Photoperiodic Entrainment Research Articles

Transcriptomic Plasticity of the Circadian Clock in Response to Photoperiod: A Study in Male Melatonin-Competent Mice.

Seasonal daylength, or circadian photoperiod, is a pervasive environmental signal that profoundly influences physiology and behavior. In mammals, the central circadian clock resides in the suprachiasmatic nuclei (SCN) of the hypothalamus where it receives retinal input and synchronizes, or entrains, organismal physiology and behavior to the prevailing light cycle. The process of entrainment induces sustained plasticity in the SCN, but the molecular mechanisms underlying SCN plasticity are incompletely understood. Entrainment to different photoperiods persistently alters the timing, waveform, period, and light resetting properties of the SCN clock and its driven rhythms. To elucidate novel candidate genes for molecular mechanisms of photoperiod plasticity, we performed RNA sequencing on whole SCN dissected from mice raised in long (light:dark [LD] 16:8) and short (LD 8:16) photoperiods. Fewer rhythmic genes were detected in mice subjected to long photoperiod, and in general, the timing of gene expression rhythms was advanced 4-6 h. However, a few genes showed significant delays, including Gem. There were significant changes in the expression of the clock-associated gene Timeless and in SCN genes related to light responses, neuropeptides, gamma aminobutyric acid (GABA), ion channels, and serotonin. Particularly striking were differences in the expression of the neuropeptide signaling genes Prokr2 and Cck, as well as convergent regulation of the expression of 3 SCN light response genes, Dusp4, Rasd1, and Gem. Transcriptional modulation of Dusp4 and Rasd1 and phase regulation of Gem are compelling candidate molecular mechanisms for plasticity in the SCN light response through their modulation of the critical NMDAR-MAPK/ERK-CREB/CRE light signaling pathway in SCN neurons. Modulation of Prokr2 and Cck may critically support SCN neural network reconfiguration during photoperiodic entrainment. Our findings identify the SCN light response and neuropeptide signaling gene sets as rich substrates for elucidating novel mechanisms of photoperiod plasticity. Data are also available at http://circadianphotoperiodseq.com/, where users can view the expression and rhythmic properties of genes across these photoperiod conditions.

Revealing the profound influence of diapause on gene expression: Insights from the annual transcriptome of the copepod Calanus finmarchicus.

Annual rhythms are observed in living organisms with numerous ecological implications. In the zooplanktonic copepod Calanus finmarchicus, such rhythms are crucial regarding its phenology, body lipid accumulation, and global carbon storage. Climate change drives annual biological rhythms out of phase with the prevailing environmental conditions with yet unknown but potentially catastrophic consequences. However, the molecular dynamics underlying phenology are still poorly described. In a rhythmic analysis of C. finmarchicus annual gene expression, results reveal that more than 90% of the transcriptome shows significant annual rhythms, with abrupt and dramatic upheaval between the active and diapause life cycle states. This work explores the implication of the circadian clock in the annual timing, which may control epigenetic mechanisms to profoundly modulate gene expression in response to calendar time. Results also suggest an increased light sensitivity during diapause that would ensure the photoperiodic entrainment of the endogenous annual clock.

Optogenetic stimulation of VIPergic SCN neurons induces photoperiodic-like changes in the mammalian circadian clock.

Circadian clocks play key roles in how organisms respond to and even anticipate seasonal change in day length, or photoperiod. In mammals, photoperiod is encoded by the central circadian pacemaker in the brain, the suprachiasmatic nucleus (SCN). The subpopulation of SCN neurons that secrete the neuropeptide VIP mediates the transmission of light information within the SCN neural network, suggesting a role for these neurons in circadian plasticity in response to light information that has yet to be directly tested. Here, we used in vivo optogenetic stimulation of VIPergic SCN neurons followed by ex vivo PERIOD 2::LUCIFERASE (PER2::LUC) bioluminescent imaging to test whether activation of this SCN neuron subpopulation can induce SCN network changes that are hallmarks of photoperiodic encoding. We found that optogenetic stimulation designed to mimic a long photoperiod indeed altered subsequent SCN entrained phase, increased the phase dispersal of PER2 rhythms within the SCN network, and shortened SCN free-running period-similar to the effects of a true extension of photoperiod. Optogenetic stimulation also induced analogous changes on related aspects of locomotor behaviour in vivo. Thus, selective activation of VIPergic SCN neurons induces photoperiodic network plasticity in the SCN that underpins photoperiodic entrainment of behaviour.

