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     

Adaptations to fasting in the American mink (<Emphasis Type="Italic">Mustela vison</Emphasis>): nitrogen metabolism

The aim of this study was to investigate the adaptations of protein metabolism to seasonal fasting in an actively wintering boreal carnivore. Fifty farm-bred male American minks Mustela vison were divided into a fed control group and four experimental groups fasted for 2, 3, 5 or 7 days. The responses of nitrogen metabolism to wintertime food deprivation were determined by measuring the rate of weight loss, the tissue total protein concentrations and the plasma amino acid, urea, ammonia, uric acid and total protein levels. The mink has relatively poor adaptations to food deprivation, as it is not able to prolong phase II of fasting with fat as the major metabolic fuel. Instead, the species has to derive a part of its energy requirements from the breakdown of body proteins. The end product of protein catabolism—urea— accumulates in its circulation, and the mink may not be able to recycle urea-N. Although the mink can still have a high body fat percent at the end of the 7-day fast, it appears to enter phase III of fasting with stimulated proteolysis during this period.     

Blockade of fatty acid oxidation mimics phase II-phase III transition in a fasting bird,the king penguin

This study tests the hypothesis that the metabolic and endocrine shift characterizing the phase II-phase III transition during prolonged fasting is related to a decrease in fatty acid (FA) oxidation. Changes in plasma concentrations of various metabolites and hormones and in lipolytic fluxes, as determined by continuous infusion of [2-(3)H]glycerol and [1-(14)C]palmitate, were examined in vivo in spontaneously fasting king penguins in the phase II status (large fat stores, protein sparing) before, during, and after treatment with mercaptoacetate (MA), an inhibitor of FA oxidation. MA induced a 7-fold decrease in plasma beta-hydroxybutyrate and a 2- to 2.5-fold increase in plasma nonesterified fatty acids (NEFA), glycerol, and triacylglycerols. MA also stimulated lipolytic fluxes, increasing the rate of appearance of NEFA and glycerol by 60-90%. This stimulation might be partly mediated by a doubling of circulating glucagon, with plasma insulin remaining unchanged. Plasma glucose level was unaffected by MA treatment. Plasma uric acid increased 4-fold, indicating a marked acceleration of body protein breakdown, possibly mediated by a 2.5-fold increase in circulating corticosterone. Strong similarities between these changes and those observed at the phase II-phase III transition in fasting penguins support the view that entrance into phase III, and especially the end of protein sparing, is related to decreased FA oxidation, rather than reduced NEFA availability. MA could be therefore a useful tool for understanding mechanisms underlying the phase II-phase III transition in spontaneously fasting birds and the associated stimulation of feeding behavior.     

Bone marrow fat mobilization in relation to lipid and protein catabolism during prolonged fasting in barn owls

 To assess the role of bone marrow fat in survival during a period of negative energy balance, we investigated the relationship between the time-course of marrow fat mobilization and the metabolic states associated with body fuel utilization during a prolonged fast. In order to mimic the winter fast of the barn owl (Tyto alba), captive birds were subjected to fasts of various durations at 5 °C ambient temperature. Body mass and plasma metabolites were used to determine the metabolic state at the end of fasting. Skeleton composition remained unchanged throughout phase II of fasting, during which the birds essentially rely on lipid fuels. During the following phase III, characterized by an increase in net body protein breakdown, the lipid mass in skeleton marrow decreased sharply by 78%, concomitant with an increase of the bone water content. This marrow fat mobilization occurred in all parts of the skeleton. This observation supports the hypothesis that bone marrow fat is not only involved in local nutrition, but can also be used as a lipid reserve for total energy requirements. However, in contrast to other fat deposits, marrow fat is mobilized only during phase III of the fast, when the last shift from lipid to protein fuel metabolism occurs. Thus, metabolic and/or hormonal changes associated with this transition could be involved in bone marrow fat mobilization. Lastly, our results suggest that the measurement of bone marrow fat can be used as an accurate index of the nutritional status (i.e. phase II or phase III) in barn owls. Accepted: 4 July 1996     

