Annual changes in body mass in captive Svalbard ptarmigan: role of changes in locomotor activity and food intake

The influence of changes in the amount of locomotor activity on the annual body mass cycle was investigated in captive Svalbard ptarmigan kept indoors at thermoneutrality and exposed to seasonal changes in daylength or continuous light from the summer onwards. In both groups there was a close correlation between locomotor activity and metabolic rate. Only birds exposed to changes in daylength showed an annual cycle in locomotor activity, with low activity in autumn and mid-winter and a peak in spring. The birds permanently exposed to continuous light had a relatively low activity throughout the year with no systematical changes. Body mass began to increase in both groups in early autumn and the food intake was elevated during most of the following fattening period. It is concluded that elevated food intake is the prime factor involved in autumnal fattening in captive Svalbard ptarmigan. Body mass increased significantly faster under decreasing daylength compared with continuous light, associated with a lower activity as well as a higher food intake. The birds exposed to continuous light maintained a high body mass and a relatively low activity level during spring. In birds exposed to changing daylength, body mass fell from late February onwards, which is about 3 months later than in outdoor caged or free-living Svalbard ptarmigan. In the birds exposed to increasing daylength a fourfold increase in the amount of locomotor activity occurred from February to April. This increased activity was correlated with a negative energy balance and may be casually associated with the fall in body mass in these birds. Under outdoor conditions, elevated locomotor activity in spring may be responsible for a continuation of the decline in body mass commencing in November, despite a slight tendency for an increased food intake towards the end of this period.Abbreviations BM body mass - CE _f caloric equivalent of food - EAE energy assimilation efficiency - EE energy expenditure - FI food intake - LA locomotor activity - LL continuous light - LD simulated annual changes in daylength - MEI metabolizable energy intake - MR metabolic rate - RQ respiratory quotient     

Protein utilization during starvation in fat and lean Svalbard ptarmigan (Lagopus mutus hyperboreus)

Summary Body protein sparing during starvation has been examined in fat and lean Svalbard ptarmigan. Protein utilization was determined from daily N excretion and from the rate of decrease in body mass. Changes in plasma concentrations of -hydroxybutyrate, free fatty acids, glucose, and uric acid were also recorded. When fat birds were starved for 15 days protein catabolism initially fell (phase I) and was thereafter kept low (phase II). This was evident from the temporal pattern in both N excretion and body mass loss. In two birds, N excretion eventually increased, revealing enhanced protein catabolism and thus a third phase of starvation. Changes in protein utilization were paralleled by changes in plasma uric acid. Approximately 9% of the energy demand was covered by breakdown of body protein during phase II. The importance of fat catabolism in providing energy was indicated by markedly elevated plasma levels of -hydroxybutyrate and free fatty acids. When lean birds were starved for 5 days there appeared to be no phase II. The temporal pattern of body mass loss indicated phase I and III but that of N excretion only phase III. The relative contribution of body protein to energy demand increased from 22% at day 2 to 41% at the end of starvation and was paralleled by increased plasma uric acid. When data from lean and fat birds were pooled, the changes in uric acid and N excretion were highly correlated (r=0.92, P<0.001), indicating that plasma uric acid is a reliable index of protein breakdown in starving Svalbard ptarmigan. In conclusion, starving fat Svalbard ptarmigan have a much greater capacity to spare body protein than lean birds. Fat birds effectively reduce protein catabolism and maintain this at a low level whereas starving lean birds increase protein catabolism.Abbreviations -OHB -hydroxybutyrate - BM body mass - BMR basal metabolic rate; dne daily nitrogen excretion - FFA free fatty acids - MR metabolic rate     

Seasonal and daily variations in plasma melatonin in the high-arctic Svalbard ptarmigan (Lagopus mutus hyperboreus).

