Prehatching temperatures drive inter-annual cohort differences in great tit metabolism.

Basal metabolic rate (BMR) constitutes the lowest metabolic rate in a resting animal and is, therefore, considered to reflect the energetic cost of maintenance in endotherms. BMR is a reversible plastic trait that changes with environmental and ecological circumstances, albeit being heritable and susceptible to selection. Inter-individual variation within populations of small birds is substantial, and while many of the drivers of such variation have been identified, many remain unexplained. We studied winter BMR variation of juveniles over a 15-year period in a wild population of great tits Parus major at the northern border of their distribution. BMR during winter consistently changed between years, even after controlling for environmental factors, suggestive of a non-reversible developmental plasticity shaping the adult metabolic phenotype. BMR in cohorts of wintering great tits varied among winters as a response to minimum ambient temperatures experienced early in life, during the prehatching period. This developmental plasticity might be adaptive if temperatures experienced by growing embryos would metabolically prime them to an environment that they will likely encounter in future life. However, in line with a more unpredictable future climate, the risk of phenotype-environment mismatch is likely to lead to certain cohorts being poorly adapted to prevailing winter conditions, resulting in wider annual fluctuations in population size.

Mitochondrial 2,4-dienoyl-CoA reductase (Decr) deficiency and impairment of thermogenesis in mouse brown adipose tissue.

A large number of studies have demonstrated significance of polyunsaturated fatty acids (PUFAs) for human health. However, many aspects on signals translating PUFA-sensing into body homeostasis have remained enigmatic. To shed light on PUFA physiology, we have generated a mouse line defective in mitochondrial dienoyl-CoA reductase (Decr), which is a key enzyme required for ß-oxidation of PUFAs. Previously, we have shown that these mice, whose oxidation of saturated fatty acid is intact but break-down of unsaturated fatty acids is blunted, develop severe hypoglycemia during metabolic stresses and fatal hypothermia upon acute cold challenge. In the current work, indirect calorimetry and thermography suggested that cold intolerance of Decr-/- mice is due to failure in maintaining appropriate heat production at least partly due to failure of brown adipose tissue (BAT) thermogenesis. Magnetic resonance imaging, electron microscopy, mass spectrometry and biochemical analysis showed attenuation in activation of lipolysis despite of functional NE-signaling and inappropriate expression of genes contributing to thermogenesis in iBAT when the Decr-/- mice were exposed to cold. We hypothesize that the failure in turning on BAT thermogenesis occurs due to accumulation of unsaturated long-chain fatty acids or their metabolites in Decr-/- mice BAT suppressing down-stream propagation of NE-signaling.

Mass or pace? Seasonal energy management in wintering boreal passerines.

Research on winter energy management in small vertebrates has focused on the regulation of body mass (BM) within a framework of starvation-predation trade-off. Winter-acclimatized birds exhibit a seasonal increase in both BM and basal metabolic rate (BMR), although the patterns of co-variation between the two traits remain unknown. We studied this co-variation in three different species of wild titmice, great, blue and willow tits, originating from two boreal regions at different latitudes. Seasonal change in BM and BMR was inter-dependent, particularly in the great tit; however, by contrast, no seasonal change was observed in the willow tit. BMR changed non-linearly in concert with BM with a peak in midwinter for both blue and great tits, whereas such non-linear pattern in willow tit was opposite and independent of BM. Surprisingly, BMR appears to be more sensitive to ambient temperatures than BM in all three species studied. Energy management is a multifaceted strategy that cannot be fully understood without considering reserve levels and energy expenditure simultaneously. Thus, our study indicates that the prevailing conceptual framework based on variation in BM alone is insufficient to understand seasonal energy management in small wintering passerines.

Northward expanding resident species benefit from warming winters through increased foraging rates and predator vigilance.

Species distributions shift northwards due to climate change, but the ecological mechanisms allowing range expansions are not fully understood. Most studies have concentrated on breeding seasons, but winter warming may also be important. Wintering distributions are restricted by food availability and temperature, which may also interact. Foraging in cold conditions requires adaptations as individuals have to be efficient in foraging, while staying warm and vigilant for predators. When the ambient temperature declines, foraging rates should be reduced due to increased time spent on warming behaviours. In addition, predator vigilance should decline, because more time has to be invested in foraging. Cold weather should limit northward expanding southern species in particular, while northern species should perform better in cold conditions. We tested this by studying temperature responses (between 0 and - 35 °C) among wintering birds at feeders. We compared foraging behaviours of two northward expanding southern species, the great tit (Parus major) and the blue tit (Cyanistes caeruleus) to a northern species, the willow tit (Poecile montanus). Foraging rate and vigilance decreased, and warming behaviour increased when temperatures declined. Importantly, the performance in these traits was poorer in the southern species compared to the willow tit. Furthermore, the response to decreasing temperatures in foraging rates and warming behaviour was stronger in the great tits than willow tits. As the winters become warmer, these mechanisms should increase wintering success of southern species wintering at high latitudes, and lead to higher survival, increased population growth, and consequent range expansion.

