Outdoor light at night (LAN) is correlated with eveningness in adolescents

External zeitgebers synchronize the human circadian rhythm of sleep and wakefulness. Humans adapt their chronotype to the day-night cycle, the strongest external zeitgeber. The human circadian rhythm shifts to evening-type orientation when daylight is prolonged into the evening and night hours by artificial light sources. Data from a survey of 1507 German adolescents covering questions about chronotype and electronic screen media use combined with nocturnal satellite image data suggest a relationship between chronotype and artificial nocturnal light. Adolescents living in brightly illuminated urban districts had a stronger evening-type orientation than adolescents living in darker and more rural municipalities. This result persisted when controlling for time use of electronic screen media, intake of stimulants, type of school, age, puberty status, time of sunrise, sex, and population density. Time spent on electronic screen media use-a source of indoor light at night-is also correlated with eveningness, as well as intake of stimulants, age, and puberty status, and, to a lesser degree, type of school and time of sunrise. Adequate urban development design and parents limiting adolescents' electronic screen media use in the evening could help to adjust adolescents' zeitgeber to early school schedules when they provide appropriate lighting conditions for daytime and for nighttime.     

Further Evidence for the Influence of Photoperiod at Birth on Chronotype in a Sample of German Adolescents

Individuals differ in their circadian preferences (chronotype). There is evidence in the literature to support a season-of-birth effect on chronotype but the evidence is not convincing. In part, the relationship is obscured by a number of methodological differences between studies, including the measures used to define morningness, the way in which the seasons were categorized, and the sample size. This study adds to the literature in several ways. First, we adopt a new approach to categorizing the photoperiod rather than the calendar season; thus we prefer to use the term photoperiod at birth. Second, we used two measures of morningness. Third, we used a large and homogeneous German sample. The results show that adolescents (n?=?2905) born during the increasing photoperiod (Feb–Apr) had a significantly later midpoint of sleep (MSFsc) than those born during the decreasing photoperiod (Aug–Oct). A similar pattern was found for the Composite Scale of Morningness (CSM). Furthermore, both measures of chronotype demonstrated a significant quadratic function over a 1-yr cycle. When looking at each of six consecutive years separately, the Composite Scale of Morningness suggests a cosine rhythm linked to increasing and decreasing photoperiods that becomes weaker in amplitude with increasing age. Despite the strengths in our study, the effect of photoperiod at birth on chronotype remains small. Future studies may require larger sample sizes, may need to explore how neonatal light exposure modulates chronotype, and may need to track how puberty and adolescent lifestyle habits mask the photoperiod effect. (Author correspondence: vollmer@ph-heidelberg.de)     

The Pattern of Entrainment of the Human Sleep-Wake Rhythm by the Natural Photoperiod in the North

Recently, it was shown that the sleep-wake rhythm of the inhabitants of the temperate zone is entrained to sun time. In the North, significant seasonal changes in the photoperiod may interfere with entrainment of the circadian system to sunlight. This investigation assessed the influence of photoperiod characteristics on the sleep length and sleep-wake rhythm of residents of high latitude. The study was conducted in four towns and six villages located between 59.5°N and 67.6°N latitude between the months of October and May from 2009 through 2011 and included 2822 subjects aged 10 to 97 yrs, 1621 of whom were females and 1201 males. The chronotype and sleep length of the subjects were assessed using the Munich Chronotype Questionnaire. The instructions for the questionnaire stressed the need to specify the sleep-wake schedule during the week preceding the date of completing the questionnaire. The study found that the length of sleep and the chronotype of the inhabitants surveyed in Northern European Russia depend on age, sex, type of settlement, and place of residence. The time of sunrise was a stronger predictor of sleep length and chronotype than the time of sunset and day length. A later chronotype and shorter sleep length were found for the subjects during the equinox (sunrise at 06:00?h) than under long-photoperiod conditions (sunrise at 04:00–05:00?h). During short-photoperiod conditions (sunrise at 07:00–10:00?h), no significant changes in the self-reported sleep-wake rhythm were found. The time of sunrise had the strongest impact on the sleep-wake rhythm of 30- to 97-yr-old persons. Sunrise had a stronger influence on chronotype and sleep length in January to May, when the days become longer, than in October to December, when the days become shorter. Age- and season-associated changes were found in the entrainment of the sleep-wake rhythm by photoperiod in the North. (Author correspondence: borisenkov@physiol.komisc.ru)     