Uric acid and urea in relation to protein catabolism in long-term fasting geese

Summary Five ganders were subjected to an experimental fast comparable to that which spontaneously occurs during breeding in domestic geese, and during migration and breeding in various wild birds. Plasma uric acid and urea concentrations, and their excretion as a proportion of total nitrogen excretion, were studied in relation to daily change in body mass per unit body mass, dm/mdt. This variable has previously been found to reflect changes in protein catabolism over the three phases of fast: I, dm/mdt and protein utilization both decrease; II, they are maintained at a low value; and III, they increase. In the fed state, daily total nitrogen excretion was 5 gN·24 h^–1; uric acid, ammonia and urea accounted for 51, 15 and 5% respectively. The high remaining proportion of, excreted nitrogen (29%), after subtraction of uric acid-N, ammonia-N and urea-N to total nitrogen, accords with the literature. During fasting, the changes in daily excretion of uric acid, urea, ammonia and total nitrogen followed a pattern essentially similar to that for dm/mdt. Uric acid accounted for a progressively increasing fraction of total nitrogen, up to 76% at the end of phase III, while urea remained at a constant 5%. Plasma concentrations of both uric acid and urea followed similar trends during the fast, in particular both increasing during phase III, i.e. when there was a rise in nitrogen exrection. This suggests they could be used as an index in field investigations, to determine whether birds which naturally fast in connection with specific activities have entered into the situation where proteins are no longer spared.     

Breeding and moulting fasts in macaroni penguins: Do birds exhaust their fat reserves?

• 1.1. Plasma concentrations of urea, uric acid and total lipid were compared in pre- and late-fast breeding and moulting macaroni penguins (Eudyptes chrysolophus) to test the hypothesis that birds exhaust their lipid reserves and initiate marked protein utilisation towards the end of natural fasts.
• 2.2. Male and female macaroni penguins fasted for a minimum of 29–32 days and 20 days during the breeding and moult fasts, and the difference in body weight over the sample period (reflecting body weight loss) was 31–34% and 41–47%, respectively.
• 3.3. There was no significant increase in plasma urea or uric acid at the end of either fast, nor any decrease in plasma lipid concentrations compared to pre-fast birds.
• 4.4. These results suggest that macaroni penguins continue to rely mainly on lipid reserves during the later stages of natural fasts. This is consistent with post-fast body composition data for other small penguin species.

     

Intermittent fasting during winter and spring affects body composition and reproduction of a migratory duck

We compared food intake, body mass and body composition of male and female black ducks (Anas rubripes) during winter (January-March). Birds were fed the same complete diet ad libitum on consecutive days each week without fasting (control; nine male; nine female) or with either short fasts (2 day x week(-1); nine male; nine female), or long fasts (4 day x week(-1); eleven male; twelve female). We continued treatments through spring (March-May) to measure the effect of intermittent fasts on body mass and egg production. Daily food intake of fasted birds was up to four times that of unfasted birds. Weekly food intake of males was similar among treatments (364 g x kg(-1) x week(-1)) but fasted females consumed more than unfasted females in January (363 g x kg(-1) x week(-1) vs. 225 g x kg(-1) x week(-1)). Although both sexes lost 10-14% body mass, fasted females lost less mass and lipid than unfasted females during winter. Total body nitrogen was conserved over winter in both sexes even though the heart and spleen lost mass while the reproductive tract and liver gained mass. Intermittent fasting increased liver, intestinal tissue and digesta mass of females but not of males. Fasting delayed egg production in spring but did not affect size, fertility or hatching of the clutch. Females on long fasts were still heavier than controls after laying eggs. Thus black ducks combine flexibility of food intake with plasticity of digestive tract, liver and adipose tissue when food supply is interrupted during winter. Females modulate body mass for survival and defer reproduction when food supply is interrupted in spring.     

