Metabolic characteristics of midwater zooplankton: Ammonia excretion,O:N rations,and the effect of starvation

The nature of protein catabolism in a wide range of species of midwater zooplankton was investigated. The weight-specific ammonia excretion rates (g NH₃–N g^–1 dry wt h^–1, y) decline exponentially with minimum depth of occurreece (MDO, x), y=163.4 x^{–0.479±0.212} (95%ci) (CI=confidence interval), when temperature is held constant. The change in ammonia excretion can be partially explained by the decrease in percent protein (%P) with MDO, %P=80.17 MDO^{–0.148±0.122 (95%ci)} The atomic O:N ratio of freshly caught zooplankters ranged from 9.1 to 91, with most measurements between 9 and 25. Detailed studies were carried out on the response of one of the species studied (Gnathophausia ingens) to starvation (28 d). After 14 d of starvation the average ammonia excretion rate declined by more than 75% to less than 1 g NH₃–N g^–1 wet wt h^–1, although the average oxygen consumption declined by only 13% within the first 7 d of starvation and then remained stable. This differential response of oxygen consumption and ammonia excretion to starvation resulted in an increase in the average O:N ratio of starved animals from an initial 33 to 165 after 21 d. The average O:N ratios of fed mysids remained below 38 during the experiment. G. ingens maintains a relatively uniform metabolic rate during starvation by relying more heavily on its large lipid stores than when being fed.     

The effect of laboratory conditions on the extrapolation of experimental measurements to the ecology of marine zooplankton. IV. Changes in respiration and excretion rates of boreal zooplankton species maintained under fed and starved conditions

Herbivorous zooplankton species (Calanus plumchrus, Paracalanus parvus and Euphausia pacifica) and carnivorous species (Parathemisto pacifica and Pleurobrachia pileus) collected from Saanich Inlet, British Columbia, Canada, were maintained in the laboratory under fed and starved conditions. Respiration rate and excretion rates of ammonia and inorganic phosphate were measured successively on the same batch populations of each species in different feeding conditions. Respiration rate remained at a constant level or increased during the feeding experiment but decreased progressively in starved individuals. Herbivorous, but not carnivorous, species showed a rapid decrease in both excretion rates for the first few days of an experiment irrespective of feeding conditions. However, the general level of excretion rates of fed specimens was higher than that of starved ones. The O:N, N:P and O:P ratios were calculated from respiration, ammonia excretion and phosphate excretion and discussed in relation to metabolic substrates of animals during the experiment. A marked difference was shown in the O:N ratio between fed hervivores (>16) and fed carnivores (7 to 19), suggesting highly protein-oriented metabolism in the latter. One unknown factor causing variation in excretion rates is speculated to be the physiological stress on animals during sampling from the field. It is suggested that the laboratory measurement of realistic excretion rates of zooplankton is difficult owing to their large fluctuations, but this is not the case with respiration rate.     

Effect of heterozygosity on metabolism during starvation in the American oysterCrassostrea virginica

Crassostrea virginica were collected in winter from the Nissequogue estuary, Long Island, New York, USA and starved for 42 d at 20°C in 28‰ salinity. Oxygen consumption (VO₂) and ammonia excretion (VNH₄), measured at about 20-d intervals, and changes in dry weight and carbohydrate content were tested for relationship with heterozygosity at six enzyme loci. Rate of decline in tissue dry weight during starvation was size-dependent, small oysters losing weight faster than large individuals. There was a significant effect of heterozygosity, the rate of weight loss decreasing with increasing heterozygosity. No significant correlation was found between heterozygosity and carbohydrate depletion rate, VO₂ or VNH₄. Carbohydrate content was positively correlated with size before and after starvation. Carbohydrate depletion contributed less to weight loss in small than in large oysters. This suggests that more lipid and protein was catabolised in small oysters. VO₂ and VNH₄ declined during the course of the experiment. There was good agreement between energy loss estimated from decline in dry weight and energy loss estimated from physiological measurements. We conclude that heterozygotes conserve weight more efficiently under temperature and nutritive stress and that this may be expected to enhance viability and growth.     

Experimental evaluation of the energy balance in Octopus vulgaris, fed ad libitum on a high-lipid diet

A complete energy balance equation was estimated for the common octopus Octopus vulgaris at a constant temperature of 20°C, fed ad libitum on anchovy fillet (Engraulis encrasicolus). Energy used for growth and respiration or lost with faeces and excreted ammonia was estimated, along with total energy consumption through food, for six specimens of O. vulgaris (with masses between 114 and 662 g). The energy balance equation was estimated for the specimens at 10-day intervals. During each 10-day interval, food consumed, body mass increase and quantity of faeces voided were measured. The calorific values of octopus flesh, anchovy flesh and faeces were measured by bomb calorimetry. Oxygen consumption and ammonia excretion rates were monitored for each specimen during three 24-h experiments and daily oxygen consumption and ammonia excretion were estimated. It was found that 58% of the energy consumed was used for respiration. The amount of energy invested in somatic and gonadal growth represented 26% of the total energy budget. The energy discarded through faeces was 13% of consumed energy. The estimated assimilation efficiency (AE) values of O. vulgaris feeding on anchovy (80.9–90.7%) were lower than the AE values estimated for other cephalopod species with different diets of lower lipid content such as crabs or mussels. Specific growth rates (SGR) ranged 0.43–0.95 and were similar to those reported for other high-lipid diets (bogue, sardine) and lower than SGR values found for low-lipid, high-protein diets (squid, crab, natural diet). Ammonia excretion peak (6 h after feeding) followed the one of oxygen consumption (1 h after feeding). The values of atomic oxygen-to-nitrogen (O:N) ratio indicated a protein-dominated metabolism for O. vulgaris.     