Time Course of Neurobehavioral Alertness During Extended Wakefulness in Morning- and Evening-Type Healthy Sleepers

The aim of the study was to evaluate the influence of chronotype (morning-type versus evening-type) living in a fixed sleep-wake schedule different from one's preferred sleep schedules on the time course of neurobehavioral performance during controlled extended wakefulness. The authors studied 9 morning-type and 9 evening-type healthy male subjects (21.4?±?1.9 yrs). Before the experiment, all participants underwent a fixed sleep-wake schedule mimicking a regular working day (bedtime: 23:30?h; wake time: 07:30?h). Then, following two nights in the laboratory, both chronotypes underwent a 36-h constant routine, performing a cognitive test of sustained attention every hour. Core body temperature, salivary melatonin secretion, objective alertness (maintenance of wakefulness test), and subjective sleepiness (visual analog scale) were also assessed. Evening-types expressed a higher level of subjective sleepiness than morning types, whereas their objective levels of alertness were not different. Cognitive performance in the lapse domain remained stable during the normal waking day and then declined during the biological night, with a similar time course for both chronotypes. Evening types maintained optimal alertness (i.e., 10% fastest reaction time) throughout the night, whereas morning types did not. For both chronotypes, the circadian performance profile was correlated with the circadian subjective somnolence profile and was slightly phase-delayed with melatonin secretion. Circadian performance was less correlated with circadian core body temperature. Lapse domain was phase-delayed with body temperature (2–4?h), whereas optimal alertness was slightly phase-delayed with body temperature (1?h). These results indicate evening types living in a fixed sleep-wake schedule mimicking a regular working day (different from their preferred sleep schedules) express higher subjective sleepiness but can maintain the same level of objective alertness during a normal waking day as morning types. Furthermore, evening types were found to maintain optimal alertness throughout their nighttime, whereas morning types could not. The authors suggest that evening-type subjects have a higher voluntary engagement of wake-maintenance mechanisms during extended wakefulness due to adaptation of their sleep-wake schedule to social constraints. (Author correspondence: jack.taillard@gmail.com)     

Functional diversities of two activity components of circadian rhythm in genetical splitting mice (CS strain)

CS mice, an inbred strain, showed two distinctive characteristics in the circadian rhythm of locomotor activity: (1) large variation in the freerunning period, and (2) spontaneous rhythm splitting under continuous darkness. In the splitting rhythm there was a positive correlation between the freerunning period of the evening component and the activity time of the morning component. The phase-shifting effect of a 15-min light pulse was examined on the two activity components of the splitting rhythm. There were significant differences in the amount of light-induced phase response between the two components. A light pulse during the late subjective night induced a phase advance shift only in the morning component, while a light pulse during the early subjective night induced a phase delay shift only in the evening component. These results indicate functional diversities of the two activity components in the circadian locomotor rhythm of CS mice, and suggest that the circadian system in CS mice consists of two mutually coupled oscillators which have different circadian periods and different responsiveness to light. The CS mouse is a useful model to explore a genetic background of oscillator coupling in the circadian system of nocturnal rodents. Accepted: 19 November 1998     

