Seasonal Variation In Day Length Research Articles

Epigenetic‐based control of flowering and seed development in plants: A review

AbstractPlants grown at different latitudes perceive and interpret seasonal variations in day length (photoperiod) and exposure to cold winter temperatures (vernalization). These factors control the expression of various genes involved in flowering, depending on the variations in photoperiod and vernalization. Epigenetic regulatory systems have evolved in plants to process environmental signals. Gene expression is modified through chromatin remodelling and small RNAs in response to seasonal changes in both annual and perennial plants. Key regulators of flowering, such as FLOWERING LOCUS C (FLC) and FLOWERING LOCUS T (FT), interact with other floral regulatory factors and undergo chromatin remodelling in response to seasonal cues. The Polycomb repressive complex (PRC) controls the expression of flowering‐related genes in photoperiodic flowering regulation. FLC acts as a potent suppressor by down‐regulating the expression of genes that promote flowering. Methylation, particularly in the context of CHG, CG and CHH, plays a critical role in embryogenesis. This review briefly explores and describes the regulation of flowering mechanisms in response to day‐length variations, cold exposure (vernalization) and seed development in plants.

Grape-Seed Procyanidin Extract (GSPE) Seasonal-Dependent Modulation of Glucose and Lipid Metabolism in the Liver of Healthy F344 Rats.

Seasonality is gaining attention in the modulation of some physiological and metabolic functions in mammals. Furthermore, the consumption of natural compounds, such as GSPE, is steadily increasing. Consequently, in order to study the interaction of seasonal variations in day length over natural compounds’ molecular effects, we carried out an animal study using photo-sensitive rats which were chronically exposed for 9 weeks to three photoperiods (L6, L18, and L12) in order to mimic the day length of different seasons (winter/summer/and autumn-spring). In parallel, animals were also treated either with GSPE 25 (mg/kg) or vehicle (VH) for 4 weeks. Interestingly, a seasonal-dependent GSPE modulation on the hepatic glucose and lipid metabolism was observed. For example, some metabolic genes from the liver (SREBP-1c, Gk, Acacα) changed their expression due to seasonality. Furthermore, the metabolomic results also indicated a seasonal influence on the GSPE effects associated with glucose-6-phosphate, D-glucose, and D-ribose, among others. These differential effects, which were also reflected in some plasmatic parameters (i.e., glucose and triglycerides) and hormones (corticosterone and melatonin), were also associated with significant changes in the expression of several hepatic circadian clock genes (Bmal1, Cry1, and Nr1d1) and ER stress genes (Atf6, Grp78, and Chop). Our results point out the importance of circannual rhythms in regulating metabolic homeostasis and suggest that seasonal variations (long or short photoperiods) affect hepatic metabolism in rats. Furthermore, they suggest that procyanidin consumption could be useful for the modulation of the photoperiod-dependent changes on glucose and lipid metabolism, whose alterations could be related to metabolic diseases (e.g., diabetes, obesity, and cardiovascular disease). Furthermore, even though the GSPE effect is not restricted to a specific photoperiod, our results suggest a more significant effect in the L18 condition.

Variations of microbial communities and substrate regimes in the eastern Fram Strait between summer and fall.

Seasonal variations in day length and temperature, in combination with dynamic factors such as advection from the North Atlantic, influence primary production and the microbial loop in the Fram Strait. Here, we investigated the seasonal variability of biopolymers, microbial abundance and microbial composition within the upper 100 m during summer and fall. Flow cytometry revealed a shift in the autotrophic community from picoeukaryotes dominating in summer to a 34-fold increase of Synechococcus by fall. Furthermore, a significant decline in biopolymers concentrations covaried with increasing microbial diversity based on 16S rRNA gene sequencing along with a community shift towards fewer polymer-degrading genera in fall. The seasonal succession in the biopolymer pool and microbes indicates distinct metabolic regimes, with a higher relative abundance of polysaccharide-degrading genera in summer and a higher relative abundance of common taxa in fall. The parallel analysis of DOM and microbial diversity provides an important baseline for microbe-substrate relationships over the seasonal cycle in the Arctic Ocean.

Seasonal Variation in the Brain μ-Opioid Receptor Availability.