Nurr1 deficiency shortens free running period, enhances photoentrainment to phase advance, and disrupts circadian cycling of the dopamine neuron phenotype

Neurological and neuropsychiatric disorders, including addiction, schizophrenia, and Parkinson’s disease (PD), involve dysfunction in midbrain dopamine (DA) neurotransmission with severity of disease symptoms and progression associated with disrupted circadian rhythms. The nuclear transcription factor Nurr1, essential for DA neuron (DAN) development, survival, and maintenance, is also known to interact with circadian rhythm regulating clock proteins. In the Nurr1-null heterozygous (+/-) mice, a Nurr1 deficient model which reproduces some of the alterations in DA function found in schizophrenia and PD, we measured, using wheel-running activity, the free running period (tau) and photoperiod entrainment. Because Nurr1 has a role in regulating the DA phenotype, we also measured the circadian fluctuations in the number of DANs using tyrosine hydroxylase (TH) immunofluorescence. In Nurr1 +/- mice, tau was significantly shorter and entrainment to a 6 h earlier shift in the dark cycle was accelerated. The Nurr1 wild-type (+/+) mice cycled DAN numbers across time, with a significantly greater number (∼2-fold increase) of DANs at zeitgeber time (ZT) 0 than ZT12. The +/- mice, however, did not cycle the DA phenotype, as no differences in DAN numbers were observed between ZT0 and ZT12. Additionally, the +/- mice had significantly fewer DANs at ZT0 but not at ZT12 as compared to +/+ mice. Based these data, circadian rhythms and fluctuations in the DA phenotype requires normal Nurr1 function. A better understanding is needed of the mechanisms regulating the DA phenotype and subsequent neurotransmission across the circadian cycle and how this is altered in circadian rhythm and DA neurotransmission-associated disorders.

Different as night and day: Behavioural and life history responses to varied photoperiods in Daphnia magna.

Nearly all animal species have utilized photoperiod to cue seasonal behaviours and life history traits. We investigated photoperiod responses in keystone species, Daphnia magna, to identify molecular processes underlying ecologically important behaviours and traits using functional transcriptomic analyses. Daphnia magna were photoperiod‐entrained immediately posthatch to a standard control photoperiod of 16 light/ 8 dark hours (16L:8D) relative to shorter (4L:20D, 8L:16D, 12L:12L) and longer (20L:4D) day length photoperiods. Short‐day photoperiods induced significantly increased light‐avoidance behaviours relative to controls. Correspondingly, significant differential transcript expression for genes involved in glutamate signalling was observed, a critical signalling pathway in arthropod light‐avoidance behaviour. Additionally, period circadian protein and proteins coding F‐box/LRR‐repeat domains were differentially expressed which are recognized to establish circadian rhythms in arthropods. Indicators of metabolic rate increased in short‐day photoperiods which corresponded with broadscale changes in transcriptional expression across system‐level energy metabolism pathways. The most striking observations included significantly decreased neonate production at the shortest day length photoperiod (4L:20D) and significantly increased male production across short‐day and equinox photoperiods (4L:20D, 8L:16D and 12L:12D). Transcriptional expression consistent with putative mechanisms of male production was observed including photoperiod‐dependent expression of transformer‐2 sex‐determining protein and small nuclear ribonucleoprotein particles (snRNPs) which control splice variant expression for genes like transformer. Finally, increased transcriptional expression of glutamate has also been shown to induce male production in Daphnia pulex via photoperiod‐sensitive mechanisms. Overall, photoperiod entrainment affected molecular pathways that underpin critical behavioural and life history traits in D. magna providing fundamental insights into biological responses to this primary environmental cue.

Preliminary observations on the effect of light and temperature on the hatching success and rate of Lepidurus arcticus eggs

