Annual variation in energy reserves in male three-spined stickleback, Gasterosteus aculeatm L. (Pisces, Gasterosteidae)

Wet and dry weights of tissue were measured and concentrations of glycogen, lipid and protein were estimated for the liver, gonad and carcass of male sticklebacks from an annual population collected each month over one complete year. In young-of-the-year there is one period of rapid weight gain, in all three body regions (liver, carcass and gonads) but particularly of the carcass, in the autumn and a second in spring and early summer. This is accompanied by an increase in the water content of all three body regions. The gonadosomatic index also increases sharply in spring and early summer. Young sticklebacks accumulate lipid and glycogen slowly during the autumn and winter of their first year of life and more rapidly from late winter to early summer. Thus, the period of most rapid accumulation of these reserves coincides with the time when body weight and gonad maturation are also increasing sharply. Lipid and glycogen levels fall during the reproductive season in those males that breed, so that by July they are reduced to 43% and 37% (respectively) of their peak values in May. Levels of protein increase throughout the year as the fish grow, but in breeding males by July the concentration of protein in the carcass falls to 70% of pre-breeding levels. Breeding males therefore reach the end of the reproductive season with their total energy reserves severely reduced, and consequently they suffer a very high mortality. In contrast, adult males that fail to reproduce survive beyond the breeding season. They continue to gain weight and to accumulate lipid and glycogen from August to September, but these energy reserves fall (to levels comparable to those of post-breeding fish) in December, when these fish probably die. These results demonstrate that in male sticklebacks, growth and gonad maturation can be sustained in parallel with the accumulation of energy reserves, which are subsequently extensively depleted as a result of reproductive activities.     

Larval history influences post-metamorphic condition in a coral-reef fish

Upon settlement, many fishes undergo an energetically costly metamorphic period that requires substantial nutritional reserves. Larval growth and the accumulation of lipids prior to metamorphosis are likely to influence growth and survival following this critical period. On the Caribbean island of St. Croix, I investigated relationships between larval growth, early life-history characteristics, and post-metamorphic lipid content in the bluehead wrasse Thalassoma bifasciatum. Lipid reserves remaining after metamorphosis were positively related (r2 = 0.62) to the width of the metamorphic band; thus, this otolith-derived trait may be used to estimate the condition at emergence of survivors collected at some later time. In contrast, pelagic larval duration, average larval growth, and otolith size at settlement were negatively related to post-metamorphic lipid content. Interestingly, the trend for slower growth among fish in good condition was not consistent over the entire pelagic larval duration. Analyses of daily larval growth histories indicated that fish with high lipid reserves grew rapidly in the last week prior to settlement, but relatively slowly during the early phases of larval life; those emerging with low lipid concentrations, however, displayed strikingly opposite patterns. These contrasting patterns of growth and energy storage were consistent at two sites and over three recruitment events. Otolith chemistry data suggested that differences in growth histories and body condition were consistent with the hypothesis of larval development in distinct oceanic environments (characterized by Pb concentration); but, within a water mass, differences reflected life-history trade-offs between growth and energy storage. The results have implications for understanding the processes driving juvenile survival, which may be condition dependent.     

Quantitative seasonal changes in the protein, lipid and energy content of the carcass, ovaries and liver of adult female plaice, Pleuronectes platessa L.

A detailed study was made of the seasonal changes in energy reserves in adult female plaice, Pleuronectes platessa L. The weights of lipid, protein, glycogen and ash in the carcass, liver and ovaries were measured in females approximately 35 cm long on ten occasions during one year. These data were converted to data for a'real'growing female by using a growth curve to take account of increasing length during the year. Growth of the whole body and of the body components was assumed to be isometric.
Plaice show a marked seasonal cycle of body weight and energy content. A six year old female plaice of 330 g, 32.6 cm long and with an energy content of 406 kcal in May, assimilates 560 kcal before December when it ceases to feed. Of this, 234 kcal is used to produce eggs, 159 kcal is used for metabolism during starvation, and the remainder 167 kcal, is used in growth. There is an increase in length to 35.9 cm and in weight to 462 g over the year. From December until March, when plaice do not feed but grow large ovaries, 40% of the protein in the body is utilised, 33% is devoted to egg production and 7% is metabolised to provide energy; 64% of the lipid is used, only 14% being used for egg production but 50% is metabolised for energy. Lipid supplies 75% of the energy for metabolism and so forms the major reserve; 42% of the energy assimilated during the year is devoted to reproduction. The main source of lipid and protein reserves is the carcass.     

