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.     

Carcass composition of spur-winged geese with comments on reproduction and moulting

Spur-winged geese at Barberspan, an agricultural area in South Africa, had an average carcass composition of 64% water, 22% protein, 6% lipid, 6% minerals and 2% carbohydrate with an energy content of 21.6 kJ g^-1 DW. There were significant changes during the year in protein and lipid levels; they were lowest during the annual flightless wing moult. Protein and lipid levels also declined during breeding but this event had less effect on body reserves than moulting. The relationships between various body measurements and mineral content of the carcass (as an estimate of skeletal weight) were examined.     

Sexual dimorphism in the energetics of reproduction and growth of North Sea plaice, Pleuronectes platessa L.

The chemical composition and energy content of North Sea plaice during the spawning period were examined in mature males and females and in immature fish, to study differences in the allocation of energy over reproduction and somatic growth between the sexes. At the beginning of the spawning period mature males and females had equal dry weights of lipid that were 70% higher than in immatures. Protein content in mature males was equal to that in immatures but was 23 % higher in mature females. Immature males and females did not differ in chemical composition. At the end of the spawning period, spent and immature fish had equal lipid contents, but protein content in spent females was 10% lower than in spent males, and 17% lower than in immatures. Gross energy content of the body decreased by 44% (65·2 to 36·3 J cm^-3) in mature females, 27% (55·0 to 40·OJ cm^-3) in mature males, and 9% (48·7 to 44·2J cm^-3) in immatures. Energy content of plaice eggs was estimated at 6·60 kJ per 1000 eggs. Reproductive investment was estimated from the energy loss during the spawning period and included the energy of sex products and spawning metabolism. Somatic growth comprised the annual increase in energy content of fish. The pattern of energy allocation over reproduction and somatic growth differed between males and females. Males started their reproduction at a smaller length and a younger age and allocated a higher proportion of the available energy into reproduction than females. Available energy resources for somatic growth and reproduction (surplus production) were equal between the sexes up to a length of about 30 cm. Beyond this length male surplus production levelled off whereas female surplus production continued to increase. The differences in surplus production and the allocation patterns are discussed. For female plaice the energy allocated into egg production was estimated as between 48 and 64% of the total amount of energy lost during spawning. The remaining energy is used for metabolism during the spawning period, yielding an estimate of the metabolic rate of mature females of between 6·4 and 9·1 kJ day^-1. A maximum estimate of the metabolic rate of mature males was 7·4 kJ day^-1.     

Estimation of carcass fat and protein in northern pintails (Anas acuta) during spring migration

Foraging in stopover areas influences nutritional condition of birds during spring migration. Our purpose was to determine if body mass, percent carcass water, and serum biochemistry would predict energy reserves (carcass fat and protein) in northern pintails (Anas acuta) at a spring staging area, Lake St. Pierre in Québec, Canada (46 degrees 11 'N, 73 degrees 08 'W). Northern pintails were collected during spring 1997 (14 April-9 May). In this staging area, body mass and percent body water successfully estimated carcass protein and fat in male northern pintails, but only carcass protein in females. None of the seven blood parameters we used accurately estimated nutritional reserves in staging northern pintails. These findings suggest that investigators must use direct estimates of carcass reserves to examine nutrient reserve requirements for egg production, migration, or body maintenance during spring migration.     

Body condition changes of sand martins (Riparia riparia) during breeding, and a comparison with fledgling condition

The body condition of adult sand martins during the breeding season and a sample of fledglings was investigated by carcass analysis and by examination of live birds in the field. Fat scoring of live individuals was a reliable predictor of an individual's lipid content in most instances. Body mass changes during breeding were related to changes in the size of birds' lipid reserves, pectoral muscle mass, body water, and to development of the reproductive system. In both males and females, reserve lipid declined between the onset of breeding and nestling rearing. Pre-breeding males had significantly greater pectoral muscle masses than did nestling rearing females. The potential significance of protein in pectoral muscles as an energy reserve during breeding was small compared with lipid reserves. By comparing potential energy reserves in body lipid and protein with daily energy expenditure, it was calculated that neither incubating nor nestling rearing adults could survive a day of normal activity without feeding. The selective premium on adults optimizing their use of time and energy for self-maintenance behaviour during breeding is therefore likely to be great.     