This study presents the daily rhythm of melatonin secretion throughout one year in a bird from the northern hemisphere, the Svalbard ptarmigan (Lagopus mutus hyperboreus), which lives naturally at 76-80 degrees N. Eight Svalbard ptarmigan were caged outdoors at 70 degrees N and blood sampled throughout one day each month for 13 months. At this latitude, daylight is continuous between May and August, but there is a short period of civil twilight around noon from late November to mid January. There was no daily rhythm in plasma melatonin in May-July. Plasma melatonin levels varied significantly throughout the day in all other months of the year, with the nighttime increase reflecting the duration of darkness. The highest mean plasma concentration occurred at midnight in March (110.1 +/- 16.5 pg/ml) and represented the annual peak in estimated daily production. Around the winter solstice, melatonin levels were significantly reduced at noon and elevated during the nearly 18 h of consecutive darkness, and the estimated mean daily production of melatonin was significantly reduced. Thus, at the times of the year characterized by light-dark cycles, melatonin may convey information concerning the length of the day and, therefore, progression of season. The nearly undetectable low melatonin secretion in summer and the reduced amplitude and production in midwinter indicate a flexible circadian system that may reflect an important adaptation to life in the Arctic.     

Energetic consequences of a major change in habitat use: endangered Brent Geese Branta bernicla hrota losing their main food resource

Coastal seagrasses are declining at increasing rates worldwide, forcing herbivores previously reliant on these habitats to abandon them in search of alternative ways to fulfil their daily energy budgets. After two decades of declining seagrass abundance in Mariager Fjord, Denmark, the Svalbard breeding population of Light‐bellied Brent Geese Branta bernicla hrota has experienced substantial changes in habitat use at this traditional autumn staging area. Declines in seagrasses have caused birds to depend increasingly on Sea Lettuce Ulva lactuca in recent years, and forced birds into terrestrial habitats such as saltmarsh and winter wheat. In contrast to those birds exploiting aquatic habitats, birds relying on these new habitats showed higher energy expenditure and failed to balance their energy budget. Eelgrass (Zostera) was energetically superior to other food resources, with marine Ulva being second best. Predicted body mass development under two different scenarios indicate that present habitat use resulted in a midwinter body mass around 122 g lower than just 20 years ago, equivalent to c. 9.4% of Brent Goose body weight. Even after controlling for inter‐annual differences in thermoregulatory costs, the effect of changes in habitat use translated into a body mass reduction of c. 56 g, which could adversely affect survival and future reproduction. Flyway‐wide declines in Zostera abundance and further reductions in traditional habitats due to climate change give cause to reassess projected population trends and consequent management implications for the East Atlantic flyway population of Light‐bellied Brent Geese.     

Demographic consequences of age-structure in extreme environments: population models for arctic and alpine ptarmigan

Organisms living in arctic and alpine environments are increasingly impacted by human activities. To evaluate the potential impacts of global change, a better understanding of the demography of organisms in extreme environments is needed. In this study, we compare the age-specific demography of willow ptarmigan (Lagopus lagopus) breeding at arctic and subalpine sites, and white-tailed ptarmigan (L. leucurus) breeding at an alpine site. Rates of egg production improved with age at the alpine and subalpine sites, but the stochastic effects of nest and brood predation led to similar rates of annual fecundity among 1-, 2-, and 3+-year-old females. All populations had short generation times (T<2.7 years) and low net reproductive rates (R ₀<1.2). Stable age distributions were weighted towards 1-year-old females in willow ptarmigan (>59%), and to 3+-year-old females in white-tailed ptarmigan (>47%). High damping ratios (ρ>3.2) indicated that asymptotic estimates were likely to match natural age distributions. Sensitivity and elasticity values indicated that changes in juvenile survival would have the greatest impact on the finite rate of population change (λ) in willow ptarmigan, whereas changes to the survival of 3+-year-old females would have a greater effect in white-tailed ptarmigan. High survivorship buffers white-tailed ptarmigan in alpine environments against the potential effects of climate change on annual fecundity, but may make the species more sensitive to the effects of pollutants or harvesting on adult survival. Conversely, processes that reduce annual fecundity would have a greater impact on the population viability of willow ptarmigan in arctic and subalpine environments. If these same demographic patterns prove to be widespread among organisms in extreme environments, it may be possible to develop general recommendations for conservation of the biological resources of arctic and alpine ecosystems.     

Huddling house sparrows remain euthermic at night,and conserve body mass

An advantage of huddling in the cold is that the individual animals involved can maintain body temperature while saving energy. Since house sparrows Passer domesticus biblicus store little fat, but inhabit relatively cold climates, we tested the hypothesis that they huddle at night. While recording body temperature and body mass of 18 house sparrows when they were either caged individually, or free in an aviary, we observed that when free in the aviary, the sparrows huddled at low ambient temperatures and more birds huddled, in tighter and tighter formation, as ambient temperature decreased. However, their body temperatures were not significantly different from when they spent the night individually caged. When free to huddle, the birds lost significantly less body mass during the course of a day than when individually caged. This reduction in body mass loss may be of particular importance during periods of adverse environmental conditions, especially for small birds that manage their energy budgets on a daily basis.     

