finals (N = 144, 72 female) were included and normalized on individual levels based on the average swim times over race types (heat, semifinal, and final) per individual for each stroke, distance and Olympic venue. Normalized swim times were analyzed with a linear mixed model and a sine fitted model. Swim performance was better during finals as compared to semi-finals and heats. Performance was strongly affected by time-of-day, showing fastest swim times in the late afternoon around 17:12 h, indicating 0.32% improved performance relative to 08:00 h. This stu...
Human thermoregulation is strictly regulated by the preoptic area of the hypothalamus, which is d... more Human thermoregulation is strictly regulated by the preoptic area of the hypothalamus, which is directly influenced by the suprachiasmatic nucleus (SCN). The main input pathway of the SCN is light. Here, thermoregulatory effects of light were assessed in humans in a forced desynchrony (FD) design. The FD experiment was performed in dim light (DL, 6 lux) and bright white light (BL, 1300 lux) in 8 men in a semi-randomized within-subject design. A 4 × 18 h FD protocol (5 h sleep, 13 h wake) was applied, with continuous core body temperature (CBT) and skin temperature measurements at the forehead, clavicles, navel, palms, foot soles and toes. Skin temperature parameters indicated sleep-wake modulations as well as internal clock variations. All distal skin temperature parameters increased during sleep, when CBT decreased. Light significantly affected temperature levels during the wake phase, with decreased temperature measured at the forehead and toes and increased navel and clavicular s...
International journal of legal medicine, Jan 5, 2017
Trace deposition timing reflects a novel concept in forensic molecular biology involving the use ... more Trace deposition timing reflects a novel concept in forensic molecular biology involving the use of rhythmic biomarkers for estimating the time within a 24-h day/night cycle a human biological sample was left at the crime scene, which in principle allows verifying a sample donor's alibi. Previously, we introduced two circadian hormones for trace deposition timing and recently demonstrated that messenger RNA (mRNA) biomarkers significantly improve time prediction accuracy. Here, we investigate the suitability of metabolites measured using a targeted metabolomics approach, for trace deposition timing. Analysis of 171 plasma metabolites collected around the clock at 2-h intervals for 36 h from 12 male participants under controlled laboratory conditions identified 56 metabolites showing statistically significant oscillations, with peak times falling into three day/night time categories: morning/noon, afternoon/evening and night/early morning. Time prediction modelling identified 10 ...
In species that initiate multiple broods in a single season there is usually a trade-off between ... more In species that initiate multiple broods in a single season there is usually a trade-off between the number of young in the first brood, and the timing and occurrence of subsequent broods in the same season. The hypothesis that experimental reduction of clutch size reduces the duration of post-hedging care was tested in great tits, Parus major, and the effect
Proceedings of the Royal Society B: Biological Sciences, 2013
Seasonal recurrence of biological processes (phenology) and its relationship to environmental cha... more Seasonal recurrence of biological processes (phenology) and its relationship to environmental change is recognized as being of key scientific and public concern, but its current study largely overlooks the extent to which phenology is based on biological time-keeping mechanisms. We highlight the relevance of physiological and neurobiological regulation for organisms’ responsiveness to environmental conditions. Focusing on avian and mammalian examples, we describe circannual rhythmicity of reproduction, migration and hibernation, and address responses of animals to photic and thermal conditions. Climate change and urbanization are used as urgent examples of anthropogenic influences that put biological timing systems under pressure. We furthermore propose that consideration ofHomo sapiensas principally a ‘seasonal animal’ can inspire new perspectives for understanding medical and psychological problems.
With time-place learning, animals link a stimulus with the location and the time of day. This abi... more With time-place learning, animals link a stimulus with the location and the time of day. This ability may optimize resource localization and predator avoidance in daily changing environments. Time-place learning is a suitable task to study the interaction of the circadian system and memory. Previously, we showed that time-place learning in mice depends on the circadian system and Cry1 and/or Cry2 clock genes. We questioned whether time-place learning is Cry specific or also depends on other core molecular clock genes. Here, we show that Per1/Per2 double mutant mice, despite their arrhythmic phenotype, acquire time-place learning similar to wild-type mice. As well as an established role in circadian rhythms, Per genes have also been implicated in the formation and storage of memory. We found no deficiencies in short-term spatial working memory in Per mutant mice compared to wild-type mice. Moreover, both Per mutant and wild-type mice showed similar long-term memory for contextual fea...
Light-induced improvements in alertness are more prominent during nighttime than during the day, ... more Light-induced improvements in alertness are more prominent during nighttime than during the day, suggesting that alerting effects of light may depend on internal clock time or wake duration. Relative contributions of both factors can be quantified using a forced desynchrony (FD) designs. FD designs have only been conducted under dim light conditions (<10 lux) since light above this amount can induce non-uniform phase progression of the circadian pacemaker (also called relative coordination). This complicates the mathematical separation of circadian clock phase from homeostatic sleep pressure effects. Here we investigate alerting effects of light in a novel 4 × 18 h FD protocol (5 h sleep, 13 h wake) under dim (6 lux) and bright light (1300 lux) conditions. Hourly saliva samples (melatonin and cortisol assessment) and 2-hourly test sessions were used to assess effects of bright light on subjective and objective alertness (electroencephalography and performance). Results reveal (1)...