Seasonal rhythms influence mood and sociability. The brain μ-opioid receptor (MOR) system modulates a multitude of seasonally varying socioemotional functions, but its seasonal variation remains elusive with no previously reported in vivo evidence. Here, we first conducted a cross-sectional study with previously acquired human [11C]carfentanil PET imaging data (132 male and 72 female healthy subjects) to test whether there is seasonal variation in MOR availability. We then investigated experimentally whether seasonal variation in daylength causally influences brain MOR availability in rats. Rats (six male and three female rats) underwent daylength cycle simulating seasonal changes; control animals (two male and one female rats) were kept under constant daylength. Animals were scanned repeatedly with [11C]carfentanil PET imaging. Seasonally varying daylength had an inverted U-shaped functional relationship with brain MOR availability in humans. Brain regions sensitive to daylength spanned the socioemotional brain circuits, where MOR availability peaked during spring. In rats, MOR availabilities in the brain neocortex, thalamus, and striatum peaked at intermediate daylength. Varying daylength also affected the weight gain and stress hormone levels. We conclude that cerebral MOR availability in humans and rats shows significant seasonal variation, which is predominately associated with seasonal photoperiodic variation. Given the intimate links between MOR signaling and socioemotional behavior, these results suggest that the MOR system might underlie seasonal variation in human mood and social behavior.SIGNIFICANCE STATEMENT Seasonal rhythms influence emotion and sociability. The central μ-opioid receptor (MOR) system modulates numerous seasonally varying socioemotional functions, but its seasonal variation remains elusive. Here we used positron emission tomography to show that MOR levels in both human and rat brains show daylength-dependent seasonal variation. The highest MOR availability was observed at intermediate daylengths. Given the intimate links between MOR signaling and socioemotional behavior, these results suggest that the MOR system might underlie seasonal variation in human mood and social behavior.

Seasonal variation of temperature regulation: do thermoregulatory responses “spring” forward and “fall” back?

Seasonal variations in day length and light intensity can affect the circadian rhythm as well as some characteristics of temperature regulation. We investigated characteristics of autonomic (ATR), behavioural (BTR) and nocturnal (NTR) temperature regulation during spring and autumn. Eleven participants underwent experiments in both seasons. To assess ATR, participants performed a 30-min bout of submaximal upright exercise on a cycle ergometer, followed by 100min of water immersion (28°C). Thresholds for the onset of shivering and sweating and vasomotor response were measured. BTR was assessed using a water-perfused suit, with participants regulating the water-perfused suit temperature (Twps) within a range, considered as thermally comfortable. The Twps changed in a saw-tooth manner from 10 to 50°C; by depressing a switch, the direction of the Twps changed, and this limit defined the thermal comfort zone (TCZ) for each participant. A 24-h proximal (calf)-distal (toe) skin temperature gradient (∆Tc-t) was measured to assess NTR. Initiation of vasomotor tone, shivering and sweating was similar between trials. Width of the TCZ was 8.1°C in spring and 8.6°C in autumn (p = 0.1), with similar upper and lower regulated temperatures. ∆Tc-t exhibited a typical circadian rhythm with no difference between seasons. Minor changes in skin temperature and oxygen consumption (p˂0.05) between the seasons may indicate a degree of seasonal adaptation over the course of winter and summer, which persisted in spring and autumn. Other factors, such as country, race, sex and age could however modify the outcome of the study.

Changes in key traits versus depth and latitude suggest energy-efficient locomotion, opportunistic feeding and light lead to adaptive morphologies of marine fishes.