Reduced metabolic cost of locomotion in Svalbard rock ptarmigan (Lagopus muta hyperborea) during winter

The Svalbard rock ptarmigan, Lagopus muta hyperborea experiences extreme photoperiodic and climatic conditions on the Arctic archipelago of Svalbard. This species, however, is highly adapted to live in this harsh environment. One of the most striking adaptations found in these birds is the deposition, prior to onset of winter, of fat stores which may comprise up to 32% of body mass and are located primarily around the sternum and abdominal region. This fat, while crucial to the birds' survival, also presents a challenge in that the bird must maintain normal physiological function with this additional mass. In particular these stores are likely to constrain the respiratory system, as the sternum and pelvic region must be moved during ventilation and carrying this extra load may also impact upon the energetic cost of locomotion. Here we demonstrate that winter birds have a reduced cost of locomotion when compared to summer birds. A remarkable finding given that during winter these birds have almost twice the body mass of those in summer. These results suggest that Svalbard ptarmigan are able to carry the additional winter fat without incurring any energetic cost. As energy conservation is paramount to these birds, minimising the costs of moving around when resources are limited would appear to be a key adaptation crucial for their survival in the barren Arctic environment.     

Foraging strategies of Great Cormorants Phalacrocorax carbo carbo wintering north of the Arctic Circle

This study describes how 30 Great Cormorants Phalacrocorax carbo carbo managed to catch sufficient food for their daily energetic needs under conditions of reduced daylight and cold while wintering north of the Arctic Circle. Activity observations showed that the Great Cormorants' daily foraging pattern was generally bimodal, with morning and evening feeding peaks. They compensated for shorter daylengths in midwinter by starting to forage later and ending progressively earlier at lower light intensities. Fishing constituted only a minor part of their time–activity budget, and was one of the most efficient reported in marine birds. The Great Cormorants spent less than 60 minutes a day fishing in midwinter. Although subzero ambient temperatures and blizzards contributed to increased heat loss in midwinter, this potential energy loss did not seem to be compensated for by an increase in fish intake. Instead the Great Cormorants seemed to economize energy expenditure by halving the time spent at sea, and halving the number but doubling the mass of each fish taken.     

Seasonal changes in daily metabolic patterns of Lacerta viridis

Summary Lacerta viridis maintained under natural photoperiodic conditions show daily and seasonal changes in metabolic rates and body temperature (T _b) as well as seasonal differences in sensitivity to temperature change. At all times of the year lizards have a daily fluctuation in oxygen consumption, with higher metabolic rates during the light phase of the day when tested at a constant ambient temperature (T _a) of 30°C. Rhythmicity of metabolic rate persists under constant darkness, but there is a decrease in the amplitude of the rhythm.Oxygen consumption measured at various T_as shows significant seasonal differences at T _as above 20°C. Expressed as the Arrhenius activation energy, metabolic sensitivity of Lacerta viridis shows temperature dependence in autumn, which changes to metabolic temperature independence in spring at T _as above 20°C. The results indicate a synergic relationship between changing photoperiod and body temperature selection, resulting in seasonal metabolic adjustment and seasonal adaptation.Abbreviations ANOVA analysis of variance - LD long day (16 h light) - SD short day (8 h light) - T _a ambient temperature - T _b body temperature     

New findings in body mass regulation in zebra finches (Taeniopygia guttata) in response to photoperiod and temperature