INFLUENCE OF SEASONALITY ON CIRCADIAN MOTOR ACTIVITY RHYTHM IN COMMON MARMOSETS DURING PUBERTY

The effect of puberty on circadian rhythmicity in nonhuman primates has been little studied, even though it has been demonstrated that puberty-related changes in circadian activity rhythm occur in a number of species, including humans. To characterize the motor activity rhythm during puberty in common marmosets (Callithrix jacchus), six animals was continuously monitored by actimeters between their 5th and 12th months of age. The animals were housed with their families in outdoor cages under seminatural conditions. Onset of puberty was determined from fecal estrogen and progesterone levels in females and androgen levels in males. The spectral power of the circadian component stabilized later in the last two animals to enter puberty. The bimodal characteristic of the active phase in this species became progressively more apparent over the course of the months in which the mean temperature was highest, irrespective of the animal's age. Although the onset of activity advanced after entry into puberty, this parameter showed a strong correlation with sunrise, indicating that seasonality influences this variable. Neither age nor climatic factors included in the regression model influenced the differences in phase angles between sunrise and onset of activity, and between sunset and offset of activity. Total activity was the only parameter influenced by age in the regression model, showing an increase after entry into puberty. Despite the evidence of pubertal influence on both the circadian component and total activity, under seminatural conditions seasonal factors may have a more important effect on motor activity rhythm in common marmosets. (Author correspondence: E-mail: carolina@cb.ufrn.br)     

Crosstalk Between Environmental Light and Internal Time in Humans

Daily exposure to environmental light is the most important zeitgeber in humans, and all studied characteristics of light pattern (timing, intensity, rate of change, duration, and spectrum) influence the circadian system. However, and due to lack of current studies on environmental light exposure and its influence on the circadian system, the aim of this work is to determine the characteristics of a naturalistic regimen of light exposure and its relationship with the functioning of the human circadian system. Eighty-eight undergraduate students (18–23 yrs) were recruited in Murcia, Spain (latitude 38°01′N) to record wrist temperature (WT), light exposure, and sleep for 1 wk under free-living conditions. Light-exposure timing, rate of change, regularity, intensity, and contrast were calculated, and their effects on the sleep pattern and WT rhythm were then analyzed. In general, higher values for interdaily stability, relative amplitude, mean morning light, and light quality index (LQI) correlated with higher interdaily stability and relative amplitude, and phase advance in sleep plus greater stability in WT and phase advance of the WT circadian rhythm. On the other hand, a higher fragmentation of the light-exposure rhythm was associated with more fragmented sleep. Naturalistic studies using 24-h ambulatory light monitoring provide essential information about the main circadian system input, necessary for maintaining healthy circadian tuning. Correcting light-exposure patterns accordingly may help prevent or even reverse health problems associated with circadian disruption. (Author correspondence: angerol@um.es)     

Impact of Perinatal Photoperiod on the Chronotype of 11- to 18-Year-Olds in Northern European Russia

The study investigates the effect of the month of birth and ambient light conditions at birth on sleep length and chronotype among residents of high latitudes. The authors surveyed 1172 persons (609 girls, 563 boys) age 11 to 18 yrs living in five villages and four towns located between 59.5°N and 67.6°N latitude. Survey participation was voluntary and anonymous. Sleep length and chronotype were assessed using the Munich chronotype questionnaire (MCTQ). The study showed the sleep length and chronotype of the children and adolescents depended on sex, age, type of settlement (town/village), and latitude of residence. Latitude exerted a stronger impact on sleep length and chronotype of children and adolescents living in villages than on those of their urban counterparts. Month of birth had no effect on sleep length and chronotype. There was a significant effect of the time of sunrise, sunset, and day length at birth on the chronotype of children and adolescents. A later chronotype was observed in the sample of young persons living above the Arctic Circle who were born during the polar day and polar night. (Author correspondence: borisenkov@physiol.komisc.ru)     

Early- and Late-Emerging Drosophila melanogaster Fruit Flies Differ in Their Sensitivity to Light During Morning and Evening