Exogenous corticosterone mimics a late fasting stage in captive Adelie penguins (Pygoscelis adeliae)

Fasting is part of penguin's breeding constraints. During prolonged fasting, three metabolic phases occur successively. Below a threshold in body reserves, birds enter phase III (PIII), which is characterized by hormonal and metabolic shifts. These changes are concomitant with egg abandonment in the wild and increased locomotor activity in captivity. Because corticosterone (CORT) enhances foraging activity, we investigated the variations of endogenous CORT, and the effects of exogenous CORT on the behavioral, hormonal, and metabolic responses of failed breeder Adélie penguins. Untreated and treated captive male birds were regularly weighed and sampled for blood while fasting, and locomotor activity was recorded daily. Treated birds were implanted with various doses of CORT during phase II. Untreated penguins entering PIII had increased CORT (3.5-fold) and uric acid (4-fold; reflecting protein catabolism) levels, concomitantly with a rise in locomotor activity (2-fold), while prolactin (involved in parental care in birds) levels declined by 33%. In CORT-treated birds, an inverted-U relationship was obtained between CORT levels and locomotor activity. The greatest increase in locomotor activity was observed in birds implanted with a high dose of CORT (C100), locomotor activity showing a 2.5-fold increase, 4 days after implantation to a level similar to that of birds in PIII. Moreover, uric acid levels increased three-fold in C100-birds, while prolactin levels declined by 30%. The experimentally induced rise in CORT levels mimicked metabolic, hormonal, and behavioral changes, characterizing late fasting, thus supporting a role for this hormone in the enhanced drive for refeeding occurring in long-term fasting birds.     

Energy metabolism and body temperature of barn owls fasting in the cold

Energetic adaptation to fasting in the cold has been investigated in a nocturnal raptor, the barn owl (Tyto alba), during winter. Metabolic rate and body temperature (Tb) were monitored in captive birds, (1) after acute exposure to different ambient temperatures (Ta), and (2) during a prolonged fast in the cold (4 degrees C), to take into account the three characteristic phases of body fuel utilization that occur during a long-term but reversible fast. In postabsorptive birds, metabolic rate in the thermoneutral zone was 4. 1+/-0.1 W kg-1 and increased linearly below a lower critical temperature of 23 degrees C. Metabolic rate was 70% above basal at +4 degrees C Ta. Wet thermal conductance was 0.22 W kg-1 degrees C-1. During fasting in the cold, the mass-specific resting metabolic rate decreased by 16% during the first day (phase I) and remained constant thereafter. The amplitude of the daily rhythm in Tb was only moderately increased during phase II, with a slight lowering (0. 6 degrees C) in minimal diurnal Tb, but rose markedly in phase III with a larger drop (1.4 degrees C) in minimal diurnal Tb. Refeeding the birds ended phase III and reversed the observed changes. These results indicate that diurnal hypothermia may be used in long-term fasting barn owls and could be triggered by a threshold of body lipid depletion, according to the shift from lipid to protein fuel metabolism occurring at the phase II/phase III transition. The high cost of regulatory thermogenesis and the limited use of hypothermia during fasting may contribute to the high mortality of barn owls during winter.     

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.     

Physiological adaptations to fasting in an actively wintering canid, the Arctic blue fox (Alopex lagopus)