Elucidating mechanisms for insect body size: partial support for the oxygen-dependent induction of moulting hypothesis.

Body size is a key life history trait, and knowledge of its mechanistic basis is crucial in life history biology. Such knowledge is accumulating for holometabolous insects, whose growth is characterised and body size affected by moulting. According to the oxygen-dependent induction of moulting (ODIM) hypothesis, moult is induced at a critical mass at which oxygen demand of growing tissues overrides the supply from the tracheal respiratory system, which principally grows only at moults. Support for the ODIM hypothesis is controversial, partly because of a lack of proper data to explicitly test the hypothesis. The ODIM hypothesis predicts that the critical mass is positively correlated with oxygen partial pressure (PO2 ) and negatively with temperature. To resolve the controversy that surrounds the ODIM hypothesis, we rigorously test these predictions by exposing penultimate-instar Orthosia gothica (Lepidoptera: Noctuidae) larvae to temperature and moderate PO2  manipulations in a factorial experiment. The relative mass increment in the focal instar increased along with increasing PO2 , as predicted, but there was only weak suggestive evidence of the temperature effect. Probably owing to a high measurement error in the trait, the effect of PO2  on the critical mass was sex specific; high PO2  had a positive effect only in females, whereas low PO2  had a negative effect only in males. Critical mass was independent of temperature. Support for the ODIM hypothesis is partial because of only suggestive evidence of a temperature effect on moulting, but the role of oxygen in moult induction seems unambiguous. The ODIM mechanism thus seems worth considering in body size analyses.

[Birds' sense of direction]. / Lintujen suuntavaisto.

Birds utilize several distinct sensory systems in a flexible manner in their navigation. When navigating with the help of landmarks, location of the sun and stars, or polarization image of the dome of the sky, they resort to vision. The significance of olfaction in long-range navigation has been under debate, even though its significance in local orientation is well documented. The hearing in birds extends to the infrasound region. It has been assumed that they are able to hear the infrasounds generated in the mountains and seaside and navigate by using them. Of the senses of birds, the most exotic one is the ability to sense magnetic fields of the earth.

Different Seasonal Patterns in Song System Volume in Willow Tits and Great Tits.

In most species of seasonally breeding songbirds studied to date, the brain areas that control singing (i.e. the song control system, SCS) are larger during the breeding season than at other times of the year. In the family of titmice and chickadees (Paridae), one species, the blue tit (Cyanistes caeruleus), shows the typical pattern of seasonal changes, while another species, the black-capped chickadee (Poecile atricapillus), shows, at best, very reduced seasonal changes in the SCS. To test whether this pattern holds up in the two Parid lineages to which these two species belong, and to rule out that the differences in seasonal patterns observed were due to differences in geography or laboratory, we compared the seasonal patterns in two song system nuclei volumes (HVC and Area X) in willow tits (Poecile montanus), closely related to black-capped chickadees, and in great tits (Parus major), more closely related to blue tits, from the same area around Oulu, Finland. Both species had larger gonads in spring than during the rest of the year. Great tit males had a larger HVC in spring than at other times of the year, but their Area X did not change in size. Willow tits showed no seasonal change in HVC or Area X size, despite having much larger gonads in spring than the great tits. Our findings suggest that the song system of willow tits and their relatives may be involved in learning and producing nonsong social vocalizations. Since these vocalizations are used year-round, there may be a year-round demand on the song system. The great tit and blue tit HVC may change seasonally because the demand is only placed on the song system during the breeding season, since they only produce learned vocalizations during this time. We suggest that changes were not observed in Area X because its main role is in song learning, and there is evidence that great tits do not learn new songs after their first year of life. Further study is required to determine whether our hypothesis about the role of the song system in the learned, nonsong vocalizations of the willow tit and chickadee is correct, and to test our hypothesis about the role of Area X in the great tit song system.

USF1 deficiency activates brown adipose tissue and improves cardiometabolic health.