Dormancy, which arrests development, is a well-known survival strategy among animals living in the Arctic to overcome harsh periods. It is not clear if the dormant state in notostracans is controlled endogenously (diapause) or exogenously (quiescence). For Lepidurus arcticus, it is unknown how it responds to the photoperiod entrainment, if it has a biological clock and if it has a rhythmic expression of the clock genes. We studied the hatching success of resting eggs at four constant temperatures (5, 10, 15 and 25 °C) and under different illumination regimes [continuous light (LL) and continuous dark (DD)]. It was assumed that light and temperature are both important triggers, with temperature having the most pronounced effect. In our experiment, hatching occurred only at 5 and 10 °C, while we did not observe hatching at 15 and 25 °C. The highest percentage of eggs hatched was at 10 °C in LL (60%); the lowest was at 5 °C in DD (18%). The percentages hatched at 5 °C in LL (24%) and at 10 °C in DD (26%) were similar. Our results indicate that both temperature and light had a significant and interacting effect on hatching in L. arcticus, with temperature being the dominant factor controlling the process. This suggests that changes in temperature affecting the Arctic may significantly impact phenology of this key species in the region. Given that no hatching was observed at 15 °C or above, the persistence of this species may be at risk in areas were arctic lakes are expected to warm to such levels during the summer months.

Shedding new light upon circadian emergence rhythmicity in the mountain pine beetle (Coleoptera: Curculionidae: Scolytinae)

AbstractThe phenological behaviours of temperate insects can be highly controlled by photoperiod. Some foundational studies of the mountain pine beetle, Dendroctonus ponderosae (Hopkins) (Coleoptera: Curculionidae), documented a diurnal emergence rhythm that was asynchronous with maximum daily temperatures in the field and persisted under constant temperature and light conditions. In the 1970s, researchers hypothesised that this emergence rhythm was regulated by an endogenous circadian mechanism. Reflecting upon these historical data, we consider that a diurnal pattern of D. ponderosae emergence may result from photoperiodic entrainment of the circadian clock during the immature stages. Mechanistically, we suggest that the long-wavelength-sensitive opsin that we previously found to be expressed across D. ponderosae life stages could mediate, from beneath the bark, the input of light–dark cycle cues that are usually required for entrainment of the insect circadian clock.

Melatonin-independent Photoperiodic Entrainment of the Circannual TSH Rhythm in the Pars Tuberalis of the European Hamster.

Adaptation of biological rhythms to a seasonal environment in circannual mammals is achieved via the synchronization of intrinsic circannual rhythms to the external year by photoperiod. In mammals, the photoperiodic information is integrated to seasonal physiology via the pineal hormone melatonin regulation of pars tuberalis (PT) TSHβ expression and its downstream control of hypothalamic dio2 gene expression. In the circannual European hamster, however, photoperiodic entrainment of the circannual clock is possible in pinealectomized animals. The present study explores whether the TSHβ expression in the PT and the downstream hypothalamic pathways are regulated by photoperiod in European hamsters in the absence of melatonin. All animals were kept on an accelerated photoperiodic regime, which compressed the natural year to a 6-month cycle. Sham-operated European hamsters and half of the pinealectomized European hamsters entrained their annual cycle in reproduction, body weight, and activity pattern to this cycle, whereas the other half of the pinealectomized animals followed only each second cycle. In all animals, PT TSHβ and hypothalamic dio2 expressions were higher in hamsters displaying a summer physiological state than in those in winter state. Moreover, in agreement with their seasonal state, reproductive animals (summer state) showed higher expression of rfrp and lower expression of kiss1-genes encoding central regulators of the reproductive axis-than those animals in reproductive quiescence (winter state), indicating the hypothalamic integration of the photoperiodic signal even in pinealectomized animals. The appropriate occurrence of a well-characterized activity pattern indicative of a so-called sensitive phase to short photoperiod suggested that the SCN constructs the melatonin-independent photoperiodic message. This message is sufficient to entrain the circannual rhythm in TSHβ expression in the PT and the downstream hypothalamic neuroendocrine pathway through a yet unknown pathway. These results reinforce the hypothesis that the PT is the site for the integration of circannual and photoperiodic information.

Modeling the photoperiodic entrainment of the plant circadian clock

The circadian clock is an endogenous 24 hour rhythm that helps organisms anticipate and adapt to daily and seasonal variations in environment, such as the day/night cycle or changing temperatures. The plant clock is a complex network of transcription factors that regulate each other, forming interlocked feedback loops. Most of its components are light-regulated in some way, making the system highly sensitive to changes in light conditions. Here, we explore the mechanisms by which the plant clock adapts to changing day length. We first present some experimental data illustrating the variety of behaviors found in seedlings exposed to external day/night cycles different from 24h. We then use a mathematical model to characterize the response of the clock to a wide range of external cycle lengths and photoperiods. We show the existence of several domains of periodic entrainment with different ratios between the external cycle length and the period of the clock, and the presence of quasiperiodic and chaotic behaviors outside of the entrainment range. We simulate knockout mutants with impaired clock function and theoretical variants with diminished light sensitivity to highlight the role of a complex network and multiple light inputs in keeping the clock entrained over a wide range of conditions.