The authors derived early and late populations of fruit flies showing increased incidence of emergence during morning or evening hours by imposing selection for timing of emergence under 12:12?h light/dark (LD) cycles. From previous studies, it was clear that the increased incidence of adult emergence during morning and evening hours in early and late populations was a result of evolution of divergent and characteristic emergence waveforms in these populations. Such characteristic waveforms are henceforth referred to as “evolved emergence waveforms” (EEWs). The early and late populations also evolved different circadian clocks, which is evident from the divergence in their clock period (τ) and photic phase response curve (PRC). Although correlation between emergence waveforms and clock properties suggests functional significance of circadian clocks, τ and PRCs do not satisfactorily explain the early and late emergence phenotypes. In order to understand the functional significance of the PRC for early and late emergence phenotypes, the authors investigated whether circadian clocks of these flies exhibit any difference in photosensitivity under entrained conditions. Such differences would suggest that the light requirement for circadian entrainment of the emergence rhythm in early and late populations is different. To test this, they examined if early and late flies differ in their light utilization behavior, first by assaying their emergence rhythm under complete photoperiod and then in three different skeleton photoperiods. The results showed that early and late populations require different durations of light during the morning and evening to achieve their EEWs, suggesting that for the circadian entrainment of the emergence rhythm, early and late flies utilize light from different parts of the day. (Author correspondence: vsharma@jncasr.ac.in or vksharmas@gmail.com)     

EVENING DAYLIGHT MAY CAUSE ADOLESCENTS TO SLEEP LESS IN SPRING THAN IN WINTER

Sleep restriction commonly experienced by adolescents can stem from a slower increase in sleep pressure by the homeostatic processes and from phase delays of the circadian system. With regard to the latter potential cause, the authors hypothesized that because there is more natural evening light during the spring than winter, a sample of adolescent students would be more phase delayed in spring than in winter, would have later sleep onset times, and because of fixed school schedules would have shorter sleep durations. Sixteen eighth-grade subjects were recruited for the study. The authors collected sleep logs and saliva samples to determine their dim light melatonin onset (DLMO), a well-established circadian marker. Actual circadian light exposures experienced by a subset of 12 subjects over the course of 7 days in winter and in spring using a personal, head-worn, circadian light measurement device are also reported here. Results showed that this sample of adolescents was exposed to significantly more circadian light in spring than in winter, especially during the evening hours when light exposure would likely delay circadian phase. Consistent with the light data, DLMO and sleep onset times were significantly more delayed, and sleep durations were significantly shorter in spring than in winter. The present ecological study of light, circadian phase, and self-reported sleep suggests that greater access to evening daylight in the spring may lead to sleep restriction in adolescents while attending school. Therefore, lighting schemes that reduce evening light in the spring may encourage longer sleep times in adolescents. (Author correspondence: figuem@rpi.edu)     

Dim Light at Night Increases Immune Function in Nile Grass Rats,a Diurnal Rodent

With the widespread adoption of electrical lighting during the 20th century, human and nonhuman animals became exposed to high levels of light at night for the first time in evolutionary history. This divergence from the natural environment may have significant implications for certain ecological niches because of the important influence light exerts on the circadian system. For example, circadian disruption and nighttime light exposure are linked to changes in immune function. The majority of studies investigating the effects of light exposure and circadian disruption on the immune system use nocturnal rodents. In diurnal species, many hormones and immune parameters vary with secretion patterns 180° out of phase to those of nocturnal rodents. Thus, the authors investigated the effects of nighttime light exposure on immunocompetence in diurnal Nile grass rats (Arvicanthis niloticus). Rats were housed in either standard 14-h light (L):10-h dark (D) cycles with L ～150 lux and D 0 lux or dim light at night (dLAN) cycles of LD 14:10 with L ～150 lux and D 5 lux for 3 wks, then tested for plasma bactericidal capacity, as well as humoral and cell-mediated immune responses. Rats exposed to dLAN showed increased delayed-type hypersensitivity pinna swelling, which is consistent with enhanced cell-mediated immune function. dLAN rats similarly showed increased antibody production following inoculation with keyhole lymphocyte hemocyanin (KLH) and increased bactericidal capacity. Daytime corticosterone concentrations were elevated in grass rats exposed to nighttime dim light, which may have influenced immunological measures. Overall, these results indicate nighttime light affects immune parameters in a diurnal rodent. (Author correspondence: fonken.1@osu.edu)     