Understanding patterns and processes governing biodiversity along broad-scale environmental gradients, such as depth or latitude, requires an assessment of not just taxonomic richness, but also morphological and functional traits of organisms. Studies of traits can help to identify major selective forces acting on morphology. Currently, little is known regarding patterns of variation in the traits of fishes at broad spatial scales. The aims of this study were (a) to identify a suite of key traits in marine fishes that would allow assessment of morphological variability across broad-scale depth (50-1200m) and latitudinal (29.15-50.91°S) gradients, and (b) to characterize patterns in these traits across depth and latitude for 144 species of ray-finned fishes in New Zealand waters. Here, we describe three new morphological traits, namely fin-base-to-perimeter ratio, jaw-length-to-mouth-width ratio, and pectoral-fin-base-to-body-depth ratio. Four other morphological traits essential for locomotion and food acquisition that are commonly measured in fishes were also included in the study. Spatial ecological distributions of individual fish species were characterized in response to a standardized replicated sampling design, and morphological measurements were obtained for each species from preserved museum specimens. With increasing depth, fishes, on average, became larger and more elongate, with higher fin-base-to-perimeter ratio and larger jaw-length-to-mouth-width ratio, all of which translates into a more eel-like anguilliform morphology. Variation in mean trait values along the depth gradient was stronger at lower latitudes for fin-base-to-perimeter ratio, elongation and total body length. Average eye size peaked at intermediate depths (500-700m) and increased with increasing latitude at 700m. These findings suggest that, in increasingly extreme environments, fish morphology shifts towards a body shape that favours an energy-efficient undulatory swimming style and an increase in jaw-length vs. mouth width for opportunistic feeding. Furthermore, increases in eye size with both depth and latitude indicate that changes in both the average ambient light conditions as well as seasonal variations in day-length can act to select ecomorphological adaptations in fishes.

Effects of photoperiod on life-history and thermal stress resistance traits across populations of Drosophila subobscura.

IntroductionOrganisms use environmental cues to match their phenotype with the future availability of resources and environmental conditions. Changes in the magnitude and frequency of environmental cues such as photoperiod and temperature along latitudes can be used by organisms to predict seasonal changes. While the role of temperature variation on the induction of plastic and seasonal responses is well established, the importance of photoperiod for predicting seasonal changes is less explored.Materials and methodsHere we studied changes in life‐history and thermal stress resistance traits in Drosophila subobscura in response to variation in photoperiod (6:18, 12:12 and 18:6 light:dark cycles) mimicking seasonal variations in day length. The populations of D. subobscura were collected from five locations along a latitudinal gradient (from North Africa and Europe). These populations were exposed to different photoperiods for two generations, whereafter egg‐to‐adult viability, productivity, dry body weight, thermal tolerance, and starvation resistance were assessed.ResultsWe found strong effects of photoperiod, origin of populations, and their interactions on life‐history and stress resistance traits. Thermal resistance varied between the populations and the effect of photoperiod depended on the trait and the method applied for the assessment of thermal resistance.PerspectivesOur results show a strong effect of the origin of population and photoperiod on a range of fitness‐related traits and provide evidence for local adaptation to environmental cues (photoperiod by population interaction). The findings emphasize an important and often neglected role of photoperiod in studies on thermal resistance and suggest that cues induced by photoperiod may provide some buffer enabling populations to cope with a more variable and unpredictable future climate.

Assortative mating by flowering time and its effect on correlated traits in variable environments.

Reproductive timing is a key life‐history trait that impacts the pool of available mates, the environment experienced during flowering, and the expression of other traits through genetic covariation. Selection on phenology, and its consequences on other life‐history traits, has considerable implications in the context of ongoing climate change and shifting growing seasons. To test this, we grew field‐collected seed from the wildflower Mimulus guttatus in a greenhouse to assess the standing genetic variation for flowering time and covariation with other traits. We then created full‐sib families through phenological assortative mating and grew offspring in three photoperiod treatments representing seasonal variation in daylength. We find substantial quantitative genetic variation for the onset of flowering time, which covaried with vegetative traits. The assortatively‐mated offspring varied in their critical photoperiod by over two hours, so that families differed in their probability of flowering across treatments Allocation to flowering and vegetative growth changed across the daylength treatments, with consistent direction and magnitude of covariation among flowering time and other traits. Our results suggest that future studies of flowering time evolution should consider the joint evolution of correlated traits and shifting seasonal selection to understand how environmental variation influences life histories.