Biochemical composition,condition index,and energy value of Ostrea peulchana (D'Orbigny): relationships with the reproductive cycle

Seasonal changes in body weight, biochemical composition, energy value, and condition index of Ostrea puelchana (D'Orbigny, 1846) from Laguna Punta Cero (Chubut, Argentina), were studied between February 1983 and April 1984. These values were determined for the soft tissues of the whole animal and, separately, from the adductor muscle, digestive gland-gonad complex, and a fraction containing all remaining organs. The main muscle components were proteins whereas glycogen and lipids were stored in the digestive gland-gonad complex. Weight and energy reserves decreased in winter during the beginning of the reproductive cycle and starvation period. In spring, there was a rapid growth of the gonad coincident with an increase in glycogen, lipid, and protein contents. In summer the glycogen content decreased with spawning, and proteins showed the highest values. The glycogen index appears to be suitable as an indicator of the physiological state. During the post-spawning period, in the autumn, the animals attained their best condition in quantity of flesh with a shigh energy value.     

Growth, maturation and reproductive investment in Arctic charr

Size and rates of growth in a cohort of 1 + Arctic charr housed in standard conditions were tracked over 12 months (December to December) and mature and immature males and females compared retrospectively. In both sexes, maturing fish were larger than non-maturing ones. In males, this size differential was the result of differences in growth in winter and early spring, but not in the remainder of the study period. In females, size differentials resulted mainly from growth rate differences immediately prior to breeding. In females but not in males, gonadosomatic index was predicted by growth rates in the months leading up to maturation, and among the females that matured, faster growing fish produced more eggs. Lipid reserves in July were correlated negatively with growth during the previous 7 months and, in females only, lipid reserves were significantly lower in maturing fish than in non-maturing fish, indicating that mobilization of lipid energy reserves in maturing fish had commenced by this time. Variation in investment in gonadal tissue, measured as gonadosomatic index, was not explained by variation in July lipid reserves for either males or females. However, July lipid reserves were negatively correlated with egg number, so females investing more in ova exhibited greater depletion of lipid reserves. These results are discussed in the context of the relationship between body condition and the onset of maturation in salmonids, relative investment in reproduction and sexual differences in the cost of reproduction.     

Seasonal changes in condition indices in adult mature and non-maturing burbot, Lota lota (L.), in the north-eastern Bothnian Bay, northern Finland

The population structure and seasonal changes in condition factors of the burbot in a shallow coastal region of the north-eastern Bothnian Bay are described. The significance of the so-called rest years is examined by comparing condition indices in immature or sterile and mature burbot. The somatic condition index ( K ₂), liver index ( K ₁), intestine index ( K ₁) and gonad index ( K_g ) are determined monthly in terms of organ weight in relation to body length. Approximately 30% of the whole catch of 1052 burbot were non-maturing but were 40 cm or more in length. K ₂, K ₁ and K ₁ were lowest in the autumn, when the first sign of gonad recrudescence was observed in mature burbot. The non-maturing burbot were never in poorer condition than mature ones. As mature and non-maturing burbot dissipated their energy stores during the warmest period of the summer, it is concluded that burbot spending a rest year do not accumulate and store energy reserves over the summer for the next year, and that such rest years, if they exist, do not occur for nutritional reasons.     