Circadian clock properties of fruit flies Drosophila melanogaster exhibiting early and late emergence chronotypes

The role of circadian clocks in timing daily behaviors is widely acknowledged, and while empirical evidence suggests that clock period is correlated with the preferred phase of a rhythmic behavior (chronotype), other clock properties have also been hypothesized to underlie chronotype variation. Here, we report that fruit fly Drosophila melanogaster populations exhibiting evening emergence chronotype (late) are characterized by higher incidence of behavioral arrhythmicity in constant dim light, wider range of entrainment, reduced rates of re-entrainment to simulated jet-lag and higher amplitude of both entrained and free-running rhythms as compared to those exhibiting morning emergence chronotype (early). Our results thus highlight the role of circadian clock properties such as zeitgeber sensitivity, amplitude and coupling in driving chronotype variation.     

CHRONOTYPE,SLEEP LENGTH,AND SCHOOL ACHIEVEMENT OF 11- TO 23-YEAR-OLD STUDENTS IN NORTHERN EUROPEAN RUSSIA

Residing at northern latitudes for long periods of time is associated with sleep disturbances and internal desynchronization, which are considered to be causes of chronic diseases in old age. In children and teenagers, they result in a poor school achievement, psychological problems, and increase in consumption of stimulants. In this paper, we analyze the relationship between both chronotype and sleep length and the variables of age, sex, place of residence, type of settlement (village/city), latitude and longitude of residence, and school achievement of young inhabitants of northern European Russia. We surveyed 1101 children and teenagers between 11 to 23 yrs of age living in four settlements located between 59° and 67° North latitude and 33° and 60° East longitude. The Munich chronotype questionnaire (MCTQ) was used in the study, and all participants were also required to answer a question about their school achievements. An analysis of covariance (ANCOVA) was used to assess the influence of the analyzed factors on sleep length and chronotype. Self-reported sleep length of teenagers depended moderately on age, whereas the place of residence, latitude, and type of settlement only had a weak effect. Chronotype strongly depended on place of residence and longitude; it moderately depended on latitude and age; and it weakly depended on sex and type of settlement. The sleep length of village teenagers was 46?min longer than that of urban teenagers. The authors found a 1?h and 18?min phase delay of the sleep-wake rhythm (as a marker of chronotype) in teenagers moving in the East-West direction and a 16-min delay moving in the South-North direction within one time zone. There was a weak, but significant, positive correlation between chronotype and time of sunrise. There was about a 2-fold stronger influence of chronotype than sleep length on achievement of school children and college students. We conclude that socioeconomic factors exert a significant influence on sleep length and that climatic conditions exert a significant influence on the chronotype of teenagers in the northern latitudes. (Author correspondence: borisenkov@physiol.komisc.ru)     

Chronotype and environmental light exposure in a student population

ABSTRACT

In humans and most other species, changes in the intensity and duration of light provide a critical set of signals for the synchronisation of the circadian system to the astronomical day. The timing of activity within the 24 h day defines an individual’s chronotype, i.e. morning, intermediate or evening type. The aim of this study was to investigate the associations between environmental light exposure, due to geographical location, on the chronotype of university students. Over 6 000 university students from cities in the Northern Hemisphere (Oxford, Munich and Groningen) and Southern Hemisphere (Perth, Melbourne and Auckland) completed the Munich ChronoType Questionnaire. In parallel, light measures (daily irradiance, timing of sunrise and sunset) were compiled from satellite or ground stations at each of these locations. Our data shows that later mid-sleep point on free days (corrected for oversleep on weekends MFS_sc) is associated with (i) residing further from the equator, (ii) a later sunset, (iii) spending more time outside and (iv) waking from sleep significantly after sunrise. However, surprisingly, MSF_sc did not correlate with daily light intensity at the different geographical locations. Although these findings appear to contradict earlier studies suggesting that in the wider population increased light exposure is associated with an earlier chronotype, our findings are derived exclusively from a student population aged between 17 and 26 years. We therefore suggest that the age and occupation of our population increase the likelihood that these individuals will experience relatively little light exposure in the morning whilst encountering more light exposure later in the day, when light has a delaying effect upon the circadian system.     