Small birds in temperate zones increase body mass in winter (Lehikoinen, 1987). We investigated daily and annual variation of body mass and fat reserves of locally-reared Australian zebra finches Taeniopygia guttata , by exposing them to different photoperiods, feeding periods, and temperatures.
Experiments with long and short photoperiods and long and short feeding periods, showed that long photoperiods increased body mass and fat reserves of the zebra finches, and readiness to breed, independently of the actual feeding period.
Furthermore, the zebra finches in indoor aviaries with constant temperature (22-24°C) and in outdoor aviaries with ambient temperature, both exposed to the natural daylength changes of Bielefeld, Germany (52 °N), had high dawn body mass in summer (12.9 and 12.0-12.4g, respectively) and low in winter (10.7 and 11.1 g, respectively). Thirty to sixty percent of these mass changes were related to changes in fat reserves, so that the finches had only 0.1-0.2g of metabolizable fat reserves in short photoperiods (or in winter), which increased up to 1.5g in long photoperiods (or in summer).
Indoor finches consumed more seeds in summer than in winter (3.3 vs. 2.7g/day), while outdoor finches consumed 4-5g of seeds per day throughout the year, which probably represented the limit of energy intake for a 11-13g bird (Kirkwood, 1983). Nightly mass loss, increasing from 0.7g in summer up to 2.0g in winter, was highly positively correlated with night length, not influenced by ambient temperature. Foraging before dawn and after dusk, roosting with well-filled crops, and decreasing body mass and fat reserves, seem to be adaptations of zebra finches for survival in winter. The summer fattening probably accelerates reproduction in this opportunistic breeder, by allocating more time to reproductive behaviour and more endogenous nutrients to egg-formation.     

Monitoring Svalbard rock ptarmigan: Distance sampling and occupancy modeling

The only resident terrestrial herbivorous bird species in high-Arctic Svalbard, Norway is the endemic Svalbard rock ptarmigan (Lagopus muta hyperborea) of which little is known of its population dynamics. We assessed temporal and spatial variability of the pre-breeding population of Svalbard rock ptarmigan males using: 1) distance sampling to estimate density (2000–2009) and 2) occupancy modeling to determine the proportion of survey points being occupied in relation to a habitat index for ptarmigan habitat suitability (2005–2009). Data were collected using a point-transect sampling design. We split the analysis according to type of survey point (non-random, random, and survey points combined). Our estimated spring densities were low (1.3–3.1 territorial male/km², non-random survey points, 2000–2009) with limited annual variability. The best models describing occupancy rates of territorial males at 2 different spatial scales (ptarmigan males observed ≤250 m and ≤450 m from the sampling point) were independent of spatial scales and the type of survey points. Occupancy dynamics were related to the habitat index whereas detection probability was year dependent. Extinction probability was negatively related to habitat quality (good habitats had lower extinction probability). We could not estimate the habitat effect on colonization precisely because initial occupancy rates were high at both spatial scales (estimated average initial occupancy at scale ≤250 m = 0.96; scale ≤450 m = 0.97). Colonization appeared to be positively related to the habitat index for the random survey points (including mainly marginal habitats), but the small sample size led to large uncertainty in the parameter estimate. Detection probabilities varied greatly between study years, thus demonstrating the importance of estimating detection probability annually. We recommend that future surveys are stratified with respect to habitat quality and to integrate the 2 methodologies in population monitoring of Svalbard rock ptarmigan. © 2011 The Wildlife Society.     

Intermittent feeding in a migratory omnivore: digestion and body composition of American black duck during autumn

Birds fast intermittently during weather disturbances and migration. We tested responses of black duck to lost feeding days during autumn mass gain. Nine adult males were fed a pelleted diet (1.5% fat, 15.8% protein, and 18.3% neutral detergent fiber) and caged indoors during September and October (12 h light; 17 degrees -24 degrees C) to measure balances over 14 d when fed ad lib. each day and fasted intermittently for 2 d wk(-1) (short fast) or 4 d wk(-1) (long fast). Body mass (1,081 g), body water content, and metabolizable intakes of energy and protein were maintained as daily intakes of dry matter increased to 1.65 (short fast) and 2.35 (long fast) times the unfasted level. Intermittent feeding reduced metabolizability of dry matter, energy, protein, and acid detergent fiber. Concentrations of Mn provided similar estimates of metabolizability to direct measures in unfasted birds but underestimated measures of birds on long fasts. Fasting regimes continued outdoors for 9 wk when temperatures declined to -9 degrees C. Birds on short fasts were heavier (1,373 vs. 1,241 g) and fatter (159 vs. 58 g) than those on long fasts, while body water (894 g) and protein (316 g) were similar between groups after 5 wk. Birds on long fasts subsequently gained mass when fed daily, but those on short fasts lost mass when fed each day. Omnivorous waterfowl combine ingestive and digestive flexibility with plasticity of body lipid to contend with uncertain food availability.     