This study investigated the physiological adaptations to fasting using the farmed blue fox (Alopex lagopus) as a model for the endangered wild arctic fox. Sixteen blue foxes were fed throughout the winter and 32 blue foxes were fasted for 22 d in Nov-Dec 2002. Half of the fasted blue foxes were food-deprived again for 22 d in Jan-Feb 2003. The farmed blue fox lost weight at a slower rate (0.97-1.02% body mass d(-1)) than observed previously in the arctic fox, possibly due to its higher initial body fat content. The animals experienced occasional fasting-induced hypoglycaemia, but their locomotor activity was not affected. The plasma triacylglycerol and glycerol concentrations were elevated during phase II of fasting indicating stimulated lipolysis, probably induced by the high growth hormone concentrations. The total cholesterol, HDL- and LDL-cholesterol, urea, uric acid and total protein levels and the urea:creatinine ratio decreased during fasting. Although the plasma levels of some essential amino acids increased, the blue foxes did not enter phase III of starvation characterized by stimulated proteolysis during either of the 22-d fasting procedures. Instead of excessive protein catabolism, it is liver dysfunction, indicated by the increased plasma bilirubin levels and alkaline phosphatase, alanine aminotransferase and aspartate aminotransferase activities, that may limit the duration of fasting in the species.     

Comparative fuel metabolism in Gentoo and King Penguins: adaptation to brief versus prolonged fasting

To compare fuel utilization in large birds adapted to brief or prolonged fasting, protein and lipid utilization were quantified in the Gentoo Penguin (Pygoscelis papua) and the King Penguin (Aptenodytes patagonica). The inshore feeder Gentoo Penguin fasts for only a few days in its colony, while King Penguin chicks starve for several months in the subantarctic winter and male King Penguins starve for 5–6 weeks at the beginning of their breeding cycle. After an initial decrease in both daily body mass loss and nitrogen excretion during the first days of food deprivation, these two parameters thereafter stabilized at low values. At that time, protein utilization accounted for 15% of total energy expenditure in Gentoo Penguins and only 6% in King Penguin chicks during winter, the remainder (85% and 94%, respectively) being provided by fat oxidation. Similar percentages in fuel metabolism as seen in chicks during winter were reached in fasting adult King Penguins and spring chicks. However, a seasonal adaptation occurs in fasting chicks because energy expenditure is lower during winter. As previously described in starved mammals, the effectiveness in protein sparing could be related to the initial adiposity of the birds: the larger the fat stores (about 9% and 30% in Gentoo Penguins and winter chicks of King Penguins, respectively), the longer the fast duration and the better the level of protein conservation.     

Early changes in plasma hormones and metabolites during fasting in king penguin chicks

Summary Chicks of the king penguin (Aptenodytes patagonica) can tolerate a fast of 4–6 months during the subantarctic winter. The aim of this work was to study their initial response to food deprivation. Nine chicks were starved for 18 days. Two phases of starvation were defined according to changes in the specific daily loss in body mass: it decreased by 92% in phase I (6.6±0.3 days) and remained steady and low in phase II. Phase I was marked by a large decline in protein utilization, indicated by decreases in plasma levels of alanine (58%), uric acid (89%) and urea (76%) together with a decrease in circulating corticosterone (60%) and thyroxine (75%). In phase I, plasma insulin concentration decreased (61%) in some birds, but did not change in others; plasma pancreatic glucagon was stable whereas gut-glucagon decreased by 75%. Free fatty acids and -hydroxybutyrate concentrations gradually rode during the fast to 5 to 6 times pre-fast levels. Glycemia remained unchanged. Phase II was characterized by no change in plasma concentrations of protein-derived metabolites and by no or little change in circulating hormone levels. From comparison with previous data, we conclude that there are similar early adjustments to food deprivation in king penguin chick, rat and man: (1) a decrease in resting metabolic rate, (2) a decrease in protein utilization, and (3) mobilization of fat stores. The key, adaptations to long-term fasting in these species are therefore effectiveness in protein sparing and ability to prolong this situation.Abbreviations GLI glucagon-like immunoreactivity - PG pancreatic glucagon - -OHB -hydroxybutyrate - FFA free fatty acids - T ₃ triiodothyronine - T ₄ thyroxine     

Lipolytic and metabolic response to glucagon in fasting king penguins: phase II vs. phase III