USF1 (upstream stimulatory factor 1) is a transcription factor associated with familial combined hyperlipidemia and coronary artery disease in humans. However, whether USF1 is beneficial or detrimental to cardiometabolic health has not been addressed. By inactivating USF1 in mice, we demonstrate protection against diet-induced dyslipidemia, obesity, insulin resistance, hepatic steatosis, and atherosclerosis. The favorable plasma lipid profile, including increased high-density lipoprotein cholesterol and decreased triglycerides, was coupled with increased energy expenditure due to activation of brown adipose tissue (BAT). Usf1 inactivation directs triglycerides from the circulation to BAT for combustion via a lipoprotein lipase-dependent mechanism, thus enhancing plasma triglyceride clearance. Mice lacking Usf1 displayed increased BAT-facilitated, diet-induced thermogenesis with up-regulation of mitochondrial respiratory chain complexes, as well as increased BAT activity even at thermoneutrality and after BAT sympathectomy. A direct effect of USF1 on BAT activation was demonstrated by an amplified adrenergic response in brown adipocytes after Usf1 silencing, and by augmented norepinephrine-induced thermogenesis in mice lacking Usf1. In humans, individuals carrying SNP (single-nucleotide polymorphism) alleles that reduced USF1 mRNA expression also displayed a beneficial cardiometabolic profile, featuring improved insulin sensitivity, a favorable lipid profile, and reduced atherosclerosis. Our findings identify a new molecular link between lipid metabolism and energy expenditure, and point to the potential of USF1 as a therapeutic target for cardiometabolic disease.

The activation of hepatic and muscle polyamine catabolism improves glucose homeostasis.

The mitochondrial biogenesis and energy expenditure regulator, PGC-1α, has been previously reported to be induced in the white adipose tissue (WAT) and liver of mice overexpressing spermidine/spermine N (1)-acetyltransferase (SSAT). The activation of PGC-1α in these mouse lines leads to increased number of mitochondria, improved glucose homeostasis, reduced WAT mass and elevated basal metabolic rate. The constant activation of polyamine catabolism produces a futile cycle that greatly reduces the ATP pools and induces 5'-AMP-activated protein kinase (AMPK), which in turn activates PGC-1α in WAT. In this study, we have investigated the effects of activated polyamine catabolism on the glucose and energy metabolisms when targeted to specific tissues. For that we used a mouse line overexpressing SSAT under the endogenous SSAT promoter, an inducible SSAT overexpressing mouse model using the metallothionein I promoter (MT-SSAT), and a mouse model with WAT-specific SSAT overexpression (aP2-SSAT). The results demonstrated that WAT-specific SSAT overexpression was sufficient to increase the number of mitochondria, reduce WAT mass and protect the mice from high-fat diet-induced obesity. However, the improvement in the glucose homeostasis is achieved only when polyamine catabolism is enhanced at the same time in the liver and skeletal muscle. Our results suggest that the tissue-specific targeting of activated polyamine catabolism may reveal new possibilities for the development of drugs boosting mitochondrial metabolism and eventually for treatment of obesity and type 2 diabetes.

Interpopulation variation in contour feather structure is environmentally determined in great tits.

BACKGROUND: The plumage of birds is important for flying, insulation and social communication. Contour feathers cover most of the avian body and among other functions they provide a critical insulation layer against heat loss. Feather structure and composition are known to vary among individuals, which in turn determines variation in the insulation properties of the feather. However, the extent and the proximate mechanisms underlying this variation remain unexplored. METHODOLOGY/PRINCIPAL FINDINGS: We analyzed contour feather structure from two different great tit populations adapted to different winter regimes, one northern population in Oulu (Finland) and one southern population in Lund (Sweden). Great tits from the two populations differed significantly in feather structure. Birds from the northern population had a denser plumage but consisting of shorter feathers with a smaller proportion containing plumulaceous barbs, compared with conspecifics from the southern population. However, differences disappeared when birds originating from the two populations were raised and moulted in identical conditions in a common-garden experiment located in Oulu, under ad libitum nutritional conditions. All birds raised in the aviaries, including adult foster parents moulting in the same captive conditions, developed a similar feather structure. These feathers were different from that of wild birds in Oulu but similar to wild birds in Lund, the latter moulting in more benign conditions than those of Oulu. CONCLUSIONS/SIGNIFICANCE: Wild populations exposed to different conditions develop contour feather differences either due to plastic responses or constraints. Environmental conditions, such as nutrient availability during feather growth play a crucial role in determining such differences in plumage structure among populations.

Specific appetite for carotenoids in a colorful bird.