Interactive effects of temperature and photoperiod on the daily activity and energy metabolism of pouched mice (Saccostomus campestris: Cricetidae) from southern Africa

The daily activity and energy metabolism of pouched mice (Saccostomus campestris) from two localities in southern Africa was examined following warm (25 °C) and cold (10 °C) acclimation under long (LD 14:10) and short (LD 10:14) photoperiol. There was no differential effect of photoperiod on the daily activity or metabolism of pouched mice from the two localities examined, which suggests that reported differences in photoresponsivity between these two populations were not the result of differences in daily organisation. Neverthe-less, there was a significant increase in metabolism at 10 °C, irrespective of photoperiod, even though seven cold-acclimated animals displayed bouts of spontaneous torpor and saved 16.4–36.2% of their daily energy expenditure. All but one of these bouts occurred under short photoperiod, which suggests that short photoperiod facilitated the expression of torpor and influenced the daily energy metabolism of these individuals. As expected for a noctureal species, the amount of time spent active increased following acclimation to short photoperiod at 25 °C. However, there was a reduction in mean activity levels under short photoperiod at 10 °C, possibly because the stimulation of activity by short photoperiod was masked by a reduction in activity during bouts of spontaneous torpor. Cold temperature clearly had an overriding effect on the daily activity and metabolism of this species by necessitating an increase in metabolic heat production and eliciting spontaneous torpor which overrode the effect of short photoperiod on activity at an ambient temperature of 10 °C.Abbreviations 3-ANOVA three-way analysis of variance - %ACT percentage of time spent active - ADMR average daily metabolic rate - M _b body mass - MR metabolic rate - MR_dark metabolic rate recorded during the dark phase - MR_light metabolic rate recorded during the light phase - NST non-shivering thermogenesis - RQ respiratory quotient - STPD standard temperature and pressure, dry - T _a ambient temperature - T _b body temperature - VO₂ oxygen consumption     

Photosynthetic and growth responses of three freshwater algae to phosphorus limitation and daylength

1. Three common species of freshwater phytoplankton, the diatom Nitzschia sp., green alga Sphaerocystis schroeteri and cyanobacterium Phormidium luridum, were grown under contrasting daylengths [18 : 6 h light : dark cycles (LD) versus 6 : 18 h LD] and phosphorus (P) regimes (P‐sufficient versus 1 μm P). The rates of growth and photosynthesis, as well as growth efficiencies and pigment concentrations, were compared among treatments. 2. The growth and photosynthetic parameters of the three species depended on both P status and daylength in a species‐specific way. The responses to P limitation depended on daylength and, conversely, the responses to daylength depended on P status. 3. Growth rates and the maximum rates of photosynthesis (P_max) of all species decreased under P limitation under both light regimes. However, the decrease of P_max because of P limitation was greater under long daylength. The P_max of the green alga S. schroeteri decreased the most (ca. sixfold) under P limitation compared with the other two species. The photosynthesis saturation parameter I_k also decreased under P limitation; the decline was significant in Nitzschia and Sphaerocystis. P‐limitation significantly increased photoinhibition (β) in Nitzschia and Sphaerocystis, but not in Phormidium. The excess photochemical capacity (the ratio of the maximum photosynthesis rate to the photosynthesis rate at the growth irradiance), characterising the ability to utilise fluctuating light, was significantly lower under P limitation. 4. The growth efficiency (growth rate normalised to daylength) declined with increasing daylength in all species. Under short daylength the cyanobacterium Phormidium had the lowest growth efficiency of the three species. 5. The cellular chlorophyll a concentration in both Nitzschia and Sphaerocystis was significantly higher under short daylength, but only under P‐sufficient conditions. In Nitzschia, under short daylength, P‐limitation significantly decreased cellular chlorophyll concentration. In contrast, P‐limitation increased cellular chlorophyll concentration in Sphaerocystis, but under long daylength only. The ratio of chlorophyll a to b in the green alga also declined under short daylength and under P‐limited conditions.     