Lipid class dynamics and storage depots in juvenile weakfish Cynoscion regalis and their application to condition assessment

Lipid class dynamics, the pattern of change in the primary form and location of lipid stores and their relationship with standard length (L(S) ), were investigated in collections of young-of-the-year weakfish Cynoscion regalis for the purpose of determining the utility of this analysis as an indication of condition. The separation of total lipids into individual classes and the analysis of potential storage depots revealed the general patterns of lipid class dynamics and energy storage in C. regalis during their period of juvenile estuarine residency. Phospholipid and cholesterol exhibited moderate but variable (8·1-40·0 and 1·3-21·5 mg g(-1) , respectively) concentrations across the entire juvenile period and were the predominant lipid classes in juveniles <100 mm L(S) , while wax ester concentrations were low (c. 1 mg g(-1) ) and exhibited the least amount of variability among lipid classes. Triacylglycerols (TAG) and free fatty acids (FFA) exhibited similar dynamics, with relatively low concentrations (<15 mg g(-1) ) in individuals ≤100 mm L(S) . In larger juveniles both TAG and FFA concentrations generally increased rapidly, though there was considerable variability in both measures (0·0-199·7 and 0·0-49·7 mg g(-1) , respectively). Increasing levels of lipids, primarily in the form of TAG, with size indicated an accumulation of energy reserves with growth, thus providing an indication of individual condition for larger juveniles. Separate analysis of liver, viscera and the remaining carcass indicated that liver and viscera did not represent a significant depot of TAG reserves. Analysis of samples derived from whole juvenile C. regalis thus provided an accurate estimate of energy reserves.     

Influence of maturity status on the annual cycles of feeding and growth in Arctic charr reared at constant temperature

Temporal changes in feeding and growth of immature (1 +), maturing male (1 +), and mature male and female (3+) Arctic charr were studied by monitoring feed intake and growth of individually-marked fish for over 1 year. The fish were held at a constant temperature (4°C) under conditions of liberal feed supply. Feed intake and growth remained low in the period December-April, with only 20–60% of the fish having fed on each occasion that feed intake was monitored. Feed intake and growth increased markedly during the late spring and summer, but the increases were delayed by approximately 1–2 months in the immature fish compared with their maturing counterparts. Maturing fish decreased feeding during the late summer and by September there had been an almost complete cessation of feeding. This was reflected in changes in fish body weight and condition, both of which declined from August-September onwards. By contrast, the immature fish peaked in weight and condition in September, and weight loss during the autumn tended to be less rapid than observed for the maturing fish. The fish appeared to reduce feeding once a'threshold condition' of 1–4—1–5 had been attained. It is suggested that the fish may become anorexic once there has been replenishment of the energy reserves required for overwintering and, in the case of maturing individuals, for the completion of gonadal growth and Spawning.     

Effects of induced variation in anuran larval development on postmetamorphic energy reserves and locomotion

Anuran larvae exhibit high levels of phenotypic plasticity in growth and developmental rates in response to variation in temperature and food availability. We tested the hypothesis that alteration of developmental pathways during the aquatic larval stage should affect the postmetamorphic performance of the Iberian painted frog (Discoglossus galganoi). We exposed tadpoles to different temperatures and food types (animal- vs. plant-based diets) to induce variation in the length of the larval period and body size at metamorphosis. In this species, larval period varied with temperature but was unaffected by diet composition. In contrast, size at metamorphosis was shaped by the interaction between food quality and temperature; tadpoles fed on an animal-based diet became bulkier metamorphs than those fed on plant-based food at high (22°C) but not at low (12°C) temperature. Body condition of newly metamorphosed frogs was unrelated to the temperature or food type experienced during the premetamorphic stage. Frogs maintained at high temperature during the larval period showed reduced jumping ability, especially when fed on the plant-based diet. However, when considering size-independent jumping ability, cold-reared individuals exhibited the lowest performance, and herbivores reared at 17°C the highest. Cold-reared (12°C) frogs accumulated larger amounts of energy reserves than individuals raised at 17°C or 22°C. This was still the case after correction for differences in body mass, thus indicating some size-independent effect of developmental temperature. Despite the higher lipid content of the carnivorous diet, the differences in energy reserves between herbivores and carnivores were relatively weak and associated with differences in body size. These results suggest that the consequences of environmental variation in the larval habitat can extend to the terrestrial phase and influence juvenile growth and survival.     