This study aims to determine how glucagon intervenes in the regulation of fuel metabolism, especially lipolysis, at two stages of a spontaneous long-term fast characterized by marked differences in lipid and protein availability and/or utilization (phases II and III). Changes in the plasma concentration of various metabolites and hormones, and in lipolytic fluxes as determined by continuous infusion of [2-3H]glycerol and [1-14C]palmitate, were examined in vivo in a subantarctic bird (king penguin) before, during, and after a 2-h glucagon infusion. In the two fasting phases, glucagon infusion at a rate of 0.025 microg. kg(-1). min(-1) induced a three- to fourfold increase in the plasma concentration and in the rate of appearance (Ra) of glycerol and nonesterified fatty acids, the percentage of primary reesterification remaining unchanged. Infusion of glucagon also resulted in a progressive elevation of the plasma concentration of glucose and beta-hydroxybutyrate and in a twofold higher insulinemia. These changes were not significantly different between the two phases. The plasma concentrations of triacylglycerols and uric acid were unaffected by glucagon infusion, except for a 40% increase in plasma uric acid in phase II birds. Altogether, these results indicate that glucagon in a long-term fasting bird is highly lipolytic, hyperglycemic, ketogenic, and insulinogenic, these effects, however, being similar in phases II and III. The maintenance of the sensitivity of adipose tissue lipolysis to glucagon could suggest that the major role of the increase in basal glucagonemia observed in phase III is to stimulate gluconeogenesis rather than fatty acid delivery.     

Physiological responses to food deprivation in the house sparrow, a species not adapted to prolonged fasting

Many wild birds fast during reproduction, molting, migration, or because of limited food availability. Species that are adapted to fasting sequentially oxidize endogenous fuels in three discrete phases. We hypothesized that species not adapted to long fasts have truncated, but otherwise similar, phases of fasting, sequential changes in fuel oxidization, and similar changes in blood metabolites to fasting-adapted species. We tested salient predictions in house sparrows (Passer domesticus biblicus), a subspecies that is unable to tolerate more than ～32 h of fasting. Our main hypothesis was that fasting sparrows sequentially oxidize substrates in the order carbohydrates, lipids, and protein. We dosed 24 house sparrows with [(13)C]glucose, palmitic acid, or glycine and measured (13)CO(2) in their breath while they fasted for 24 h. To ascertain whether blood metabolite levels reflect fasting-induced changes in metabolic fuels, we also measured glucose, triacylglycerides, and β-hydroxybutyrate in the birds' blood. The results of both breath (13)CO(2) and plasma metabolite analyses did not support our hypothesis; i.e., that sparrows have the same metabolic responses characteristic of fasting-adapted species, but on a shorter time scale. Contrary to our main prediction, we found that recently assimilated (13)C-tracers were oxidized continuously in different patterns with no definite peaks corresponding to the three phases of fasting and also that changes in plasma metabolite levels accurately tracked the changes found by breath analysis. Notably, the rate of recently assimilated [(13)C]glycine oxidization was significantly higher (P < 0.001) than that of the other metabolic tracers at all postdosing intervals. We conclude that the inability of house sparrows to fast for longer than 32 h is likely related to their inability to accrue large lipid stores, separately oxidize different fuels, and/or spare protein during fasting.     

The effect of flight, fasting and p,p'-DDT on thyroid hormones and corticosterone in Gambel's white-crowned sparrow, Zonotrichia leucophrys gambelli

