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The Qinghai-Tibetan Plateau, with low precipitation, low oxygen partial pressure, and temperatures routinely dropping below −30 °C in winter, presents several physiological challenges to its fauna. Yet it is home to many endemic mammalian species, including the plateau pika (Ochotona curzoniae). How these small animals that are incapable of hibernation survive the winter is an enigma. Measurements of daily energy expenditure (DEE) using the doubly labeled water method show that pikas suppress their DEE during winter. At the same body weight, pikas in winter expend 29.7% less than in summer, despite ambient temperatures being approximately 25 °C lower. Combined with resting metabolic rates (RMRs), this gives them an exceptionally low metabolic scope in winter (DEE/RMRt = 1.60 ± 0.30; RMRt is resting metabolic rate at thermoneutrality). Using implanted body temperature loggers and filming in the wild, we show that this is achieved by reducing body temperature and physical activity. Thyroid hormone (T(3) and T(4)) measurements indicate this metabolic suppression is probably mediated via the thyroid axis. Winter activity was lower at sites where domestic yak (Bos grunniens) densities were higher. Pikas supplement their food intake at these sites by eating yak feces, demonstrated by direct observation, identification of yak DNA in pika stomach contents, and greater convergence in the yak/pika microbiotas in winter. This interspecific coprophagy allows pikas to thrive where yak are abundant and partially explains why pika densities are higher where domestic yak, their supposed direct competitors for food, are more abundant.

Summary In many species, photo-identification could be used as an alternative to artificial marking to provide data on demographic parameters. However, unless the population is very small or fragmented, software may be required to pre-screen and reject most image pairs as potential matches. Depending on the species and method used to obtain images, currently available software may falsely reject some matches. We estimate the false rejection rate (FRR) of the ExtractCompare (EC) program when used to pre-screen images of female grey seals. Filtering images manually to reduce the FRR involves subjective assessment of image quality, reduces the amount of data available and may bias the results in favour of relatively well-marked individuals. The data may contain individuals identified only from the left side or the right side, as well as individuals identified from both sides. Missed matches resulting from false rejections by pre-screening software and/or inclusion of individuals identified only from opposite sides cause some individuals to generate multiple encounter histories. We describe an open population model for data of this type which, given a measured risk of missing a match between a randomly selected pair of images of the same individual, provides maximum likelihood (ML) estimates of initial population size, survival/emigration and immigration/recruitment by calculating the expected frequency of any encounter history that could be generated. As a case study for the method, we used EC to pre-screen photographs of female grey seals on a breeding colony and generate encounter histories over five successive seasons. Allowing for the measured FRR, we calculated ML estimates for comparison with estimates from previous studies. We also used the model with encounter histories simulated using the same FRR to give the same mixture of left side, right side and both sides histories and derived ML estimates for comparison with the values used to drive the simulation. With FRR set at up to 33%, the method gave estimates of the abundance and survival parameters used in the simulation model that were biased by at most 4·7% up and 3% down, respectively. The results of the grey seal case study were consistent with previous estimates of apparent survival and trends in abundance.

Energy stores are critical for successful breeding, and longitudinal studies require nonlethal methods to measure energy stores ("body condition"). Nonlethal techniques for measuring energy reserves are seldom verified independently. We compare body mass, size-corrected mass (SCM), plasma lipids, and isotopic dilution with extracted total body lipid content in three seabird species (thick-billed murres Uria lomvia, all four measures; northern fulmars Fulmarus glacialis, three measures; and black-legged kittiwakes Rissa tridactyla, two measures). SCM and body mass were better predictors of total body lipids for the species with high percent lipids (fulmars; R2 = 0.5-0.6) than for the species with low percent lipids (murres and kittiwakes; R2 = 0.2-0.4). The relationship between SCM and percent body lipids, which we argue is often a better measure of condition, was also poor (R20.2) for species with low lipids. In a literature comparison of 17 bird species, percent lipids was the only predictor of the strength of the relationship between mass and total body lipids; we suggest that SCM be used as an index of energy stores only when lipids exceed 15% of body mass. Across all three species we measured, SCM based on the ordinary least squares regression of mass on the first principal component outperformed other measures. Isotopic dilution was a better predictor of both total body lipids and percent body lipids than were mass, SCM, or plasma lipids in murres. Total body lipids decreased through the breeding season at both sites, while total and neutral plasma lipid concentrations increased at one site but not another, suggesting mobilization of lipid stores for breeding. A literature review showed substantial variation in the reliability of plasma markers, and we recommend isotopic dilution (oxygen-18, plateau) for determination of energy reserves in birds where lipid content is below 15%.