BACKGROUND: Since carotenoids have physiological functions necessary for maintaining health, individuals should be selected to actively seek and develop a specific appetite for these compounds. METHODOLOGY/PRINCIPAL FINDINGS: Great tits Parus major in a diet choice experiment, both in captivity and the field, preferred carotenoid-enriched diets to control diets. The food items did not differ in any other aspects measured besides carotenoid content. CONCLUSIONS/SIGNIFICANCE: Specific appetite for carotenoids is here demonstrated for the first time, placing these compounds on a par with essential nutrients as sodium or calcium.

Enhanced polyamine catabolism alters homeostatic control of white adipose tissue mass, energy expenditure, and glucose metabolism.

Peroxisome proliferator-activated receptor gamma coactivator 1 alpha (PGC-1 alpha) is an attractive candidate gene for type 2 diabetes, as genes of the oxidative phosphorylation (OXPHOS) pathway are coordinatively downregulated by reduced expression of PGC-1 alpha in skeletal muscle and adipose tissue of patients with type 2 diabetes. Here we demonstrate that transgenic mice with activated polyamine catabolism due to overexpression of spermidine/spermine N(1)-acetyltransferase (SSAT) had reduced white adipose tissue (WAT) mass, high basal metabolic rate, improved glucose tolerance, high insulin sensitivity, and enhanced expression of the OXPHOS genes, coordinated by increased levels of PGC-1 alpha and 5'-AMP-activated protein kinase (AMPK) in WAT. As accelerated polyamine flux caused by SSAT overexpression depleted the ATP pool in adipocytes of SSAT mice and N(1),N(11)-diethylnorspermine-treated wild-type fetal fibroblasts, we propose that low ATP levels lead to the induction of AMPK, which in turn activates PGC-1 alpha in WAT of SSAT mice. Our hypothesis is supported by the finding that the phenotype of SSAT mice was reversed when the accelerated polyamine flux was reduced by the inhibition of polyamine biosynthesis in WAT. The involvement of polyamine catabolism in the regulation of energy and glucose metabolism may offer a novel target for drug development for obesity and type 2 diabetes.

Elevated plasma concentrations of haptoglobin in European brown bears during hibernation.

Haptoglobin (Hp), a hemoglobin-binding protein, is known as an acute phase protein and increases during the acute phase of inflammation in most mammals. We reported previously in brown bears that the mean Hp concentrations were higher in blood samples obtained in winter than those in spring. To examine a possible relation of the seasonal variations of Hp to hibernation, in the present study, we measured the plasma concentrations of Hp as well as some other acute phase proteins (alpha(2)-macroglobulin, alpha(1)-antitrypsin, C-reactive protein) in 6 European brown bears (Ursus arctos), from which blood samples were obtained at 5-6 different months of year including February, the time of hibernation. The Hp concentrations showed clear seasonal variations, being highest in February. The alpha(2)-macroglobulin concentrations also showed a similar but much smaller rise in February, but those of alpha(1)-antitrypsin and C-reactive protein did not show any seasonal variations. Our results suggest that the seasonal variation of plasma Hp concentration in brown bears is associated with a hibernation-specific mechanism more than that of acute phase response.

Local adaptation to winter conditions in a passerine spreading north: a common-garden approach.

Sedentary passerine birds living in temperate and boreal regions need a high metabolic capacity for thermogenesis to survive winter conditions. As a consequence of the increased thermogenic capacity, basal energetic demands rise at a time when resources and time to acquire them decrease. In a previous study, great tits (Parus major) from two localities in Fennoscandia with contrasting winter conditions differed in their metabolic response to ambient temperature. To investigate the physiological basis underlying interpopulation differences we performed a common-garden experiment to test whether these differences were genetically based. We found basal metabolic rate to be higher in birds originating from transferred eggs from the southern population compared to the ones from the northern population, contrary to the relationship among birds living in their region of origin. Despite previous evidence suggesting that gene flow prevents local adaptation at the northern range limits of a species expanding northward, we found that great tits differ in their reaction norm to winter conditions according to the population of origin.

Thermoregulatory adaptations of the overwintering captive raccoon dog (Nyctereutes procyonoides) in boreal climate.