PACEMAKER PHASE CONTROL VERSUS MASKING BY LIGHT: SETTING THE CIRCADIAN CHRONOTYPE IN DUAL OCTODON DEGUS

There are two main processes involved in the expression of circadian rhythmicity: entrainment and masking. Whereas the first operates via the central pacemaker to anticipate predictable environmental conditions, masking (mainly induced by light) functions as a direct modulator of the circadian output signal induced by nonpredictable events. The Chilean rodent Octodon degus presents both diurnal and nocturnal chronotypes when given free access to an exercise wheel. Two steady-entrainment phases and graded masking by light seem to generate the wide variability of chronotypes in this species. The aim of this study was to characterize the differential masking by light according to the individual chronotypes, their stability over time, and the influence of wheel running availability and ambient temperature upon the degus' nocturnality. To this end, diurnal and nocturnal degus were subjected to ultradian cycles (1:1-h light-dark [LD]), with and without wheel running availability, and under both normal and high diurnal ambient temperature cycles. The present results show that diurnal and nocturnal degus present a stable masking by light, each according to its respective chronotype. Thus, whereas diurnal animals increased their activity with light, in nocturnal degus light induced a sharp drop in wheel running activity. These two types of masking responses appeared not only when the animals were synchronized to the 12:12-h LD cycle, but also under ultradian cycles. Different masking effects persisted when wheel running was made unavailable and when the animals shifted their circadian activity patterns in response to ultradian cycles or to diurnal exposure to high temperatures. In conclusion, our results show that the positive and negative masking effects of light on diurnal and nocturnal degus, respectively, seem to occur independently of relative phase control by the central pacemaker or the negative masking induced by high environmental temperatures. (Author correspondence: angerol@um.es)     

Seasonal Variation of Temporal Niche in Wild Owl Monkeys (Aotus azarai azarai) of the Argentinean Chaco: A Matter of Masking?

Among the more than 40 genera of anthropoid primates (monkeys, apes, and humans), only the South American owl monkeys, genus Aotus, are nocturnal. However, the southernmostly distributed species, Aotus azarai azarai, of the Gran Chaco may show considerable amounts of its 24-h activity during bright daylight. Due to seasonal changes in the duration of photophase and climatic parameters in their subtropical habitat, the timing and pattern of their daily activity are expected to show significant seasonal variation. By quantitative long-term activity recordings with Actiwatch AW4 accelerometer data logger devices of 10 wild owl monkeys inhabiting a gallery forest in Formosa, Argentina, the authors analyzed the seasonal variation in the temporal niche and activity pattern resulting from entrainment and masking of the circadian activity rhythm by seasonally and diurnally varying environmental factors. The owl monkeys always displayed a distinct bimodal activity pattern, with prominent activity bouts and peaks during dusk and dawn. Their activity rhythm showed distinct lunar and seasonal variations in the timing and daily pattern. During the summer, the monkeys showed predominantly crepuscular/nocturnal behavior, and a crepuscular/cathemeral activity pattern with similar diurnal and nocturnal activity levels during the cold winter months. The peak times of the evening and morning activity bouts were more closely related to the times of sunset and sunrise, respectively, than activity-onset and -offset. Obviously, they were better circadian markers for the phase position of the entrained activity rhythm than activity-onset and -offset, which were subject to more masking effects of environmental and/or internal factors. Total daily activity was lowest during the two coldest lunar months, and almost twice as high during the warmest months. Nighttime (21:00–06:00?h) and daytime (09:00–18:00?h) activity varied significantly across the year, but in an opposite manner. Highest nighttime activity occurred in summer and maximal daytime activity during the cold winter months. Dusk and dawn activity, which together accounted for 43% of the total daily activity, barely changed. The monkeys tended to terminate their nightly activity period earlier on warm and rainy days, whereas the daily amount of activity showed no significant correlation either with temperature or precipitation. These data are consistent with the dual-oscillator hypothesis of circadian regulation. They suggest the seasonal variations of the timing and pattern of daily activity in wild owl monkeys of the Argentinean Chaco result from a specific interplay of light entrainment of circadian rhythmicity and strong masking effects of various endogenous and environmental factors. Since the phase position of the monkeys' evening and morning activity peaks did not vary considerably over the year, the seasonal change from a crepuscular/nocturnal activity pattern in summer to a more crepuscular/cathemeral one in winter does not depend on a corresponding phase shift of the entrained circadian rhythm, but mainly on masking effects. Thermoregulatory and energetic demands and constraints seem to play a crucial role. (Author correspondence: hans.erkert@t-online.de)     