We explored how seasonal changes in temperature, exercise and food supply affected energy metabolism and heart rate of Inuit dogs in Greenland. Using open flow respirometry, doubly labeled water, and heart rate recording, we measured metabolic rates of the same dogs at two different locations: at one location the dogs were fed with high energy food throughout the year while at the other location they were fed with low energy food during summer. Our key questions were: is resting metabolic rate (RMR) increased during the winter season when dogs are working? Does feeding regime affect RMR during summer? What is the proportion of metabolic rate (MR) devoted to specific dynamic action (SDA), and what is the metabolic scope of working Inuit sled dogs? The Inuit dogs had an extremely wide thermoneutral zone extending down to -25 degrees C. Temperature changes between summer and winter did not affect RMR, thus summer fasting periods were defined as baseline RMR. Relative to this baseline, summer MR was upregulated in the group of dogs receiving low energy food, whereas heart rate was downregulated. However, during food digestion, both MR and HR were twice their respective baseline values. A continuously elevated MR was observed during winter. Because temperature effects were excluded and because there were also no effects of training, we attribute winter elevated MR to SDA because of the continuous food supply. Working MR during winter was 7.9 times the MR of resting dogs in winter, or 12.2 times baseline MR.

We studied factors influencing daily energy expenditures (DEE) of male least weasels (Mustela nivalis) using the doubly labelled water technique. The relationship between ambient temperature and DEE formed a triangular pattern, characterized by invariance of the maximum DEE and an inverse relationship between minimum DEE and temperature. A simple energetic model relating the DEE of male weasels to activity time (AT) and ambient temperature predicted that, across seasons, less than 10 per cent of measurements approach the upper bound of observed DEE. Male weasels were able to maintain a relatively constant maximum energy output across varying temperatures by adjusting their AT to changes in temperature. They achieved maximum energy expenditures in winter due to high thermoregulatory costs, and in spring and summer due to high levels of physical activity. This pattern exemplifies a ‘metabolic niche’ of a small mammal having extremely high energy expenditures primarily driven by ambient temperature.

The effects of dietary antioxidant supplementation on oxidative stress and life span are confused. We maintained C57BL/6 mice at 7 +/- 2 degrees C and supplemented their diet with alpha-tocopherol from 4 months of age. Supplementation significantly increased (p = 0.042) median life span by 15% (785 days, n = 44) relative to unsupplemented controls (682 days, n = 43) and also increased maximum life span (oldest 10%, p = 0.028). No sex or sex by treatment interaction effects were observed on life span, with treatment having no effect on resting or daily metabolic rate. Lymphocyte and hepatocyte oxidative DNA damage and hepatic lipid peroxidation were unaffected by supplementation, but hepatic oxidative DNA damage increased with age. Using a cDNA macroarray, genes associated with xenobiotic metabolism were significantly upregulated in the livers of female mice at 6 months of age (2 months supplementation). At 22 months of age (18 months supplementation) this response had largely abated, but various genes linked to the p21 signaling pathway were upregulated at this time. We suggest that alpha-tocopherol may initially be metabolized as a xenobiotic, potentially explaining why previous studies observe a life span extension generally when lifelong supplementation is initiated early in life. The absence of any significant effect on oxidative damage suggests that the life span extension observed was not mediated via any antioxidant properties of alpha-tocopherol. We propose that the life span extension observed following alpha-tocopherol supplementation may be mediated via upregulation of cytochrome p450 genes after 2 months of supplementation and/or upregulation of p21 signaling genes after 18 months of supplementation. However, these signaling pathways now require further investigation to establish their exact role in life span extension following alpha-tocopherol supplementation.