Condition factor and hepatosomatic index as estimates of energy status in male three-spined stickleback

Seasonal variation in direct and indirect measures of energy status was examined using estimates of glycogen, lipid and protein levels in a single cohort of male three-spined sticklebacks from an annual population collected each month over one complete year. Condition factor, somatic condition factor and hepatosomatic index (HSI) were calculated as indirect indices of energy status and the accuracy of these indirect measures as predictors of energy status was investigated. Results indicate that both condition factors were significant predictors of energy reserves (lipid, protein, glycogen and total energy), but that the proportion of variance accounted for was small. Both condition factors perform better as predictors of energy content per unit body weight. The HSI was a significant, but a weak predictor of total glycogen levels over the whole year. On a seasonal basis the relationship between HSI and energy reserves was highly variable. These indices are therefore poor predictors of energy reserves in male three-spined sticklebacks.     

The feeding and energetics of stream-resident trout in winter*

The feeding of brook and brown trout in a temperate Canadian stream was studied over two winters. Stomach analyses indicate that feeding continues throughout the winter. Both species had similar diets. Despite continuous feeding, condition factors declined significantly in early winter and remained low until the spring, suggesting an early-winter depletion of lipid reserves. Energy (calorific) intake was similar in winter to that found in late summer or spring and therefore was not the cause of low winter condition. The derived energy from feeding was insufficient to offset the costs of maintenance metabolism in the early winter period, for both immature and recently spawned fish. It is suggested that stream-dwelling trout suffer a metabolic deficit during acclimation to rapidly declining water temperatures in November/December. Such a phenomenon has important physiological implications for overwinter survival.     

Short and chubby or long and slim? Food intake,growth and body condition in free‐ranging pythons

Abstract An animal's sex and body size can influence not only its rate of food consumption, but also the way in which it allocates the resultant energy among the competing demands of maintenance, growth, reproduction and storage. A 13‐year mark–recapture study of pythons (Liasis fuscus) in tropical Australia provides extensive data on these topics. Rates of food intake and growth were highest in small pythons, and decreased more rapidly with body size in males than in females. Allocation to storage (as measured by the snake's mass relative to its body length) showed a more complex pattern. Body condition was high at hatching, but dropped rapidly as energy was allocated to growth rather than storage. Condition then increased through juvenile life, was at a maximum close to maturation, and was higher in females than in conspecific males. Body condition thereafter decreased with increasing body length. These allocation ‘decisions’ reflect the relative advantages of growth versus energy storage at different body sizes. Hatchling snakes grow rapidly (and hence become thin) because greater body size enables the snake to ingest larger prey items. Adult females amass larger energy reserves than males, because they need reserves to produce the clutch. Large snakes become thinner because their feeding rates are low, and they cannot compensate with increased prey size because large‐bodied mammalian prey are rare in our study area.     

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.     

Do gravid females become selfish? Female allocation of energy during gestation

Net energy availability depends on plasma corticosterone concentrations, food availability, and their interaction. Limited net energy availability requires energy trade-offs between self-maintenance and reproduction. This is important in matrotrophic viviparous animals because they provide large amounts of energy for embryos, as well as self-maintenance, for the extended period of time during gestation. In addition, gravid females may transmit environmental information to the embryos in order to adjust offspring phenotype. We investigated effects of variation in maternal plasma corticosterone concentration and maternal food availability (2 × 2 factorial design) during gestation on offspring phenotype in a matrotrophic viviparous lizard (Pseudemoia entrecasteauxii). Subsequently, we tested preadaptation of offspring phenotype to their postnatal environment by measuring risk-averse behavior and growth rate using reciprocal transplant experiments. We found that maternal net energy availability affected postpartum maternal body condition, offspring snout-vent length, offspring mass, offspring performance ability, and offspring fat reserves. Females treated with corticosterone allocated large amounts of energy to their own body condition, and their embryos allocated more energy to energy reserves than somatic growth. Further, offspring from females in high plasma corticosterone concentration showed compensatory growth. These findings suggest that while females may be selfish when gestation conditions are stressful, the embryos may adjust their phenotype to cope with the postnatal environment.     