GABAergic Neurotransmission in the Premammillary Nucleus of the Turkey Hypothalamus Regulates Reproductive Seasonality and the Onset of Photorefractoriness

Background/Aims: Photoperiod is a major environmental cue in temperate-zone birds which synchronizes breeding with the time of year that offers the optimal environment for offspring survival. Despite continued long photoperiods, these birds eventually become refractory to the stimulating photoperiod and their reproductive systems regress. In this study, we characterized the role of γ-aminobutyric acid (GABA)ergic neurotransmission in modulating the response of the premammillary nucleus (PMM) to a gonad stimulatory photoperiod and the onset of photorefractoriness. Methods and Results: Bilateral ablation of the PMM blocked the light-induced neuroendocrine response from occurring in photosensitive turkeys. Microarray analyses revealed an increase in GABAergic activity in the PMM of photorefractory birds as opposed to photosensitive ones, and this enhanced GABAergic activity appeared to inhibit the photoperiodic signal. Additionally, GABA_A and GABA_B receptors were expressed by dopamine-melatonin neurons in the PMM, and the administration of the GABA receptor agonist baclofen blocked the photoperiodic reproductive neuroendocrine responses. Conclusions: Consistent with the present findings, we propose that the long-sought-after mechanism underlying photorefractoriness is linked to the inhibitory actions of GABA. We suggest that (1) GABAergic interference with photoperiodic entrainment in the PMM initiates the photorefractory state and terminates the annual breeding season in temperate-zone birds, and (2) the PMM is a site of photoreception and photorefractoriness that controls the initiation and termination of avian reproductive seasonality.

Small Geographic Variation in Photoperiodic Entrainment of the Circannual Rhythm in the Varied Carpet Beetle,Anthrenus verbasci

The circannual pupation rhythm of Anthrenus verbasci is entrained to an environmental cycle by changes in photoperiod. Exposure of larvae reared under short-day conditions to long days induced a clear phase delay of the circannual rhythm. There was no notable difference in the initial phase or period of the circannual rhythm among four geographically distinct populations of A. verbasci in Japan: Takanabe (32.1°N), Osaka (34.7°N), Sendai (38.3°N), and Sapporo (43.1°N) populations. The range of photoperiodic changes effective for phase delay in the circannual pupation rhythm was compared among the four populations. Although larvae did not show a typical threshold response, but responded quantitatively to the photophase duration in intermediate conditions, the critical daylengths were calculated as those under which the pupation was delayed by 50%: 12.8 h in the Takanabe population, 13.2 h in the Osaka population, and 13.6 h in the Sendai and Sapporo populations. Thus, the critical daylength for entrainment of the circannual rhythm in A. verbasci was correlated to the habitat latitude, but the differences among the populations were much smaller than those reported in photoperiodism for induction of diapause in various insects. Consequently, the difference in the pupation time among the four geographic populations was very small under the natural photoperiod in Osaka at 20°C, and absent under the natural photoperiod and temperature in Osaka. These results suggest that A. verbasci survives and successfully produces the next generation in different geographic regions without changing the parameters of the circannual rhythm.

Immune responses to lipopolysaccharide challenge in a tropical rodent (Funambulus pennanti): Photoperiod entrainment and sex differences

Annual variation in day length (photoperiod) triggers changes in the immune system of seasonal breeders. The rationale behind this study was to delineate any sex differences in immune responses of photoperiodically entrained animals challenged against lipopolysaccharide (LPS)-induced inflammatory stress. We observed that photoperiodically entrained [short day, SD, 10 h light (L):14 h dark (D); long day, LD, 16 h L:8 h D; and natural day length, NDL, 12 h L:12 h D] male and female Indian palm squirrels, Funambulus pennanti, presented sexual dimorphism in immune status after LPS-induced stress. Females presented high humoral (anti-keyhole limpet hemocyanin immunoglobulin) and cellular immunity (lymphocyte proliferation) compared with the males of all photoperiodic conditions. Female squirrels showed reduced pro-inflammatory cytokine levels (interleukin-1β, interleukin-6, and tumor necrosis factor-α) than the males suggesting their high efficiency to recover from LPS-induced inflammatory stress. Increased duration of melatonin secretion and corticosterone concentration in squirrels experiencing SD evidently supported survival of squirrels as compared with control (NDL) and LD squirrels of both sexes. Decreased immune status in both sexes under LD condition might be due to a short melatonin signal mimicking the LDs of summer. Thus, we infer that photoperiodic entrainment via the levels of melatonin and corticosterone synergistically supported more the survival of female squirrels under LPS-induced stress.