Larks, owls, swifts, and woodcocks among fruit flies: differential responses of four heritable chronotypes to long and hot summer days

PurposeDrosophila melanogaster and our own species share (Homo sapiens) the history of relatively rapid out-of-Africa dispersal. In Eurasia, they had faced a novel adaptive problem of adjustment of their circadian rhythmicity and night sleep episode to seasonal variation in day length and air temperature. Both species usually respond to heat and a short duration of night by reduction of the amount of night sleep and prolongation of “siesta”. To further explore similarities between the two species in the ways of adjustment of their sleep–wake behavior to extreme environmental factors, this study examined the possibility to distinguish four extreme chronotypes among fruit flies and the possibility of the differential response of such chronotypes to light and heat stressors.Materials and methodsCircadian rhythms of locomotor activity and sleep–wake pattern were tested in constant darkness, and four strains of fruit flies originating from three wild populations of Africa, Europe, and the USA were selected to represent four distinct chronotypes: “larks” (early morning and evening activity peaks), “owls” (late morning and evening peaks), “swifts” (early morning and late evening peaks), and “woodcocks” (late morning and early evening peaks). The circadian rhythms and sleep efficiency of the selected chronotypes were further tested under such extreme conditions as either long day (LD20:4 at 20°C) or a combination of LD20:4 with hot temperature (29°C).ResultsDespite the identity of such experimental conditions for four chronotypes, their circadian rhythms and sleep timing showed significantly distinct patterns of response to exposure to heat and/or long days. All two-way repeated measures analysis of variances yielded a significant interaction between chronotype and time of the day (P<0.001).ConclusionAn experimental study of heritable chronotypes in the fruit fly can facilitate a search for genetic underpinnings of individual variation in vulnerability to circadian misalignment, maladaptive sleep–wake behavior, and sleep disorders.

Anthropogenic noise, but not artificial light levels predicts song behaviour in an equatorial bird.

Birds in cities start singing earlier in the morning than in rural areas; commonly this shift is attributed to light pollution. Some studies have suggested that traffic noise has a stronger influence on singing activity than artificial light does. Changes in the timing of singing behaviour in relation to noise and light pollution have only been investigated in the temperate zones. Tropical birds, however, experience little seasonal variation in day length and may be less dependent on light intensity as a modifier for reproductive behaviours such as song. To test whether noise or light pollution has a stronger impact on the dawn chorus of a tropical bird, we investigated the singing behaviour of rufous-collared sparrows (Zonotrichia capensis) in Bogota, Colombia at two times during the year. We found that birds in places with high noise levels started to sing earlier. Light pollution did not have a significant effect. Birds may begin to sing earlier in noisy areas to avoid acoustic masking by traffic later in the morning. Our results also suggest that some tropical birds may be less sensitive to variations in day length and thus less sensitive to light pollution.

Ecological effects of artificial light at night on wild plants

Summary Plants use light as a source of both energy and information. Plant physiological responses to light, and interactions between plants and animals (such as herbivory and pollination), have evolved under a more or less stable regime of 24‐h cycles of light and darkness, and, outside of the tropics, seasonal variation in day length. The rapid spread of outdoor electric lighting across the globe over the past century has caused an unprecedented disruption to these natural light cycles. Artificial light is widespread in the environment, varying in intensity by several orders of magnitude from faint skyglow reflected from distant cities to direct illumination of urban and suburban vegetation. In many cases, artificial light in the night‐time environment is sufficiently bright to induce a physiological response in plants, affecting their phenology, growth form and resource allocation. The physiology, behaviour and ecology of herbivores and pollinators are also likely to be impacted by artificial light. Thus, understanding the ecological consequences of artificial light at night is critical to determine the full impact of human activity on ecosystems. Synthesis. Understanding the impacts of artificial night‐time light on wild plants and natural vegetation requires linking the knowledge gained from over a century of experimental research on the impacts of light on plants in the laboratory and glasshouse with knowledge of the intensity, spatial distribution, spectral composition and timing of light in the night‐time environment. To understand fully the extent of these impacts requires conceptual models that can (i) characterize the highly heterogeneous nature of the night‐time light environment at a scale relevant to plant physiology; and (ii) scale physiological responses to predict impacts at the level of the whole plant, population, community and ecosystem.

Seasonal and urban effects on the endocrinology of a wild passerine.