The raccoon dog (Nyctereutes procyonoides) is a nocturnal canid thought to utilise passive wintering strategy in the boreal climate. To record the deep body temperature (T(b)), 12 farmed raccoon dogs were implanted with intra-abdominal T(b) loggers on November 26, 2003. Between December 3, 2003 and January 27, 2004 half of the animals were fasted for 8 weeks. The amplitude of the diurnal T(b) oscillations increased due to fasting. However, the mean diurnal T(b) was lower in the fasted animals only during two occasions. Unlike observed previously in other species, not only did the raccoon dogs experience hypothermia between 0600 and 1000 hr but also hyperthermia between noon and 1800 hr. The fasted animals were as active as the fed animals measured after 42-43 days of fasting and there was a significant cross-correlation between physical activity and T(b). The nocturnal period of hypothermia is probably an adaptation to save energy during food deprivation. The diurnal hyperthermia could be explained by the opportunistic foraging behaviour of the species. Opposite to the established assumptions, the raccoon dog does not seem to enter winter sleep on fur farms. In the future it is important to determine if true winter sleep occurs in nature in the species.

Force control of isometric elbow flexion with visual feedback in cold with and without shivering.

BACKGROUND: Exposure to cold impairs manual performance through effects on muscle tissue and control mechanisms. The purpose of this study was to assess the influence of increased muscle tone and shivering on ability to maintain required force during isometric flexion over a wide range of effort levels. METHODS: Lightly clad male subjects (n = 6) were exposed to thermoneutral air (TN, 27 degrees C) for 30 min, or to cold air (CA, 10 degrees C) for 30 min followed by a cold drink (1 L, 8 degrees C) to cause vigorous shivering (SH). At the end of each condition, subjects performed isometric elbow flexion at 10, 20, 40, and 80% of individual maximal voluntary contraction (MVC) for 10 s each, using digital visual feedback to control the force. We analyzed mean force output (F), the coefficient of force variation (FCv), information transmission (F/SD), and the coefficient of force auto-correlation (Ra), and the averaged electromyogram (aEMG) from elbow flexors, elbow extensor, and pectoral girdle muscles. RESULTS: Compared with TN, CA with increased muscle tone raised the aEMG by 5-30% but did not impair any of the force characteristics. In SH, F was not affected, while FCv and Ra were significantly increased at 10% MVC, while aEMG increased by 30-400% depending on the specific muscle and MVC level. CONCLUSION: Neither thermoregulatory muscle tone nor shivering influenced the control of force output during isometric elbow flexion, except that at the lowest MVC (10%) the force output was more variable during shivering.

Adaptive timing of digestion and digestion-related thermogenesis in the pigeon.

Pigeons were allowed to feed for 1 h either 2 h after lights on (morning pulse) or 3 h before lights off (evening pulse). Body temperature was measured radiotelemetrically. Faecal excretion, as an index of rate of digestion, was measured using load cells. At 22 degrees C, faecal excretion peaked just after lights-on in morning-pulse condition, but not in evening-pulse condition. In the cold (+5 degrees C), the peak was absent. We conclude that at thermoneutrality, pigeons are able to postpone a major part of digestion until late in the dark phase when their body temperature is increasing to the diurnal level. Thus, the extra heat from digestion-related thermogenesis that otherwise would be dissipated into the environment can be used for rewarming. In the cold, such a delay is not necessarily advantageous as the extra heat can always be used to substitute for thermoregulatory thermogenesis. The occurrence of concentrated period of digestion only in the morning-pulse condition was correlated to a lower food intake and lower body temperature as compared with the evening-pulse condition. Restricted feeding may thus be needed to induce adaptive timing of digestion as a mechanism of energy sparing. In addition to storage of food, timing of digestion may be a significant factor in the evolution of the crop in birds.

Cytochrome P450 enzyme activity in five herbivorous, non-passerine bird species.

We examined hepatic cytochrome P450 activity in wild and hand-reared grey partridges (Perdix perdix), capercaillies (Tetrao urogallus) and ring-necked pheasants (Phasianus colchicus), as well as the enzyme activity in a variety of tissues of hand-reared Japanese quails (Coturnix coturnix japonica) and pigeons (Columba livia). Post-mortem decrease in hepatic enzyme activity in the grey partridge was measured. Hepatic 7-ethoxyresorufin-O-deethylase activity was similar in wild and hand-reared grey partridges and pheasants, but the activity was significantly lower in wild than in hand-reared capercaillies, probably resulting from their phenolic-rich diet. In the tissues of both quails and pigeons 7-ethoxycoumarin-O-deethylase exhibited the highest and 7-pentoxyresorufin-O-deethylase the lowest activity. Hepatic enzyme activity was significantly higher than that in other tissues. In the small intestine some activity could be found, reflecting some intestinal detoxication capacity. Enzyme activity decreased by 34-69% during the 30-min sampling period, which confirmed the importance of equalising sampling time to obtain comparable data. Because the hand-reared birds in this study were fed the same commercial diets, we assumed that the enzyme activity values detected reflect species differences without any induction by dietary secondary compounds.