This study examined the effects of p,p'-dichlorodiphenyltrichloroethane (p,p'-DDT), fasting and flight on thyroid hormones and corticosterone in Gambel's White-crowned Sparrows (Zonotrichia leucophrys gambelli). Female sparrows were dosed daily with either 5 mg p,p'-DDT per kg body mass or corn oil vehicle over 3 days. On the fifth day the sparrows were divided into 3 groups: (1) unstressed - non-stressed control sparrows; (2) fasted - sparrows fasted for intervals ranging from 20 min to 9 h; or (3) flown - sparrows flown in a wind tunnel for intervals between 20 min and 2.5 h while fasting. Half the sparrows from each group received DDT (DDT-dosed sparrows) and the other half corn oil vehicle only (vehicle sparrows). Trunk blood plasma was analyzed for thyroxine, triiodothyronine and corticosterone using radioimmunoassay. In the flown group, corticosterone was elevated (DDT-dosed 35.52 ng/ml, P < or = 0.05), and thyroxine was depressed (DDT-dosed 4.09 ng/ml, P < or = 0.05; vehicle 4.33 ng/ml, P < or = 0.05). Elevated corticosterone likely decreased thyroid hormone production through a negative feedback mechanism originating at the hypothalamus. Mean triiodothyronine concentrations did not differ among any of the test groups. Relative to time fasted and flown, thyroxine decreased in flown birds dosed with DDT (P < 0.001) and triiodothyronine decreased in fasted birds dosed with DDT (P = 0.004). The increased rate of hormone diminution may be a result of the ability of DDT to induce microsomal enzyme production.     

Focus: The Science of Stress: The Impact of Maternal Antioxidants on Prenatal Stress Effects on Offspring Neurobiology and Behavior

Prenatal stress is a neuropsychiatric risk factor, and effects may be mediated by prenatal oxidative stress. Cell types in the brain sensitive to oxidative stress—cortical microglia and cortical and hippocampal interneurons—may be altered by oxidative stress generated during prenatal stress and may be neurobiological substrates for altered behavior. Our objective was to determine the critical nature of oxidative stress in prenatal stress effects by manipulating prenatal antioxidants. CD1 mouse dams underwent restraint embryonic day 12 to 18 three times daily or no stress and received intraperitoneal injections before each stress period of vehicle, N-acetylcysteine (200 mg/kg daily), or astaxanthin (30 mg/kg before first daily stress, 10 mg/kg before second/third stresses). Adult male and female offspring behavior, microglia, and interneurons were assessed. Results supported the hypothesis that prenatal stress-induced oxidative stress affects microglia; microglia ramification increased after prenatal stress, and both antioxidants prevented these effects. In addition, N-acetylcysteine or astaxanthin was effective in preventing distinct male and female interneuron changes; decreased female medial frontal cortical parvalbumin interneurons was prevented by either antioxidant; increased male medial frontal cortical parvalbumin interneurons was prevented by N-acetylcysteine and decreased male hippocampal GAD67GFP+ cells prevented by astaxanthin. Prenatal stress-induced increased anxiety-like behavior and decreased sociability were not prevented by prenatal antioxidants. Sensorimotor gating deficits in males was partially prevented by prenatal astaxanthin. This study demonstrates the importance of oxidative stress for persistent impacts on offspring cortical microglia and interneurons, but did not link these changes with anxiety-like, social, and sensorimotor gating behaviors.     

Plasma chemistry of the chinstrap penguin <Emphasis Type="Italic">Pygoscelis antarctica </Emphasis>during fasting periods: a case of poor adaptation to food deprivation?

The chinstrap penguin (Pygoscelis antarctica) is the smallest penguin species to be used to study the physiology of fasting. We analysed body-mass change and plasma chemistry of five non-breeding chinstraps during an experimental fasting period in the breeding season. We also analysed the same parameters in six fasting birds under natural conditions (during an incubation shift, which lasts about 10 days). Both groups presented similar patterns of change, showing a rapid increase in urea and uric acid plasma concentrations. Urea surpassed 3 mmol/l after 5 fasting days, while uric acid reached 1 mmol/l after 9 days. Plasma glucose levels decreased after 11 days, whereas cholesterol also showed a clear reduction during fasting. These results as a whole suggest that chinstrap penguins reached phase III after a short period in comparison with other Pygoscelis species. Body size and ecological factors could explain these inter-specific differences.     