In order to maximize their fitness, organisms in seasonal environments rely on external cues to optimally time their life‐history stages. One of the most important zeitgeber to time reproduction is the photoperiod, but further environmental cues are assessed to fine‐tune reproduction due to year‐to‐year variation in environmental conditions. However, in urbanized environments, the pervasive artificial light at night has altered the natural signal of light and darkness. Accordingly, artificial light at night was repeatedly shown to affect avian reproductive physiology and to advance seasonal reproduction in birds. However, these experiments were mainly conducted in the absence of further environmental cues to facilitate the investigation of the mechanisms which are still poorly understood. Here, we investigate whether the endocrine system of free‐ranging European blackbirds (Turdus merula) correlates with the amount of artificial light at night along a rural to urban gradient while the birds still encounter complementary environmental cues including seasonal variation in day length and temperature. Testosterone and estrone were assessed as metabolites in fecal samples and corticosterone in blood from mist‐netted blackbirds. We demonstrate that seasonal fluctuations in abiotic factors, individual conditions, but also light at night affect the reproductive and stress physiology of wild European blackbirds. Elevated artificial night light intensities were significantly positively correlated with corticosterone and negatively with female estrone levels. No effects of artificial light were found for testosterone levels. Our results suggest that female blackbirds in particular perceive even low levels of artificial light at night as a weak but chronic stressor that interacts with the hypothalamic‐pituitary‐gonadal axis and leads to a reduced secretion of reproductive hormones. These findings point out that the impacts of light pollution are diverse and we only slowly disentangle its multiple effects on physiology, ecology, and biodiversity.

Seasonally different reproductive investment in a medium-sized rodent (Cavia aperea)

Pronounced seasonal variations in day length, temperature, and resource availability characterize the temperate regions and strongly influence the animals living in these environments. To survive and reproduce successfully, animals must allocate resources among competing physiological systems, and they usually adjust their time of breeding to the most adequate season. Here, we examined whether reproductive investment in the wild guinea pig (Cavia aperea) differs across seasons. We kept animals in combined indoor–outdoor enclosures under natural light and temperature year-round. We measured littering probability, litter size, and birth weight, as well as maternal weight loss during lactation. In addition, we measured ovulation rate as a parameter to adjust reproductive investment prenatally. Our data reveal strong seasonal variations in reproductive traits despite the fact that the animals reproduced year-round. The results show a reduced reproductive investment in winter, indicated by a lower litter size and birth weight of pups, whereas investment was highest in warm seasons (summer and autumn) with higher litter size and birth weight. Maternal weight loss in lactation was highest in cold seasons even if the litter size was lower. Furthermore, we found the regulation on the proximate level of the reproductive investment, the ovulation rate, to differ significantly between the seasons.

Photoperiod modulation of aggressive behavior is independent of androgens in a tropical cichlid fish

Photoperiod is a major environmental cue that signals breeding conditions in animals living in temperate climates. Therefore, the activity of the reproductive (i.e. hypothalamic–pituitary–gonadal, HPG) axis and of the expression of reproductive behaviors, including territoriality, is responsive to changes in day length. However, at low latitudes the seasonal variation in day length decreases dramatically and photoperiod becomes less reliable as a breeding entraining cue in tropical species. In spite of this, some tropical mammals and birds have been found to still respond to small amplitude changes in photoperiod (e.g. 17min). Here we tested the effect of 2 photoperiod regimes, referred to as long-day (LD: 16L:08D) and short-day (SD: 08L:16D), on the activity of the HPG axis, on aggressive behavior and in the androgen response to social challenges in males of the tropical cichlid fish Tilapia rendalli. For each treatment, fish were transferred from a pre-treatment photoperiod of 12L:12D to their treatment photoperiod (either LD or SD) in which they were kept for 20days on stock tanks. Afterwards, males were isolated for 4days in glass aquaria in order to establish territories and initial androgen levels (testosterone, T; 11-ketotestosterone, KT) were assessed. On the 4th day, territorial intrusions were promoted such that 1/3 of the isolated males acted as residents and another 1/3 as intruders. Territorial intrusions lasted for 1h to test the effects of a social challenge under different photoperiod regimes. Photoperiod treatment (either SD or LD) failed to induce significant changes in the HPG activity, as measured by androgen levels and gonadosomatic index. However, SD increased the intensity of aggressive behaviors and shortened the time to settle a dominance hierarchy in an androgen-independent manner. The androgen responsiveness to the simulated territorial intrusion was only present in KT but not for T. The percent change in KT levels in response to the social challenge was different between treatments (SD>LD) and between male types (resident>intruder). The higher androgen response to a social challenge in residents under SD may be explained by the time course of the androgen response that due to the long time it takes to fight resolution under LD, might have been delayed. This result illustrates the importance of incorporating time response data in social endocrinology studies.