Weight change and the body reserves of female blue tits, Parus caeruleus, during the breeding season

A female blue tit lays a large clutch that is commonly heavier than her own body weight. Body reserves cannot provide for the entire clutch and little is known of the importance of these reserves during the breeding season. This study reports on weight changes in adult female blue tits during the breeding season, observed by the use of electronic balances. In addition, data from dissecting 67 adult female blue tits, at different stages of the breeding cycle, were used to investigate the changes in body weight. There was a rapid increase in body weight during a period of 4–5 days before the first egg was laid, which was explained mostly by the development of the oviduct and the production of an egg on the final day. Body weight remained constant during the laying period, before decreasing by approximately one gram after clutch completion, owing to the absence of an egg and the absorption of the oviduct. Body reserves increased during the day after clutch completion, were maintained throughout the incubation period and were mobilized during the first five days of the nestling period. Breeding female blue tits appear to deposit small body reserves to act as a short-term buffer against adverse conditions or to support the brooding phase.     

Utilization of pre-existing energy stores of female Aedes aegypti mosquitoes during the first gonotrophic cycle

Pre-existing energy reserves may play an important role in regulating the utilization of blood meal proteins in female anautogenous mosquitoes. Determining the fate of reserves derived from the sugar meal and larval food during the first gonotrophic cycle would help to elucidate the relative contributions of larval and adult nutrition to survival and reproduction. We measured the allocation of pre-blood-meal reserves to egg production or energy production during the first gonotrophic cycle by using [14C]-labeled female Aedes aegypti mosquitoes. Feeding adults [3,4-14C]-glucose labeled the glycogen and sugar stores (approximately 50%), lipid stores (approximately 25%), and protein and amino acid stores (approximately 25%). During the first gonotrophic cycle, about 60% of the glycogen and sugar stores were metabolized and all were used for energy production. About 33% of the labeled protein and 72% of the labeled amino acid stores were metabolized, with about 9% being transferred to the eggs and the rest oxidized. About 30% of the lipid was metabolized, with about 65% being transferred to the eggs and the rest oxidized. Feeding [1-14C]-oleic acid to larvae effectively labeled adult lipid stores with about 75% of the label in lipid stores and 16% in proteins and 6% in glycogen. During the first gonotrophic cycle, about 35% of the labeled lipid stores were metabolized, with equal amounts being oxidized and transferred to the eggs. None of the other maternal stores labeled by fatty acid were metabolized during the first gonotrophic cycle. These results show that carbohydrate reserves are a critical source of energy during the first gonotrophic cycle, while lipid reserves are used equally for energy production and provisioning the eggs.     

Temporal changes in fat and protein levels in the tropical Anabantid Trichogaster pectoralis (Regan)

Samples of a wild population of the Anabantid Trichogaster pectoralis (Regan) were taken during and after one of the two main breeding periods of the year. Carcasses of both sexes and female gonads were analysed for fat, protein and carbohydrate. Carbohydrates were present in insignificant amounts in all tissues. The fat content of the ovary proved to be high (over 30% of wet weight) and, as a result, ovarian energy in a ripe female formed a substantial proportion of the total energy content of the fish. However, it did appear possible for females to develop large ovaries without depleting the carcass fat stores.
Although protein levels in the carcass were significantly higher in 1 month, relative to fat levels, protein levels remained fairly constant during the period of study. Conversely fat levels showed marked fluctuations throughout the breeding period, being high at the start of the breeding period, low towards the end, finally rising again 1 month after most fish had terminated breeding. The range of fat levels appeared to be independent of the gonadal state of the fish at capture and seemed more closely related to month of capture. Two explanations are suggested; (1) changes in food availability over the period in question and/or (2) if repeated spawnings occur this could deplete fat reserves in the carcass.
In this species fat is stored largely in the muscle but some is stored around the coiled intestine. A visual scoring system was used to assess the extent of this fat deposition and it was found that the fat score was correlated with the level of fat in the carcass. Thus the level of fat around the intestine, although insignificant relative to carcass fat, could be used as a method of assessing the fatness of a fish.     

Adipokinetic hormone inhibits the formation of energy stores and egg production in the cricket Gryllus bimaculatus

Adipokinetic hormones (AKHs) mobilise lipids, carbohydrates and/or proline from insect fat body stores. In addition, AKHs inhibit lipid and protein synthesis in the fat body. In the current study, 100 pmol homologous Grybi-AKH was injected twice daily into adult female crickets, Gryllus bimaculatus, starting on the day of adult emergence. The effects of the injected AKH on the formation of energy reserves in the fat body and on egg production were measured on day 4 after adult emergence. In comparison to water-injected control animals, lipid and protein content in the fat body of the AKH-injected crickets was significantly reduced, suggesting an inhibitory effect of AKH on the formation of lipid reserves and protein stores. The content of glycogen and free carbohydrate in the fat body was significantly higher in the AKH-injected animals. The most pronounced effect of the AKH-injections was a significant reduction of ovary mass, due to the retarded maturation of the oocytes and the significantly lower number of terminal oocytes produced. It is concluded that AKH inhibits egg production indirectly by interference with the formation of energy stores in the fat body that are mobilised to fuel egg production.     