SUMMARY Many small mammals respond to seasonal changes in photoperiod by altering body mass and adiposity. These animals may provide valuable models for understanding the regulation of energy balance. Here, we present data on the field vole (Microtus agrestis) – a previously uncharacterised example of photoperiod-induced changes in body mass. We examined the effect of increased day length on body mass, food intake, apparent digestive efficiency,body composition, de novo lipogenesis and fatty acid composition of adipose tissue in cold-acclimated (8°C) male field voles by transferring them from a short (SD, 8 h:16 h L:D) to long day photoperiod (LD, 16 h:8 h L:D). During the first 4 weeks of exposure to LD, voles underwent a substantial increase in body mass, after which the average difference between body masses of LD and SD voles stabilized at 7.5 g. This 24.8% increase in body mass reflected significant increases in absolute amounts of all body components, including dry fat mass, dry lean mass and body water mass. After correcting body composition and organ morphology data for the differences in body mass, only gonads (testes and seminal vesicles) were enlarged due to photoperiod treatment. To meet energetic demands of deposition and maintenance of extra tissue, voles adjusted their food intake to an increasing body mass and improved their apparent digestive efficiency. Consequently, although mass-corrected food intake did not differ between the photoperiod groups, the LD voles undergoing body mass increase assimilated on average 8.4 kJ day-1 more than animals maintained in SD. The majority(73–77%) of the fat accumulated as adipose tissue had dietary origin. The rate of de novo lipogenesis and fatty acid composition of adipose tissue were not affected by photoperiod. The most important characteristics of the photoperiodic regulation of energy balance in the field vole are the clear delineation between phases where animals regulate body mass at two different levels and the rate at which animals are able to switch between different levels of energy homeostasis. Our data indicate that the field vole may provide an attractive novel animal model for investigation of the regulation of body mass and energy homeostasis at both organism and molecular levels.

Many seabirds in the North Sea feed on lesser sandeels Ammodytes marinus during the breed- ing season. Unprecedented breeding failures were recorded at many seabird colonies on the east coast of Britain in 2004. We used demographic, dietary and behavioural data from a long-term study of a colony of common guillemots Uria aalge, the most abundant seabird species in the North Sea, to set the 2004 season in context. Birds at this colony showed greatly reduced breeding success and those chicks that did survive left the colony in very poor condition. The main prey item fed to chicks in 2004 was sprat Sprattus sprattus rather than sandeels, and parents increased the amount of time spent foraging, frequently leaving chicks unat- tended in order to maintain a normal feeding rate. The calculated daily food intake of chicks derived from these values did not differ markedly from previous years and therefore the magnitude of the impact on chick growth and breeding success appeared dispro- portionately large. However, nutrient analyses of fish collected from birds in 2004 revealed them to be of sig- nificantly lower energy value than expected. Poor food quality therefore appeared to be the proximate cause of seabird breeding failure in 2004 giving support to the 'junk-food' hypothesis. Single-prey loaders such as guillemots will be particularly sensitive to reductions in the energy value of food items. The reasons for the poor fish condition in this part of the North Sea are currently unknown, but the results provide further evidence of major changes in the North Sea food web.

Gray seals (Halichoerus grypus) in the UK exhibit clear preferences for pupping close to pools of water on inland breeding sites. The reasons for this are unclear as seals are thought to derive all their water requirements from the metabolism of fat. Likewise, the prospect of seals overheating during the UK's cool, damp, autumnal breeding seasons, has seemed unlikely, but has not been previously explored. Here, we provide preliminary behavioral evidence of thermal stress in female gray seals breeding on the island of North Rona, Scotland. Video footage provided measures of proximity to, and the proportion of animals bathing in, pools in relation to meteorological data (temperature, mean sea-level pressure, rainfall, wind speed and direction) on four dates spread through a single breeding season. The proportion of females close to pools increased with pressure (warmer, drier conditions) and decreased on wetter days. In addition, analyses of colony-wide patterns of seal dispersion showed that females tended to be closer to pools of water at higher pressures and temperatures and at lower wind-speeds. These results provide the first evidence of thermal stress and behavioral thermoregulation through access to pools of water in a phocid breeding in temperate autumnal conditions. Postprint

Colonially breeding animals such as grey seals typically have a discrete number of breeding sites. The extent to which offspring are philopatric (return to breed at their natal site) and the prevalence of philopatry by sex of pup has fundamental consequences for the social and genetic structure within and between each colony. Grey seals born and marked by tagging and cohort brands at colonies on North Rona, Outer Hebrides since 1960 and at the Isle of May, Firth of Forth since 1981 have shown philopatry. Overall re-sight rates of 1620 pups tagged at North Rona and 1667 pups at the Isle of May ranged from 0 to 17%. Although most evidence of philopatry relates to females, males at each colony displayed philopatry. In addition, females born at N. Rona pupped closer to their natal sites than would be expected by chance (p = 0.005), providing evidence of fine scale natal site fidelity. Females born at the Isle of May did not show the same degree of fine scale natal site fidelity, but there was considerable individual variation. The spatial scale of philopatry shown by seals differed according to the local area within each island. Fidelity of known adult females to pupping sites at both colonies was high. In addition, we found an instance of mother and daughter pupping together away from the daughter's natal site. These data: (i) provide evidence of philopatry in both sexes at each site; (ii) describe occurrences of fine scale philopatry at both sites; (iii) show that mothers and offspring occupied the same areas on the colony, suggesting the possibility of highly related groups within colonies, which would provide conditions for kin-specific altruistic behaviours; (iv) suggest the possibility of individual or kin recognition, even though contact between mothers and offspring has been thought to cease after the 18 d lactation period.