Metabolic responses of homing pigeons to flight and subsequent recovery

This study examines metabolic changes occurring during short to endurance flights and during subsequent recovery in free-flying pigeons, in particular the change towards lipid utilization with increasing flight duration, lipid supply to the flight muscles, protein utilization and the time needed to metabolically recover. Eight plasma metabolite concentrations were measured in homing pigeons released from sites 20–200 km from the loft (0.3–4.8 h flight duration) just after landing and after keeping birds fasting at rest for 30 and 60 min, respectively, after their return. Birds kept in the loft fasting at rest were used as controls. Plasma free fatty acid and glycerol concentrations increased rapidly with flight duration and leveled off after about 1.5 h. This indicates a marked change towards a high and stable lipid utilization from adipose tissues within 1–2 h of flight. Plasma triglyceride levels and very-low-density lipoproteins were decreased after short flights, but subsequently regained or surpassed fasting levels at rest. This indicates that re-esterification of free fatty acids and delivery as very-low-density lipoproteins to the flight muscles to circumvent constraints of fatty acid supply, as described previously for small passerines, is not as significant in the pigeon which has a much lower mass-specific energy rate. An initial increase in plasma glucose levels and a transient decrease to fasting levels at rest was observed and may reflect the initial use and subsequent exhaustion of glycogen stores. Contrary to other birds and mammals, -hydroxy-butyrate levels increased markedly with flight duration. This may suggest a more important sparing of carbohydrates and protein as gluconeogenic precursors in the pigeon than in other species. Plasma uric acid levels increased linearly up to about 4 h flight duration. This indicates an accelerated protein breakdown during flight which may primarily serve to deliver amino acids as glucogenic precursors and citrate cycle intermediates. With increasing flight duration, the energy sources change from an initial phase based primarily on carbohydrates to a lipid-based endurance phase. It is discussed whether this metabolic change depends on the level of power output or the performed work (energy spent) since the start of flight. During the first hour of recovery, most metabolites reached or approached fasting levels at rest, indicating a marked reduction in lipolysis and protein breakdown. -hydroxy-butyrate levels remained at flight levels and glucose levels increased slightly, indicating a restoration of glycogen stores.Abbreviations VLDL very-low-density lipoproteins - FFA free fatty acids     

The Effect of Social Stress on Chronic Pain Perception in Female and Male Mice

The current investigations on social stress primarily point to the negative health consequences of being in a stressful social hierarchy. The repetitive nature of such stressors seems to affect behavioral response to pain both in rodents and humans. Moreover, a large discrepancy in the possibility of social stresses affecting pain perception in the two genders exists. The present study examined the effect of chronic social stress on nociceptive responses of both sexes by implementing of food deprivation, food intake inequality and unstable social status (cage-mate change every 3 days) for a period of 14 days in 96 Balb/c mice. In this regard we injected 20 µl formalin 2% into the plantar surface of hind paw at the end of stress period and scored pain behaviors of all subjects, then serum concentrations of proinflammatory cytokines were measured. Our results showed that there was significant difference in chronic phase of formalin test following implementation of food deprivation and inequality (P<0.05) as compared to control group, so that pain perception was decreased considerably and this decline in inequality exposed subjects was well above isolated ones (P<0.05); whereas unstable social situation did not affect pain perception. Moreover, IL-1 and IL-6 concentrations in serum of stressed mice of both genders were well above control group (p<0.05). Finally, despite chronic pain perception in control and unstable male subjects was larger than females; the decrease of chronic pain perception in male stressed animals (poverty and inequality experienced subjects) was much more than stressed females. These results revealed that although food deprivation and social inequality can induce hypoalgesia, some socioeconomic situations like social instability don''t affect pain sensation, whereas there were similar increases of proinflammatory cytokines level in all socially stressed subjects. In addition, males display larger hypoalgesic responses to inequality as compared with females.