High Dietary Calcium Reduces Body Fat Content,Digestibility of Fat,and Serum Vitamin D in Rats

Objective: This study investigated which aspect of energy balance was responsible for the decrease in body fat content of rats fed a high‐calcium, high—dairy protein diet. Research Methods and Procedures: Male Wistar rats were fed a control diet (25% kcal fat, 14% kcal protein from casein, 0.4% by weight calcium) or high‐calcium diet (25% kcal fat, 7% kcal protein from nonfat dry milk, 7% kcal protein from casein, 2.4% calcium) for 85 days. Body weights, digestible energy intakes, energy expenditures, rectal temperatures, body composition, and serum glucose, insulin, free fatty acids, triglycerides, and 1, 25‐dihydroxyvitamin D were measured. Results: Rats fed high‐calcium diet gained significantly less weight than controls and had 29% less carcass fat. Gross energy intake was not significantly different between groups, but digestible energy was 90% of gross energy in the high‐calcium diet compared with 94% in the control diet because of increased fecal excretion of dietary lipid. The difference in digestible energy intake accounted for differences in carcass energy. Body temperatures and energy expenditures of the rats were not different. The high‐calcium diet reduced serum triglycerides by 23% and serum 1, 25‐dihydroxyvitamin D by 86%. Discussion: These results confirm that a high‐calcium diet decreases body weight and fat content due to a lower digestible energy intake caused by increased fecal lipid and a nonsignificant reduction in gross energy intake.     

Seasonal changes in serum lipid composition of the plaice, Pleuronectes platessa L.

Seasonal changes in the major lipid classes of plaice serum were examined in fish caught off the north-east coast of Scotland. Both male and female plaice were sampled monthly between June 1983 and May 1984. Total lipid, total cholesterol, free cholesterol, triglycerides, phospholipids and non-esterified fatty acids were measured using commercial kits validated against standard reference methods. There were no significant differences in lipid levels between male and female fish and, although there were real differences between months for each lipid class, there was no regular pattern over the 12-month period. All the lipid class values, however, reached a maximum in September, which could be related to the increase in feeding after spawning. The smaller peak in February could be associated with the mobilization of lipid reserves for gonadal development and to overcome the effect of starvation during spawning.     

Ovarian maturity,egg development,and offspring generation of the deep-water shrimp Plesionika edwardsii (Decapoda,Pandalidae) from three isolated populations in the eastern North Atlantic

Ovarian maturity, egg development and brood size were analysed for three isolated populations (Madeira, the Canary Islands and Cape Verde Islands) of Plesionika edwardsii (Decapoda, Pandalidae) in the eastern Atlantic. Multiple colour patterns were observed at the same ovarian maturity stage, which was verified histologically, invalidating the extensive use of ovarian colour as a maturity stage criterion. The physiological size at sexual maturity, based on the maturity of the ovaries, was higher in Madeira (carapace length of 19.73?mm) and decreased to the Cape Verdes (16.39?mm). Synchronic ovarian maturation was observed during the embryo incubation process, and ovigerous females bearing embryos at the final stage of development were found throughout the year. Females are multiple spawners during the reproductive season, after which the reproductive process ends and a rest period begins. The absence of females larger than the size at sexual maturity with ovaries in Stage 1, the incubation of embryos in the final developmental stages, suggests that the resting period begins with a process that reabsorbs the energy located in the ovaries and that the resting period occurs asynchronously in females in each of the studied populations. Embryo size was independent of female body size in the three populations studied, but increased with the developmental stage. A power equation was used to describe the relationship between brood size and female body size in the three areas studied. The mean number of external Stage I embryos carried by females decreased from Madeira (n?=?7868) to the Cape Verdes (n?=?3781), where less energy, in terms of the number of embryos and the size of the eggs, was invested in reproduction. Although female size decreases from north to south, the egg number was higher in Madeira than in the Cape Verdes for the same size range.     