Hunting dogs display an extraordinary working capacity. They can work for several hours a day for a week or more and cover considerable running distances. The great physical effort displayed by hunting dogs has led to an increased desire among dog breeders for greater knowledge of their specific nutrient requirements, especially for energy and drinking water, which are crucial for endurance. Most experiments regarding sporting dogs have been conducted with Greyhounds and sled dogs, representing the extremes in dog racing, as Greyhounds are sprinters relying on both anaerobic and aerobic fuel, whereas sled dogs are long-distance runners using aerobic energy. Hunting dogs (gun dogs) are likely intermediate in their energy usage, primarily exhibiting endurance activity for several hours interrupted with bouts of sprinting (1). There is also considerable interest among dog owners regarding distance covered and actual speed of their working dogs. The distance covered is likely a major factor influencing the energy spent by the hunting dog. In addition, environmental factors such as air temperature, ground conditions, terrain, and vegetation must have significant influence, although this has to our knowledge never been studied in hunting dogs. The present pilot study aimed at measuring energy expenditure (EE) 5 and body water turnover (BWT) in hunting dogs. Three different running conditions were studied: hunting, road running in harness, which is a common used training activity for hunting dogs, and running on a treadmill, focusing on indications of differences in EE as a result of different ground conditions. The doubly labeled water technique was used to determine EE, and the repeatability of this method was examined by letting the same dogs undertake similar exercise 3 times.

WILLIAM D. PATERSON and PAULA REDMAN, Scottish Oceans Institute, Sea Mammal Research Unit, University of St. Andrews, St. Andrews, KY16 8LB, United Kingdom; LEX A. HIBY, Conservation Research Ltd., Cambridge CB22 5AL, United Kingdom; SIMON E. W. MOSS, AILSA J. HALL and PADDY POMEROY, Scottish Oceans Institute, Sea Mammal Research Unit, University of St. Andrews, St. Andrews, KY16 8LB, United Kingdom.

Capsule. Female Great Tit Parus major feeding rate behaviour was highly variable between individuals, but no effect of the doubly labelled water (DLW) procedure on female visit rate was detected, whereas visit rates by males, which were neither trapped or manipulated, increased following trapping and manipulation of their partners. Aims. To test the assumption that the subject’s behaviour is normal during the measurement period when using DLW to measure energy expenditure. Methods. Visits to the nest box by parents feeding young were counted separately for females and males on the day before and the day after the female only was trapped to measure her energy expenditure using DLW. Visit rates were also counted for control pairs. Results. Female visit rates did not differ before and after manipulation, or between experimental and control pairs, but bird behaviour was highly variable between individuals. In contrast, the visit rates of both experimental and control males, which were not trapped, increased on the second day. Conclusion. The results for female behaviour supported the assumption of normality, but a small subset of particular individuals may be prone to adverse reactions. The response of the males may have been a reaction to disturbance at the nest.

Oxidative stress is suggested to be central to the ageing process, with endogenous antioxidant defence and repair mechanisms in place to minimize damage. Theoretically, supplementation with exogenous antioxidants might support the endogenous antioxidant system, thereby reducing oxidative damage, ageing-related functional decline and prolonging life- and health-span. Yet supplementation trials with antioxidants in animal models have had minimal success. Human epidemiological data are similarly unimpressive, leading some to question whether vitamin C, for example, might have pro-oxidant properties in vivo. We supplemented cold exposed (7+/-2 degrees C) female C57BL/6 mice over their lifespan with vitamin C (ascorbyl-2-polyphosphate), widely advocated and self administered to reduce oxidative stress, retard ageing and increase healthy lifespan. No effect on mean or maximum lifespan following vitamin C treatment or any significant impact on body mass, or on parameters of energy metabolism was observed. Moreover, no differences in hepatocyte and lymphocyte DNA oxidative damage or hepatic lipid peroxidation was seen between supplemented and control mice. Using a DNA macroarray specific for oxidative stress-related genes, we found that after 18 months of supplementation, mice exhibited a significantly reduced expression of several genes in the liver linked to free-radical scavenging, including Mn-superoxide dismutase. We confirmed these effects by Northern blotting and found additional down-regulation of glutathione peroxidase (not present on macroarray) in the vitamin C treated group. We suggest that high dietary doses of vitamin C are ineffective at prolonging lifespan in mice because any positive benefits derived as an antioxidant are offset by compensatory reductions in endogenous protection mechanisms, leading to no net reduction in accumulated oxidative damage.