Influence of daylength on metabolic rate and daily water loss in the male prosimian primate Microcebus murinus

In its natural habitat, Microcebus murinus, a small malagasy prosimian primate, is exposed to seasonal shortage of water and resources. During the winter dry season, animals enter a pronounced fattening period with concurrent decrease in behavioural/physiological activities, whereas the breeding season is restricted to the rainy summer months. To determine the role of daylength on metabolic rate and water loss in this nocturnal primate, we measured body mass, oxygen consumption at 25°C (RMR), circadian water loss through urine output (UO) and evaporation (EWL) in eight males exposed to either short days (8L:16D SD) or long days (14L:10D LD), under controlled captive conditions. Exposure to SD led to a ponderal increase (maximal body mass: 125±4 g, N=8), and to significant changes in RMR and water loss, both reaching lowest values after 3 months under SD (0.84±0.04 ml O₂ h⁻¹ g⁻¹ and 38±0.3 mg H₂O g⁻¹ day⁻¹, respectively). Following exposure to LD, body mass decreased to 77±3 g (N=8), whereas both RMR and water loss, mainly through EWL, significantly increased (P<0.001), the highest value occurring after 2 months (1.51±0.08 ml O₂ h^.−1 g⁻¹ and 87±7 mgH₂O g⁻¹ day⁻¹, respectively). Moreover, independent of daylength, circadian changes in EWL were characterized by significantly reduced values during the diurnal rest. The results demonstrate that daylength variations affect the physiology of this tropical primate, allowing anticipatory adaptation to seasonal environmental constraints.     

Even a mild spell of cold weather affects feeding in migrating redheaded buntings

We investigated whether a climatic change in temperature affected daily food intake in migrating male redheaded buntings. Groups of adult male birds (n = 18) were photoinduced into migratory phenotype under increasing spring daylengths (NDL); as the birds began to exhibit night restlessness, Zugunruhe, these were allocated into groups, either with ambient (NDL, variable daily temperature: maximum – 29–44 °C and minimum – 16–33 °C; for food intake (six birds) and activity recording, six birds) until 2 weeks after they concluded migration or with constant temperature (NDT, 22 ± 1 °C; for food intake (six birds)) conditions. As day length increased March onwards, daily food intake increased (hyperphagia) in NDL and NDT groups. However, hyperphagia was slower in NDT birds as compared to NDL birds, suggesting that altered ambient temperature affects daily food intake in migrating buntings. Another group of 12 birds were held under constant daylengths (12L:12D; EDT and constant temperature 22 ± 1 °C). Although the onset of Zugunruhe was delayed under EDT, the day of onset of Zugunruhe was taken as day 0. Daily food intake and body weight before and during migration of EDT birds were compared with that of NDT and NDL groups. Daily food intake and body weight increased in all migrating birds, but hyperphagia continued post-migration in NDT birds. The study suggests that constantly suboptimal temperature despite increasing daylength, NDT, appeared to affect feeding and body weight of migratory buntings as evident from continued hyperphagia and body weight gain, even after concluding migrating activity.     

Does avian winter fat storage integrate temperature and resource conditions? A long‐term study

Theoretical models of short‐term avian behaviour suggest that small birds adaptively balance the ecological costs and benefits of winter fat to maximize survival probability. When low starvation risk eliminates benefit but not cost of fat, birds are leaner than when under high starvation risk. Most models focus on single factors affecting starvation risk and subsequent choice of adaptive body mass; however, in complex environments, more than one factor affects starvation risk. To test for multiple interacting factors affecting fat reserves, long‐term geographical data on winter fat in a ground‐feeding finch, the dark‐eyed junco Junco hyemalis were analyzed. Two measures of fat were used: (1) visible subcutaneous fat class, and (2) body mass residuals left after age, sex and wing length effects were factored out. Site means for fat measures were obtained from juncos visiting supplemental feeding sites in midwest and northwest North America. In backward elimination regression of fat class, the temperature‐snowfall interaction term and its constituent variables, proximate temperature (averaged over capture day and the preceding ten days) and snowfall (frequency over the same time interval) were significant explanators of variation. Snowfall frequency is considered to be a surrogate measure of resource deterioration. The interaction term, also found in backward regression of body mass residuals, showed that as temperature declined at low snowfall frequency, less fat was deposited than when temperature declined at high snowfall frequency. Thus, in a recently cold environment suggesting relatively high resource predictability, perceived starvation risk is low, and less costly fat is needed to reduce starvation risk compared to a cold and unpredictable resource environment. The analysis of mass residuals also yielded a significant effect of daylength, suggesting an underlying fattening programme independent of proximate environmental conditions. A longitudinal study of junco fat stores indicated that individual environmental responses contributed significantly to midwinter fat peaks. These results agree with predictions of a synergistic model of adaptive fat regulation in small birds by suggesting that a ground‐feeding bird may maximize winter survival probability by integrating multiple environmental factors affecting starvation risk.     

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)