Exploring the origin of egg protein in an oviparous water snake (Natrix maura)

Reptiles are typical capital breeders that fuel reproduction by the use of lipids stored in fat bodies. However, little is known about the origin (exogenous or endogenous) of egg protein. We have examined the origin of egg protein by means of the analysis of protein content of liver and carcass (skeletal muscle) of the oviparous snake Natrix maura throughout an annual cycle. We have also measured monthly variation of the digestive-content mass and the ovarian mass. Results showed that protein in liver peaked during vitellogenesis according to the role of the liver in the synthesis of vitellogenin. Partial correlations showed the path of protein from the prey (digestive-content) to the liver, and finally to the ovaries, as well as an inverse relation between carcass protein and ovarian mass. Carcass muscle is the only body part that may act as a potential reserve for endogenous protein, although we did not find significant variation in carcass protein during female reproduction. As females with large follicles did not stop foraging activity, we assumed that egg protein was derived from the diet as partial correlations indicated. Our results suggest that N. maura is a capital breeder for lipids and tend to be income breeder for protein. This conclusion contrasts with that observed in capital breeders for which egg protein was derived from muscle. We discuss the idea that flexibility in the origin of egg protein could affect the body condition in post-reproductive females.     

Sexual maturation in female triploid plaice, Pleuronectes platessa, and plaice × flounder, Platichthys flesus, hybrids

The gonads of immature female triploid plaice ( Pleuronectes plafcssa ) and plaice × flounder ( Platichthys flesus ) hybrids produced from cold-shocked eggs were examined. They were less than half the size of those in diploids of similar age and contained only small numbers of developing oocytes. These were similar both in size and cytological appearance to those in the controls. The bulk of the ovarian tissue was composed of nests of small undifferentiated cells resembling the oogonia in the ovaries of diploid fish. Mature, 5 year old triploid hybrids which had not produced eggs over the previous two spawning seasons were killed in January when the diploid controls were nearing spawning condition. The ovaries of these fish were smaller than those of the controls and contained degenerating oocytes at maturation stage V. The advantage of sterile female triploid fish for fish-cultivation is discussed.     

The effect of female body condition on egg laying in Lesser black-backed gulls Larus fuscus

Female Lesser black-backed gulls, culled from a moorland nesting site in Lancashire for public health reasons, were examined to compare their body condition with the number and quality of eggs that they would lay. The clutch size and stage of the laying cycle of each bird were determined by examining their ovaries. Fat condition was obtained by extraction of the whole carcass with chloroform and an index of protein condition was obtained from the lean dry weight of the flight muscles. Oviduct eggs were analysed for the amounts of fat and protein present in the yolk and albumen.
Lesser black-backed gulls are not fully indeterminate layers. Most females develop only enough enlarged ova to produce the normal clutch of three eggs, with some birds able to lay an additional one or two replacement eggs if necessary. The female's protein reserve, but not her fat reserve, declined during egg formation and was correlated with the total number of eggs she could potentially have laid. Similarly egg quality, as measured by yolk weight and protein content, was correlated with female condition but not her fat reserves. Albumen weight was not correlated with body condition.
It is shown that egg production is a demanding process in Lesser black-backed gulls and breeding success is influenced by the amount of the female's protein reserve both through its effect on egg quality and on her ability to lay replacement eggs if necessary.     

Relative contribution of lipid sources to eggs of lesser scaup

Studies of how birds mobilize nutrients to eggs have traditionally considered a continuum of possible allocation strategies ranging from income breeding (rely on food sources found on the breeding grounds) to capital breeding (rely on body reserves stored prior to the breeding season). For capital breeding, stored body reserves can be acquired either on or away from the breeding grounds, but it has been difficult to quantify the relative contribution of each, precluding identification of key habitats for acquiring nutrients for clutch formation. During 2006–2009, we explored the importance of spring‐staging habitats versus breeding‐area habitats for egg‐lipid formation in female lesser scaup Aythya affinis using stable carbon (δ¹³C) isotope analyses. Although δ¹³C values for abdominal lipid reserves brought to the breeding grounds overlapped those of local amphipods, we were able to quantify the importance of local plant carbohydrates (seeds of emergent wetland plants) to the production of eggs. We compared the importance of local wetland seeds (overall δ¹³C: ?29.1 ± 0.9‰ SD) to combined lipid stores and lipids from local amphipods (overall δ¹³C: ?23.8 ± 2.2‰). Local seeds and stored body lipids contributed equally to egg lipid formation across years but we found evidence of annual variation in their relative importance. Wetland seeds contributed 39% (SE = 10%) to egg lipid production, and the importance of this source varied by year (90% CI = 47–75% in 2006, 13–42% in 2007, 29–65% in 2008, and 7–30% in 2009). In contrast to earlier studies that suggest lesser scaup predominantly employ a capital breeding strategy, our results suggest that in some years females may attain half of their energy for clutch formation from foods on the breeding grounds.     