We have previously shown that cold-acclimated (8 degrees C) male field voles (Microtus agrestis) transferred from short (SD, 8:16 h L:D) to long photoperiod (LD, 16:8 h L:D) exhibit increases in body mass, adiposity and food intake. To assess whether these increases were associated with decreased leptin sensitivity, we infused LD and SD voles with physiological doses of murine leptin (or saline) delivered peripherally for 7 days via mini-osmotic pumps. Measurements were made of body mass (weight-reducing effect of leptin), food intake (anorectic effect of leptin) and gene expression of uncoupling protein 1 (UCP1) in brown adipose tissue (BAT) (thermogenic effect of leptin). The SD animals were sensitive to the weight-reducing effects of leptin (mean body mass decrease of 1.2 g over 7 days) and appetite-reducing effect of leptin (mean food intake decrease of 2.5 g over 7 days), whereas LD voles were resistant to the hormone treatment. The switch from a leptin-sensitive to leptin-resistant state appears to act as a desensitisation mechanism that allows voles transferred from SD to LD to ignore elevated leptin levels generated by increased body fat and accumulate adipose tissue without stimulating compensatory changes opposing the weight gain. Neither SD nor LD voles responded to infusion of leptin by changes in BAT UCP1 gene expression, suggesting dissociation of anorectic and thermogenic effects of leptin, possibly related to chronic cold exposure. Our results indicate that cold-acclimated voles show photoperiod-regulated changes in leptin sensitivity and may provide an attractive model for elucidating molecular mechanisms of leptin resistance.

Two theories of how energy metabolism should be associated with longevity, both mediated via free-radical production, make completely contrary predictions. The 'rate of living-free-radical theory' (Pearl, 1928; Harman, 1956; Sohal, 2002) suggests a negative association, the 'uncoupling to survive' hypothesis (Brand, 2000) suggests the correlation should be positive. Existing empirical data on this issue is contradictory and extremely confused (Rubner, 1908; Yan & Sohal, 2000; Ragland & Sohal, 1975; Daan et al., 1996; Wolf & Schmid-Hempel, 1989]. We sought associations between longevity and individual variations in energy metabolism in a cohort of outbred mice. We found a positive association between metabolic intensity (kJ daily food assimilation expressed as g/body mass) and lifespan, but no relationships of lifespan to body mass, fat mass or lean body mass. Mice in the upper quartile of metabolic intensities had greater resting oxygen consumption by 17% and lived 36% longer than mice in the lowest intensity quartile. Mitochondria isolated from the skeletal muscle of mice in the upper quartile had higher proton conductance than mitochondria from mice from the lowest quartile. The higher conductance was caused by higher levels of endogenous activators of proton leak through the adenine nucleotide translocase and uncoupling protein-3. Individuals with high metabolism were therefore more uncoupled, had greater resting and total daily energy expenditures and survived longest - supporting the 'uncoupling to survive' hypothesis.

Nest construction is an extremely widespread behaviour. In small endotherms the nest serves primarily to provide insulation, and thereby retard heat loss of the constructor, or its offspring. In arctic and temperate regions many small mammals build nests to protect themselves from low ambient temperatures. We measured the physical properties of nests built by short-tailed field voles Microtus agrestis that were kept in captivity under cold conditions. The most important factor influencing nest insulation was nest wall thickness; however, nests with thick walls also contained more nesting material. Insulative capacity of the nest did not reach an asymptote up to nests containing 20 g of material. Nest insulation was not correlated with resting metabolic rate, body mass or body composition of the vole that constructed the nest. However, nests built by males had greater insulation than those made by females; males also had significantly lower food intake rates when compared to females with nests. No significant difference was observed in either fat mass or whole animal thermal conductance between males and females. Thermal conductance did increase significantly with increasing body mass, although not with resting metabolic rate. Voles with nests for prolonged periods had lower food intakes than voles without nests. The absolute saving averaged 1.9 g and was independent of body mass. This was a 28% saving on intake for a 22-g vole but only an 18% saving for a 40-g individual. When voles had nests for short periods they used the energy they saved to reduce food intake and increase body mass.