A circadian system is involved in photoperiodic entrainment of the circannual rhythm of Anthrenus verbasci

In the circannual pupation rhythm of the varied carpet beetle, Anthrenus verbasci, entrainment to annual cycles is achieved by phase resetting of the circannual oscillator in response to photoperiodic changes. In order to examine whether a circadian system is involved in expression of the periodic pattern and phase resetting of the circannual rhythm as photoperiodic responses, we exposed larvae to light–dark cycles with a short photophase followed by a variable scotophase (the Nanda–Hamner protocol). When the cycle length ( T) was a multiple of 24 h, i.e., 24, 48, or 72 h, short-day effects were clearer than when T was far from a multiple of 24 h, i.e., 36 or 60 h. Exposure to light–dark cycles of T = 36 h had effects similar to exposure to long-day cycles of T = 24 h. The magnitude of phase shifts depended on the duration and the phase of exposure to the cycles of T = 36 or 60 h. It was therefore concluded that a circadian system is involved in photoperiodic time measurement for phase resetting of the circannual oscillator of A. verbasci.

The Rat Circadian Clockwork and its Photoperiodic Entrainment During Development

The mammalian circadian pacemaker is located in the suprachiasmatic nucleus (SCN), which is composed of dorsomedial (dm) and ventrolateral (vl) regions. The molecular clockwork responsible for the SCN rhythmicity consists of clock genes and their transcriptional‐translational feedback loops. The rat SCN rhythmicity and clockwork are affected by the photoperiod. The aim of this study was to elucidate development of the rat SCN rhythmicity, namely of the rhythmicity of the dm‐ and vl‐SCN and of expression of clock genes and to ascertain when the photoperiod starts to affect the SCN rhythmicity. Rhythmicity of the dm‐SCN, measured as the rhythm in spontaneous c‐FOS production, developed earlier than that of the vl‐SCN, which was measured as the rhythm in c‐FOS photoinduction. However, photoperiodic affection of the rhythmicity occurred earlier in the vl‐SCN than in the dm‐SCN. From the 4 clock genes (Per1, Per2, Cry1 and Bmal1) studied, the expression of Bmal1 and Per1 was rhythmic already in 1‐day‐old rats; at this age, the Per2 mRNA rhythm just started to form and no rhythm in Cry1 expression was detected. After the second postnatal day, all 4 genes were expressed in a rhythmic manner. Thereafter, the rhythms matured gradually via increasing amplitude. Per1 and Per2 mRNA rhythms started to be affected by the photoperiod at the 10th postnatal day. The data suggest that the rhythms in clock genes expression in the rat SCN develop mostly postnatally. The molecular clockwork may start to be photoperiod‐dependent around the 10th postnatal day.

Ontogenesis of photoperiodic entrainment of the molecular core clockwork in the rat suprachiasmatic nucleus

The molecular mechanism underlying a generation of circadian rhythmicity within the suprachiasmatic nucleus (SCN) is based on interactive negative and positive feedback loops that drive the rhythmic transcription of clock genes and translation of their protein products. In adults, the molecular mechanism is affected by seasonal changes in day length, i.e., photoperiod. The photoperiod modulates phase, waveform, and amplitude of the rhythmic clock genes expression as well as the phase relationship between their profiles. To ascertain when and how the photoperiod affects the circadian core clock mechanism during ontogenesis, the rhythmic expression of clock genes, namely of Per1, Per2, Cry1 and Bmal1 was determined in 3-, 10- and 20-day-old rat pups maintained under either a long photoperiod with 16 h of light and 8 h of darkness per day (LD 16:8) or under a short, LD 8:16 photoperiod. The daily profiles in the level of clock genes mRNA were studied in constant darkness. The photoperiod affected the profile of Per1 and Per2 mRNA in 20- and 10- but not yet in 3-day-old pups. Expression of Cry1 was affected only in 20-day-old pups, whereas expression of Bmal1 was not yet affected even in 20-day-old rats. The results demonstrate no effect of the photoperiod on 3-day-old pups, only partial entrainment of the molecular core clockwork in 10-day-old pups and a more mature, though not yet fully complete, entrainment in 20-day-old pups as compared with adult animals. The developmental interval when the photoperiod begins to entrain the core clock mechanism completely might thus occur around the time of weaning.