Daily Changes in Ultraviolet Light Levels Can Synchronize the Circadian Clock of Bumblebees (Bombus terrestris)

Endogenous circadian clocks are synchronized to the 24-h day by external zeitgebers such as daily light and temperature cycles. Bumblebee foragers show diurnal rhythms under daily light:dark cycles and short-period free-running circadian rhythms in constant light conditions in the laboratory. In contrast, during the continuous light conditions of the arctic summer, they show robust 24-h rhythms in their foraging patterns, meaning that some external zeitgeber must entrain their circadian clocks in the presence of constant light. Although the sun stays above the horizon for weeks during the arctic summer, the light quality, especially in the ultraviolet (UV) range, exhibits pronounced daily changes. Since the photoreceptors and photopigments that synchronize the circadian system of bees are not known, we tested if the circadian clocks of bumblebees (Bombus terrestris) can be entrained by daily cycles in UV light levels. Bumblebee colonies were set up in the laboratory and exposed to 12?h:12?h UV?+?:UV? cycles in otherwise continuous lighting conditions by placing UV filters on their foraging arenas for 12?h each day. The activity patterns of individual bees were recorded using fully automatic radiofrequency identification (RFID). We found that colonies manipulated in such a way showed synchronized 24-h rhythms, whereas simultaneously tested control colonies with no variation in UV light levels showed free-running rhythms instead. The results of our study show that bumblebee circadian rhythms can indeed be synchronized by daily cycles in ambient light spectral composition. (Author correspondence: r.stanewsky@qmul.ac.uk)     

Effects of Temperature on Circadian Clock and Chronotype: An Experimental Study on a Passerine Bird

Daily schedules of many organisms, including birds, are thought to affect fitness. Timing in birds is based on circadian clocks that have a heritable period length, but fitness consequences for individuals in natural environments depend on the scheduling of entrained clocks. This chronotype, i.e., timing of an individual relative to a zeitgeber, results from interactions between the endogenous circadian clock and environmental factors, including light conditions and ambient temperature. To understand contributions of these factors to timing, we studied daily activity patterns of a captive songbird, the great tit (Parus major), under different temperature and light conditions. Birds were kept in a light (L)-dark (D) cycle (12.5?L:11.5 D) at either 8°C or 18°C with ad libitum access to food and water. We assessed chronotype and subsequently tested birds at the same temperature under constant dim light (LL_dim) to determine period length of their circadian clock. Thermal conditions were then reversed so that period length was measured under both temperatures. We found that under constant dim light conditions individuals lengthened their free-running period at higher temperatures by 5.7?±?2.1?min (p?=?.002). Under LD, birds kept at 18°C started activity later and terminated it much earlier in the day than those kept under 8°C. Overall, chronotype was slightly earlier under higher temperature, and duration of activity was shorter. Furthermore, individuals timed their activities consistently on different days under LD and over the two test series under LL_dim (repeatability from .38 to .60). Surprisingly, period length and chronotype did not show the correlation that had been previously found in other avian species. Our study shows that body clocks of birds are precise and repeatable, but are, nonetheless, affected by ambient temperature. (Author correspondence: marina.lehmann@uni-konstanz.de)     