Seasonal wing colour plasticity varies dramatically between buckeye butterfly populations in different climatic zones

1. In the present study, how seasonal wing colour plasticity in buckeye butterflies (Junonia coenia Hübner) differs between populations in different climatic regions, and whether these differences are explained by regional environmental differences or reflect genetic divergences in plasticity is addressed.2. Using museum specimen data, it is shown that buckeye wing colour variation is much greater in populations from the humid and subtropical climates of the eastern U.S.A. than in populations from the desert and mediterranean climates of the western U.S.A.3. Museum specimen data further show that wing colour in eastern populations is strongly correlated with seasonal variation in day length and temperature, whereas wing colour in western populations is only weakly associated with these cues.4. Controlled incubator experiments comparing Southern California and North Carolina populations suggest that regional differences in wing pattern variation are attributable to dramatically increased robustness to environmental variation in western populations.5. Together these experiments show that while phenotypic variation can be influenced by environmental cues, the range of this variation can genetically diverge between populations in different regions.

Associations between seasonal variations in day length (photoperiod), sleep timing, sleep quality and mood: a comparison between Ghana (5°) and Norway (69°)

The hypothesis of whether day length (photoperiod) is an important zeitgeber (time-giver) for keeping the circadian rhythm entrained to a 24-hour cycle was examined, as was its association with sleep patterns and mood problems. Data were collected prospectively from a site with very large differences in daylight duration across seasons (Tromsø in Norway, 69°39'N) and a site with very small seasonal differences in daylight duration (Ghana in Accra, 5°32'N). Two hundred subjects were recruited from both sites in January. At the follow-up in August, 180 and 150 subjects in Ghana and Norway participated, respectively. Use of a weekly sleep diary indicated low to moderately strong seasonal changes in rise- and bedtime, sleep efficiency and sleep onset latency only in the northern latitude. No seasonal changes in sleep duration or night awakenings were observed. The self-report measures indicated moderate to strong seasonal differences in insomnia and fatigue, and weaker differences in depressed mood in Norway, but small to non-existing seasonal differences in Ghana. Lack of daylight was related to phase-delayed rise- and bedtimes, increased problems falling asleep, daytime fatigue and depressive mood. However, total sleep duration and sleep quality appeared unaffected.

Relationship between daylength and suicide in Finland

BackgroundMany previous studies have documented seasonal variation in suicides globally. We re-assessed the seasonal variation of suicides in Finland and tried to relate it to the seasonal variation in daylength and ambient temperature and in the discrepancy between local time and solar time.MethodsThe daily data of all suicides from 1969 to 2003 in Finland (N = 43,393) were available. The calendar year was divided into twelve periods according to the length of daylight and the routinely changing time difference between sun time and official time. The daily mean of suicide mortality was calculated for each of these periods and the 95% confidence intervals of the daily means were used to evaluate the statistical significance of the means. In addition, daily changes in sunshine hours and mean temperature were compared to the daily means of suicide mortality in two locations during these afore mentioned periods.ResultsA significant peak of the daily mean value of suicide mortality occurred in Finland between May 15th and July 25th, a period that lies symmetrically around the solstice. Concerning the suicide mortality among men in the northern location (Oulu), the peak was postponed as compared with the southern location (Helsinki). The daily variation in temperature or in sunshine did not have significant association with suicide mortality in these two locations.ConclusionsThe period with the longest length of the day associated with the increased suicide mortality. Furthermore, since the peak of suicide mortality seems to manifest later during the year in the north, some other physical or biological signals, besides the variation in daylight, may be involved. In order to have novel means for suicide prevention, the assessment of susceptibility to the circadian misalignment might help.