Reproductive effort and reproductive nutrition of female desert tortoises: essential field methods

I used three innovative, nondestructive field methods (gas dilution,doubly labeled water and radiography) to measure individualenergy and water budgets of wild, female desert tortoises (Gopherusagassizii). With these budgets, I evaluated whether body reserveshelp females produce eggs independent of rainfall and food availability.Female desert tortoises used large seasonal and annual changesin metabolism and body water, protein and energy reserves tosurvive and produce eggs. Although lipid reserves are importantto female desert tortoises, nitrogen or crude protein appearsto be the primary limiting resource for producing eggs. By reducingmetabolic rates 90%, females conserved enough body reservesto produce eggs during extreme drought conditions; this is aneffective bet-hedging reproductive pattern in an extreme andunpredictable environment.     

Fuel mobilization for energy metabolism in the silkmoth, Bombyx mori, during reproductive processes

Flight muscles of male moth, B. mori seem to utilize carbohydrate preferentially as a source of energy for all its acrobatic movements during the search for female moth. Depletion of triacylglycerol from flight muscles without affecting its level from fat body suggests that this lipid fraction serves as a source of energy in flight muscles during insemination processes. Significant depletion of triacylglycerol and glycogen from flight muscles of female moth after egg laying indicates that they are used to meet the energy requirement of female during oviposition activity. Depletion of proteins from flight muscles of male and female insects suggest that these proteins are transported to the accessory reproductive glands to meet their protein demand.     

Temperature-dependent development rates of plaice (Pleuronectes platessa L.) eggs from the Irish Sea

Experimentally derived temperature-dependent development ratesfor planktonic fish eggs are required for developing IndividualBased Models (IBMs) of early life history stages and for estimatingdaily egg production in ichthyoplankton surveys. Developmentrates of Irish Sea plaice eggs were experimentally determinedfor temperatures between 4.5 and 12° C. Differences in developmentrates were apparent when these results were compared with literaturedata for eggs from North Sea parents. On average, plaice eggsfrom the Irish Sea developed more rapidly and hatched up to2 days earlier compared with North Sea eggs incubated at thesame temperature. In terms of parental effects, larger femalestended to produce larger eggs but egg size decreased over successivebatches from the same female within a year. Temperature-dependentegg development rates were influenced by egg size in a non-linearmanner for eggs incubated at 6° C, but egg size had no effectat higher temperatures. Since most plaice eggs sampled fromthe Irish Sea during ichthyoplankton surveys in 1995 and 2000were found in waters cooler than 8° C, such effects maymodify egg development rates in the wild.     

Biochemical changes occurring during final maturation of eggs of some marine and freshwater teleosts

Biochemical changes which occur during post-vitellogenic meiotic maturation of oocytes were investigated by measuring the concentrations of various phosphorus-containing fractions of the ovaries and mature eggs of four marine species with pelagic eggs, plaice, Pleuronecles platessa , cod, Gadus morhua , haddock, Melanogrammus aeglefinus , and whiting, Merlangius merlangus , and three freshwater species with demersal eggs, rainbow trout, Salmo gairdneri , pike, Esox lucius , and powan, Coregonus lavarelus . In the former group, protein phosphate was deposited at normal vertebrate levels in the ovaries during vitellogenesis but was exhausted during maturation, its decrease correlating with the uptake of water which rendered the mature egg buoyant in sea water. In plaice, phospholipid phosphate declined slightly during maturation, while inorganic phosphate increased by an amount slightly less than that which disappeared from the protein fraction. No such changes occurred on maturation in these three fractions in the freshwater species, in which little or no water uptake occurred. In plaice, the approximately five-fold increase in the water content of the oocyte was accompanied by a corresponding five-fold increase in the amount of potassium, the major inorganic cation of the oocyte. These pelagic eggs appear to be unique among the eggs of vertebrates in their high water content ( c . 92%) and in the fact that the protein phosphate is almost entirely utilized before fertilization, while, in those freshwater fishes and other oviparous vertebrates that have been examined, the egg is of much lower water content (50–70%) and most of the protein phosphate, like the other major yolk constituents, is used during embryonic growth. This utilization at such an early stage in the life cycle constitutes an extreme example of heterochrony.