Same Temporal Niche,Opposite Rhythmicity: Two Closely Related Bioluminescent Insects With Opposite Bioluminesce Propensity Rhythms

Arachnocampa species, commonly called glowworms, are flies whose larvae use light to attract prey. Here we compare rhythmicity in two of the nine described species: the Tasmanian species, Arachnocampa tasmaniensis, which inhabits caves and wet forest, and the eastern Australian mainland species, A. flava, primarily found in subtropical rainforest. Both species show the same nocturnal glowing pattern in external (epigean) environments and the same inhibition of bioluminescence by light and both species show circadian regulation of bioluminescence. We find that the underlying circadian bioluminescence propensity rhythm (BPR) of the two species peaks at opposite phases of the day:night cycle. Larvae of A. flava, placed in constant darkness in the laboratory, bioluminesce during the subjective scotophase, typical of nocturnal animals, whereas A. tasmaniensis shows the opposite tendency, bioluminescing most intensely during the subjective photophase. In A. tasmaniensis, which are exposed to natural day:night cycles, light exposure during the day overrides the high bioluminescence propensity through negative masking and leads to a release of bioluminescence after dusk when the BPR is on the wane. A consequence is that A. tasmaniensis is able to start glowing at any phase of the light:dark cycle as soon as masking by light is released, whereas A. flava is locked into nocturnal bioluminescence. We suggest that the paradoxical BPR of A. tasmaniensis is an adaptation for living in the cave environment. Observations of bioluminescence in colonies of A. tasmaniensis located in the transition from a cave mouth to the dark zone show that glowing is inhibited by light exposure but a peak bioluminescence follows immediately after “dusk” at their location. The substantial difference in the circadian regulation of bioluminescence between the two species probably reflects adaptation to the cave (hypogean) habitat in A. tasmaniensis and the forest (epigean) habitat in A. flava. (Author correspondence: d.merritt@uq.edu.au)     

Association Study of a Variable-Number Tandem Repeat Polymorphism in the Clock Gene PERIOD3 and Chronotype in Norwegian University Students

Circadian rhythms are endogenously generated cycles involving physiological parameters, such as core body temperature, hormone levels, blood pressure, sleep, and metabolism, with a period length of around 24?h. The circadian clock in mammals is regulated by a set of clock genes that are functionally linked together, and polymorphisms in clock genes could be associated with differences in circadian rhythms. A variable-number tandem repeat (VNTR) in the human clock gene PERIOD3 (PER3) has been suggested to correlate with a morning (lark) versus evening (owl) chronotype as well as with the circadian rhythm sleep disorder “delayed sleep phase disorder” (DSPD). The authors examined 432 healthy Norwegian university students in search of further support for an association between the PER3 polymorphism and diurnal preference. The Horne-Östberg Morningness-Eveningness Questionnaire (MEQ) and Preferences Scale (PS) were used to evaluate subjective chronotype. DNA samples were genotyped with respect to the 4-repeat and 5-repeat alleles of the VNTR PER3 polymorphism, and the genotype distribution was 192 (4-4), 191 (4-5), and 49 (5-5). The authors estimated that the power to detect an association of the 4-allele with preference for morningness or eveningness was 75%. The authors found no association between the PER3 clock gene and chronotype, indicating that the proposed role of PER3 needs further clarification. (Author correspondence: teresa.osland@med.uib.no)