Photoperiod regulates clock gene rhythms in the ovine liver

To investigate the photoperiodic entrainment of peripheral rhythms in ruminants, we studied the expression of clock genes in the liver in the highly seasonal Soay sheep. Animals were kept under long (LD 16:8) or short photoperiod (LD 8:16). Daily rhythms in locomotor activity were recorded, and blood concentrations of melatonin and cortisol were measured by RIA. Per2, Bmal1, and Cry1 gene expression was determined by Northern blot analyses using ovine RNA probes in liver collected every 4 h for 24 h. Liver Per2 and Bmal1, but not Cry1, expression was rhythmic in all treatments. Under long days, peak Per2 expression occurred at end of the night with a similar timing to Bmal1, whereas, under short days the Per2 maximum was in the early night with an inverse pattern to Bmal1. There was a photoperiod × time interaction for only Per2 ( P < 0.001). The 24-h pattern in plasma cortisol matched the observed phasing of Per2 expression, suggesting that it may act as an endocrine entraining factor. The clock gene rhythms in the peripheral tissues were different in timing compared with the ovine suprachiasmatic nucleus (SCN, central pacemaker) and pars tuberalis (melatonin target tissue), and the hepatic rhythms were of lower amplitude compared with photoperiodic rodents. Thus, there are likely to be important species differences in the way the central and peripheral clockwork encodes external photoperiod.

Circadian rhythm entrainment with melatonin, melatonin receptor antagonist S22153 or their combination in mice exposed to constant light.

The ability of daily melatonin and the melatonin receptor antagonist, S22153, to entrain circadian system function was investigated in mice with atypical melatonin rhythm. B6D2F(1) mice were first synchronized to a LD 12:12 for approximately 2 wk, then exposed to continuous light (LL) until study completion. After 10-18 days of LL exposure, mice received daily subcutaneous (s.c.) melatonin at a dose of 0.1, 1 or 10 mg/kg/day (exp. 1) or daily intraperitoneal (i.p.) S22153 (20 mg/kg/day) with or without melatonin (1 mg/kg/day, exp. 2) at subjective zeitgeber time (ZT) 10 for 19 days. Then all the mice were exposed to LL for another 10 days. Spectral analysis showed that initial LL lengthened the period of both rhythms by approximately 1.5 hr as compared with LD 12:12. No entrainment of either rhythm was found in controls. Conversely, daily melatonin-only, S22153-only or their combination set the temperature and activity periods to approximately 24 hr and produced a significant increase of the circadian amplitude of both rhythms as compared with controls. However, after treatment withdrawal, the dominant period lengthened to approximately 25.5 hr in mice receiving either melatonin or S22153. On the contrary, the period remained close to 24 hr for the 10 days following withdrawal of combined S22153 and melatonin. Such sustained pharmacological resetting of circadian function could display therapeutic potential against external resynchronization resulting from defective photoperiodic entrainment.

Wheel-Running Activity Patterns of Five Species of Desert Rodents

In contrast to the extensive laboratory data on activity patterns in rodent species inhabiting temperate zones, much less is known about the activity patterns of desert rodents. In order to address this issue, we measured wheel-running activity patterns in males and females of five species of wild-trapped desert rodents (Dipodillus dasyurus, Gerbillus andersoni, Gerbillus pyramidum, Meriones shawi, and Acomys cahirinus) in long ‘summer-like’ and short, ‘winter-like’ day lengths. The specific goals of the present study were to characterize activity patterns in several desert rodent species in the laboratory and to determine if activity patterns are expressed in a seasonal or sexually dimorphic manner. Specifically, wheel-running was measured for 11 weeks in long days followed by 11 weeks in short days to test for photoperiodic entrainment as well as responsiveness to changes in the light-dark cycle. All animals exhibited rhythmic patterns of wheel-running with consistent onsets and offsets that had well-defined relations with the light-dark cycle. All individuals of G. andersoni showed nocturnal activity patterns. Most individuals of G. pyramidum had nocturnal activity patterns, but some individuals showed a short bout of activity at the beginning of the light period. Most individuals of D. dasyurus and M. shawi showed bimodal (i.e., nocturnal and diurnal) activity patterns, although some showed markedly nocturnal activity patterns. There was no sexual dimorphism in wheel running activity rhythms in any of the species examined. As expected, decreases in day length resulted in an overall increase in the duration of activity in all species. Collectively, these data provide an initial characterization of activity patterns within desert rodents in a controlled laboratory setting.