Endogenous daily (circadian) rhythms allow organisms to anticipate daily changes in the environme... more Endogenous daily (circadian) rhythms allow organisms to anticipate daily changes in the environment. Most mammals are specialized to be active during the night (nocturnal) or day (diurnal). However, typically nocturnal mammals become diurnal when energetically challenged by cold or hunger. The circadian thermo-energetics (CTE) hypothesis predicts that diurnal activity patterns reduce daily energy expenditure (DEE) compared with nocturnal activity patterns. Here, we tested the CTE hypothesis by quantifying the energetic consequences of relevant environmental factors in mice. Under natural conditions, diurnality reduces DEE by 6-10% in energetically challenged mice. Combined with night-time torpor, as observed in mice under prolonged food scarcity, DEE can be reduced by ∼20%. The dominant factor determining the energetic benefit of diurnality is thermal buffering provided by a sheltered resting location. Compared with nocturnal animals, diurnal animals encounter higher ambient temperatures during both day and night, leading to reduced thermogenesis costs in temperate climates. Analysis of weather station data shows that diurnality is energetically beneficial on almost all days of the year in a temperate climate region. Furthermore, diurnality provides energetic benefits at all investigated geographical locations on European longitudinal and latitudinal transects. The reduction of DEE by diurnality provides an ultimate explanation for temporal niche switching observed in typically nocturnal small mammals under energetically challenging conditions. Diurnality allows mammals to compensate for reductions in food availability and temperature as it reduces energetic needs. The optimal circadian organization of an animal ultimately depends on the balance between energetic consequences and other fitness consequences of the selected temporal niche.
Circadian clocks drive daily rhythms in physiology and behavior which allow organisms to anticipa... more Circadian clocks drive daily rhythms in physiology and behavior which allow organisms to anticipate predictable daily changes in the environment. In most mammals, circadian rhythms result in nocturnal activity patterns although plasticity of the circadian system allows activity patterns to shift to different times of day. Such plasticity is seen when food access is restricted to a few hours during the resting (light) phase resulting in food anticipatory activity (FAA) in the hours preceding food availability. The mechanisms underlying FAA are unknown but data suggests the involvement of the reward system and homeostatic regulation of metabolism. We previously demonstrated the isolated effect of metabolism by inducing diurnality in response to energetic challenges. Here the importance of reward timing in inducing daytime activity is assessed. The daily activity distribution of mice earning palatable chocolate at their preferred time by working in a running wheel was compared with tha...
Proceedings of the National Academy of Sciences of the United States of America, Jan 21, 2014
The mammalian circadian system synchronizes daily timing of activity and rest with the environmen... more The mammalian circadian system synchronizes daily timing of activity and rest with the environmental light-dark cycle. Although the underlying molecular oscillatory mechanism is well studied, factors that influence phenotypic plasticity in daily activity patterns (temporal niche switching, chronotype) are presently unknown. Molecular evidence suggests that metabolism may influence the circadian molecular clock, but evidence at the level of the organism is lacking. Here we show that a metabolic challenge by cold and hunger induces diurnality in otherwise nocturnal mice. Lowering ambient temperature changes the phase of circadian light-dark entrainment in mice by increasing daytime and decreasing nighttime activity. This effect is further enhanced by simulated food shortage, which identifies metabolic balance as the underlying common factor influencing circadian organization. Clock gene expression analysis shows that the underlying neuronal mechanism is downstream from or parallel to ...
Deep hibernators that spend winter in a hypothermic coma below ground can still emerge and reprod... more Deep hibernators that spend winter in a hypothermic coma below ground can still emerge and reproduce in spring at the right moment. A recent study shows that specific cells of the pituitary may harbor the internal calendar responsible for this.
Time as an ecological niche variable or &amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;... more Time as an ecological niche variable or &amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;quot;temporal niche&amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;quot; can be defined in the context of the most prominent environmental cycles, including the tidal cycle, the lunar day and month, the solar day, and the earth year. For the current review, we focus on the 24-h domain generated through the earth&amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;#39;s rotation around its axis (solar day). The daily environmental cycles of light and temperature are a dominant ecological factor generating a variety of adaptations among animals. In this review, we describe these adaptations with a special focus on the visual system and on the adaptive plasticity of activity patterns. Our goals are: (1) Underscore the importance of the 24-h time axis as critical variable in the ecological niche. (2) Highlight cases of temporal niche switches at the evolutionary timescale (phylogenetic level). (3) Review temporal niche switching within an individual&amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;#39;s lifespan. (4) Evaluate possible underlying mechanisms for temporal niche switching. (5) Describe a new hypothesis of circadian thermoenergetics which may explain several cases of temporal niche switching in mammals. With this, we hope to inspire experiments under natural conditions or more complex laboratory environments, aimed to reveal environmental factors and mechanisms underlying specific temporal programs.
Circadian clocks of adolescents typically run late-including sleep times-yet adolescents generall... more Circadian clocks of adolescents typically run late-including sleep times-yet adolescents generally are expected at school early in the morning. Due to this mismatch between internal (circadian) and external (social) times, adolescents suffer from chronic sleep deficiency, which, in turn, affects academic performance negatively. This constellation affects students' future career prospects. Our study correlates chronotype and examination performance. In total, 4734 grades were collected from 741 Dutch high school students (ages 11-18 years) who had completed the Munich ChronoType Questionnaire to estimate their internal time. Overall, the lowest grades were obtained by students who were very late chronotypes (MSFsc > 5.31 h) or slept very short on schooldays (SDw < 7.03 h). The effect of chronotype on examination performance depended on the time of day that examinations were taken. Opposed to late types, early chronotypes obtained significantly higher grades during the early...
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