Evolution of time-keeping mechanisms: early emergence and adaptation to photoperiod

Virtually all species have developed cellular oscillations and mechanisms that synchronize these cellular oscillations to environmental cycles. Such environmental cycles in biotic (e.g. food availability and predation risk) or abiotic (e.g. temperature and light) factors may occur on a daily, annual or tidal time scale. Internal timing mechanisms may facilitate behavioural or physiological adaptation to such changes in environmental conditions. These timing mechanisms commonly involve an internal molecular oscillator (a 'clock') that is synchronized ('entrained') to the environmental cycle by receptor mechanisms responding to relevant environmental signals ('Zeitgeber', i.e. German for time-giver). To understand the evolution of such timing mechanisms, we have to understand the mechanisms leading to selective advantage. Although major advances have been made in our understanding of the physiological and molecular mechanisms driving internal cycles (proximate questions), studies identifying mechanisms of natural selection on clock systems (ultimate questions) are rather limited. Here, we discuss the selective advantage of a circadian system and how its adaptation to day length variation may have a functional role in optimizing seasonal timing. We discuss various cases where selective advantages of circadian timing mechanisms have been shown and cases where temporarily loss of circadian timing may cause selective advantage. We suggest an explanation for why a circadian timing system has emerged in primitive life forms like cyanobacteria and we evaluate a possible molecular mechanism that enabled these bacteria to adapt to seasonal variation in day length. We further discuss how the role of the circadian system in photoperiodic time measurement may explain differential selection pressures on circadian period when species are exposed to changing climatic conditions (e.g. global warming) or when they expand their geographical range to different latitudes or altitudes.

Seasonal variation in daily insolation induces synchronous bud break and flowering in the tropics

In many temperate plants seasonal variation in day length induces flowering at species-specific times each year. Here we report synchronous bud break and flowering of tropical perennials that cannot be explained by seasonal changes in day length. We recorded flushing and flowering of more than 100 tropical trees, succulents and understory herbs over several years. We observed the following phenological patterns throughout the northern Neotropics: wide-ranging trees flush or flower twice a year at the Equator, but annually further north; many trees leaf out in February; in autumn, wide-ranging perennials flower 4 months earlier in Mexico than at the Equator. This latitudinal variation of phenology parallels that of the annual cycle of daily insolation, a function of day length and solar irradiation. Insolation has two annual maxima at the Equator, it rapidly increases in February at all latitudes, and between Mexico and the Equator its maximum shifts from the summer solstice to the autumn equinox. These unique, manifold correlations suggest that throughout the tropics insolation, rather than day length, may control the phenology of many perennials. Our observations significantly extend current knowledge of environmental signals involved in photoperiodic control of plant development.

Current knowledge on the photoneuroendocrine regulation of reproduction in temperate fish species

Seasonality is an important adaptive trait in temperate fish species as it entrains or regulates most physiological events such as reproductive cycle, growth profile, locomotor activity and key life-stage transitions. Photoperiod is undoubtedly one of the most predictable environmental signals that can be used by most living organisms including fishes in temperate areas. This said, however, understanding of how such a simple signal can dictate the time of gonadal recruitment and spawning, for example, is a complex task. Over the past few decades, many scientists attempted to unravel the roots of photoperiodic signalling in teleosts by investigating the role of melatonin in reproduction, but without great success. In fact, the hormone melatonin is recognized as the biological time-keeping hormone in fishes mainly due to the fact that it reflects the seasonal variation in daylength across the whole animal kingdom rather than the existence of direct evidences of its role in the entrainment of reproduction in fishes. Recently, however, some new studies clearly suggested that melatonin interacts with the reproductive cascade at a number of key steps such as through the dopaminergic system in the brain or the synchronization of the final oocyte maturation in the gonad. Interestingly, in the past few years, additional pathways have become apparent in the search for a fish photoneuroendocrine system including the clock-gene network and kisspeptin signalling and although research on these topics are still in their infancy, it is moving at great pace. This review thus aims to bring together the current knowledge on the photic control of reproduction mainly focusing on seasonal temperate fish species and shape the current working hypotheses supported by recent findings obtained in teleosts or based on knowledge gathered in mammalian and avian species. Four of the main potential regulatory systems (light perception, melatonin, clock genes and kisspeptin) in fish reproduction are reviewed.