Photoperiodic entrainment of seasonal changes in the appetite, feeding behaviour, growth rate and pelage of pony colts

AbstractRelationships among photoperiod and changes in voluntary food intake, feeding behaviour, growth and pelage were determined in seven, 2-year-old pony colts (182·4 (s.e. 5·4) kg). Individually housed colts were provided with ad libitum access to a complete pelleted diet (gross energy = 16·7 MJ/kg dry matter). Voluntary food intake (VFI, kg/ day) was calculated daily and body weights were recorded weekly throughout the 36-week study. Feeding behaviour was evaluated at approximately 4-week intervals by continuous observation (24 h), and the hair weight density (HWD, mg/cm2) of shoulder pelage was determined fortnightly. Day length was artificially manipulated to mimic the prevailing mid-summer photoperiod (16 h light: 8 h dark, 16L: 8D). After 1 week of the study (and the preceding fortnight), day length was abruptly decreased and thereafter animals were exposed to alternating 14-week periods of short (SD, 8L: 16D) and long days (LD, 16L: 8D). The mean daily VFI of individual ponies was calculated weekly and normalized for digestible energy (DE) content and metabolic body weight (DEI, MJ/kg M0·75). The average daily gain (ADG, kg/day) in body weight of each individual was calculated weekly. The apparent digestibility of dietary energy (digestibility) was determined over 72 h (no. = 6) on two occasions (days 92 to 95 and 190 to 193) during the study. Digestibility was similar in both periods (0·48, s.e. 0·01). DEI, ADG and HWD changed in a cyclic manner throughout the study. The period of the appetite cycle (24·4 (s.e. 1·3) weeks) did not differ from that of the 28-week photoperiodic regime. DEI decreased from a maximum of 1·4 (s.e. 0·03) MJ/kg M0.75 per day (day 21), to a nadir of 0·75 (s.e. 0·02) MJ/kg M0.75 per day (day 154, P &lt; 0·001) and had increased (P &lt; 0·001) to attain a second zenith (0·93 (s.e. 0·01) MJ/kg M0.75 per day) before the end of the study. Ponies ate discrete meals of similar duration, but meal frequency was associated with changes in VFI (r = 0·77) as was proportion of time spent feeding (r = 0·79). Changes in ADG reflected those of DEI. Body weight was stable for 4 weeks at the nadir of the appetite cycle. Maximal HWD was coincident with the nadir of the appetite and growth cycles. Regression of individual values for DEI on ADG described a linear relationship (R2 = 0·80) which could be used to predict the energy requirements of growing ponies maintained under similar conditions:DEI total (MJ/kg M0·75 per day) = 0·654ADG (kg/day) + 0·789 (1).The duration of the photoperiod, appetite, growth and pelage cycles were similar, suggesting a causal relationship. Physiological responses to photoperiodic change were not immediate and exhibited a delay of 5 to 8 weeks.

Photoperiodic history determines the reproductive response of rainbow trout to changes in daylength

The mechanisms underlying the photoperiodic entrainment of the endogenous circannual rhythm of maturation in the rainbow trout were investigated by subjecting December-spawning fish to abrupt changes in daylength which varied in their timing or magnitude. These protocols advanced spawning by up to 4 months. Maturation occurred in sequence in fish maintained on 18L:6D from January and February, and in fish exposed to 18L:6D from December, January and February, followed by 6L:18D in May, indicating that the abrupt increases in daylength were effective entraining cues. `Long' photoperiods of between 12 and 22 h applied in January, followed by shorter photoperiods of between 3.5 and 13.5 h from May, were equally effective in advancing maturation. Maturation was also advanced, though to a lesser extent, in fish maintained on photoperiods of 8.5 or 10 h from January, followed by a photoperiod of 1.5 h from May. In contrast, maturation was delayed in fish maintained under a constant 8.5-h photoperiod from January, and these fish also exhibited a desynchronization of spawning times characteristic of endogenous circannual rhythms in free-run. Collectively, these results indicate that photoperiodic history determines the reproductive response of rainbow trout to changes in daylength.