Thermal stress,thermal safety margins and acclimation capacity in tropical shallow waters—An experimental approach testing multiple end-points in two common fish

Tropical organisms are predicted to be among the most impacted by increasing sea surface temperatures, particularly those from intertidal habitats. In this study, a complete thermal biology assessment was conducted for two widespread tropical Atlantic shallow reef fish: Abudefduf saxatilis (damselfish) and Scartella cristata (blenny), which make extensive use of tide pools. The main objectives were to measure the time-course changes during one month in i) thermal and oxidative stress biomarkers (in gills, muscle and skin), ii) upper thermal limits, acclimation capacity and thermal safety margins and iii) body size, condition and energy reserves (total protein and lipid contents), under two temperature treatments (control – mean summer temperature, and elevated temperature − + 3 °C, as projected by climate warming scenarios for the end of this century).Results from biomarker analyses suggest that under increased temperature, both species displayed a typical response of physiological stress characterized by the activation of molecular chaperones and antioxidant protection. Both species presented a significant acclimation potential in the long term, as shown by increased critical thermal maxima values at higher temperature. However, these species may already be at risk during summer heat waves, as thermal safety margins for both species were low. Additionally, despite acclimation, some energetic tradeoffs may exist, since specimens from both species showed smaller body sizes at higher temperature (even though maintaining body condition). Finally, temperature treatments had a significant influence not only in the total amount of energy reserves (lipid contents) but also in their rate of deposition or depletion (total proteins and lipid contents). This is the first multi-end-point holistic approach to assess the impact of warming in shallow tropical water fish and it highlights the high risk that intertidal organisms are facing in both present and future sea surface temperature conditions.     

Determining seabird body condition using nonlethal measures

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 (R2 < 0.2) 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%.     

The annual pyloric cycle of Asterias rubens L. (echinodermata: Asteroidea)

Observations of biochemical changes in the pyloric caeca of Asterias rubens L. in different nutritional states and at different stages of the reproductive cycle show the existence of an annual cycle in the pyloric caeca. This pyloric cycle is closely related to that of the gonads. During the gonadal rest (May to September, when the gonads are barely visible) the pyloric caeca grow and store glycogen. During gonadal activity (October to April), they progressively autolyse and first accumulate, then liberate, lipid reserves. This winter pyloric autolysis provides the gonads with the structural material necessary for their annual growth. The summer pyloric growth must be interpreted as the accumulation of the structural material. Under normal conditions, during gonadal rest, the pyloric glycogen reserves ensure the survival of the animal. Pyloric lipid reserves appear only at the beginning of gonadal activity. Their synthesis is intimately related to the start of the gonadal activity of which they are the principal energy source. We believe that the pyloric cycle is under control of hormones that could be transported through the haemal system.The pyloric caeca are the only reserve organs of starfish. Neither the other digestive organs nor the body wall accumulate energy reserves. During starvation of medium duration (2 to 4 weeks) the pyloric caeca suffice to ensure the survival of the animal by liberating their reserve material and simultaneously starting to autolyse. During prolonged starvation, general autolysis occurs and the size of the starfish diminishes.     

Lipid dynamics in the viperine snake, Natrix maura, from the Ebro Delta (NE Spain)

Body reserves play a major role in several aspects of vertebrate biology. Accurate identification and quantification of body reserves constitute a useful contribution to the better understanding of the energetic costs of reproduction and the implication of food availability in life history traits of organisms. In this study, lipid content in fat bodies, liver and muscle of the viperine snake (Natrix maura) were measured along an active season. Samples were collected monthly from a natural population of the Ebro Delta Natural Park (NE Spain). This methodology pointed out that lipids stored in fat bodies were the main energetic source during reproduction. In addition, lipids stored in liver appeared to be critical for vitellogenesis, while lipids stored in muscle turned out to be a supplementary energetic resource to fuel reproductive effort. Relationships between changes in body reserves and prey availability in canals of the Ebro Delta were also considered. In males, lipid reserves presented a positive correlation with food availability. On the contrary, lipid reserves in female viperine snakes decreased during vitellogenesis even though food availability increased in this period, which suggests a quick transfer of body lipids to clutch. In April, when rice fields of the Ebro Delta were dry and aquatic prey was scarce for viperine snakes, males and females presented a lower lipid content in fat bodies, liver and muscle than they did in other months, showing a clear link between prey availability and body reserves during food shortage. Thus, patterns of variation in fat levels indicated that Natrix maura is a capital breeder since it acquires resources in advance and stores them until they are invested during the reproductive period. Nevertheless, the shortage in April forces Natrix maura to turn into a facultative income breeder to fuel vitellogenesis. Finally, fat reserves in body components were compared with an estimate of body condition calculated by the residuals from the regression of body mass on body length. In male viperine snakes, the estimate of body condition was correlated with fat levels, revealing that this index is useful to measure condition in living individuals. On the contrary, body condition in females was not correlated with fat levels, which suggests that it is not appropriate to infer condition in female viperine snakes since it depends on the reproductive status of the individuals.     

Winter lipid depletion of juvenile walleye pollock Theragra chalcogramma in the Doto area, northern Japan

Seasonal variation in body size and nutritional condition of juvenile walleye pollock Theragra chalcogramma was examined to elucidate the mechanism underlying their first-winter survival on the continental shelf of the Doto area, northern Japan, based on monthly samples collected over 2 years. Stored lipid mass was highest during autumn, but 93% (2004) and 80% (2005) of lipids were exhausted by the onset of winter. Lipid levels in the winter of 2004 remained low (7–14% of the autumnal maximum), and there was reduced growth rate until the spring, whereas in 2005 lipid levels were higher and more variable (10–46% of the maximum) and some growth occurred. An analysis of the allometric relationships between body size and stored energy showed that larger individuals accumulated disproportionately more energy in the autumn, but the advantage disappeared prior to the winter. In January 2004, stored lipid energy was low throughout the Doto continental shelf relative to the continental slope area. These results suggest that winter feeding opportunities on the shelf are severely limited but not completely absent. Previous studies have shown that winter temperatures on the shelf are lower than those in the slope area. It is possible that juvenile T. chalcogramma survive winter on the shelf without a high level of pre-winter lipid storage because the occasional feeding in the cold shelf water benefits energy conservation.     

Temporal changes in body mass, body composition and metabolism of gibel carp Carassius auratus gibelio during food deprivation

To investigate the response to starvation, gibel carp Carassius auratus gibelio [12.5 ± 0.03 g (mean ± SE, n = 24)] were deprived of food at 25.8 ± 0.2°C (mean ± SE, n = 56) for 56 days. Body mass, proximate composition in whole body and muscle, and respiration were measured at 7‐day intervals. Body mass decreased with prolongation of deprivation, with a significant decline recorded after 7 days deprivation. Fish lost 22% of their fresh mass and 34% of dry mass after 56 days. Fish lost 38% of the body lipid over the first 7 days, and lost body lipid at a rate of 0–11% per week over the remaining 49 days. Body protein was lost at 1–5% per week throughout deprivation. Compared with the initial composition, body lipid concentration was lower and ash concentration higher on day 7. Water as a percentage of body mass was higher after 28 days, and protein concentration lower after 42 days, than at the start of deprivation. Muscle lipid and protein concentration was lower, and % water higher, after 7 days than at the start of deprivation, whereas muscle ash concentration was relatively constant during deprivation. After 56 days, fish lost body water by 18%, body lipid by 84%, body protein by 30%, and body energy by 45%. Oxygen consumption rate dropped from day 1 to day 3, increased from day 4 to day 14, gradually decreased from day 15 to day 35, and maintained a relatively constant level from day 36 to day 56. Results of the present experiment reveal that gibel carp utilize body lipid as a major energy source in the first 7 days of food deprivation, then turn to body protein as an energy fuel when lipid reserves are heavily depleted. Oxygen consumption is maintained at a relatively low and constant level when most lipid reserves are exhausted.