Effect of starvation on the energy budget of two Asian horseshoe crab species: <Emphasis Type="Italic">Tachypleus tridentatus</Emphasis> and <Emphasis Type="Italic">Carcinoscorpius rotundicauda</Emphasis> (Chelicerata: Xiphosura)

Energy budget is one of the most studied parameters in aquatic animals under environmental challenge. To examine how prolonged starvation would affect their energy budget, respiration rate (RR), ammonia excretion rate (ER), oxygen consumption to ammonia–nitrogen excretion (O:N) ratio and scope for growth (SfG) representing the balance between energy intake and metabolic output, two Asian horseshoe crab species, Tachypleus tridentatus and Carcinoscorpius rotundicauda, were investigated in two feeding regimes (fed and starved) over a period of 7 weeks. No significant effects of species and time course, as well as their interaction, on absorption efficiency were observed in the fed treatments. For both species, RR and ER of the starved treatments significantly decreased, while their O:N ratio significantly increased during the experiment. However, such values for the fed treatments remained relatively stable over the study period. A rapid reduction in SfG was only apparent in the first week of the starved treatments for both species; thereafter, their SfG remained relatively constant. In the fed treatments, SfG of T. tridentatus was significantly lower than that of C. rotundicauda throughout the experiment. In general, C. rotundicauda showed a greater decrease in SfG under starvation than T. tridentatus, suggesting that they may have a more competitive life-history strategy for adjusting to poor nutritional conditions.     

Egg production and lipid content of Calanus glacialis in spring: indication of a food-dependent and food-independent reproductive mode

Female Calanus glacialis were collected in early May 1989 in the pack ice region of the western Barents Sea and were fed or starved over 11 wk. Both groups laid eggs continuously during this period, however, fed females laid up to six times more eggs. During the first 10 d after collection, both groups spawned at low rates. There-after, fed females strongly increased spawning rates and maintained high egg production levels over 11 wk, while the rates of starved females decreased. During starvation they lost 70% body carbon, 50% body nitrogen and 70% lipids. The wax ester portion decreased from 86 to ca. 60% of total lipids. Three phases of gonad development and lipid metabolism were distinguished: early gonad development; gonad maturation with a rapid decrease in lipids, especially wax esters; and spawning under fed and starved conditions, where in fed females food provided most of the energy, whereas in starved females the lipid content strongly decreased.     

Some aspects of nitrogen metabolism in Mytilus edulis: effects of aerial exposure

The effects of starvation and emersion on the ammonia effluxes of Mytilus edulis L. were studied. Both fed and starved groups showed similar patterns of efflux during re-immersion, indicating no compensation for starvation during anaerobic catabolism; this is taken as evidence that carbohydrate or lipid is the predominant energy source at the outset of anaerobiosis. Instead of an overshoot of ammonia excreted during the 4 h following re-immersion, all groups showed significantly reduced efflux rates independent of emersion duration. Such results suggest some conversion of ammonia during the recovery period, as no significant drop in haemolymph ammonia occurred at this time. The level of ammonia accumulation in haemolymph and mantle cavity fluid decreased exponentially with emersion duration, implying the stabilisation, at a low rate, of ammonia production after a relatively long period of emersion (16 to 24 h). Oxygen levels of the mantle cavity fluid dropped rapidly during emersion but never attained anoxia over 24 h of emersion, which may indicate some oxygen uptake by emersed mussels. It has been concluded that the behaviour and physiological responses shown by M. edulis to emersion appear to reflect a need for respiratory gas exchange more than a response to desiccation – presumably in the interest of energy-conservation. The reductions in ammonia efflux rate during emersion and, to a certain extent, immediately after re-immersion have also been considered here to be energy-saving strategies that illustrate the importance of ammonia in restoring normal levels of some amino acids following re-immersion. Received: 26 August 1997 / Accepted: 27 May 1999     

Metabolic properties of Northern krill, Meganyctiphanes norvegica, from different climatic zones. I. Respiration and excretion

Adaptive processes linked to overall metabolism were studied in terms of oxygen consumption and ammonia excretion in each of three self-contained krill populations along a climatic gradient. In the Danish Kattegat, krill were exposed to temperatures which ranged from 4°C to 16°C between seasons and a vertical temperature gradient of up to 10°C during summer. In the Scottish Clyde Sea, water temperatures varied less between seasons and the vertical temperature gradient in summer was only 3°C. Temperatures in the Ligurian Sea, off Nice, were relatively constant around 12-13°C throughout the year, with a thin surface layer (20-30 m) of warm water developing during summer. The trophic conditions were rich in the Kattegat and, particularly, in the Clyde, but comparatively poor in the Ligurian Sea. Oxygen consumption increased exponentially with increasing experimental temperature, which ranged from 4°C to 16°C. Overall respiration rates were between 19.9 and 89.9 µmol O₂ g^-1 dry wt h^-1. Krill from the Kattegat, the Clyde Sea, and the Ligurian Sea all exhibited approximately the same level of oxygen consumption (30-35 µmol O₂ g^-1 dry wt h^-1) when incubated at the ambient temperatures found in their respective environments (9°C, 5°C, and 12°C). This indicates that krill adjust their overall metabolic rates to the prevailing thermal conditions. The exception to this were the respiration rates of Ligurian krill from winter/spring, which were about twice as high as the rates from summer krill despite the fact that the thermal conditions were the same. This effect appears to result from enhanced somatic activity during a short period of increased food availability and reproduction. Accordingly, krill appears to be capable of adapting to both changing thermal and trophic conditions, especially when nutrition is a limiting factor in physiological processes.     

Metabolism of epipelagic tropical ctenophores

Measurements of respiration and excretion at 25°C were made for five species of ctenophores collected during five cruises to the Bahamas (1982–1984). The mean element-specific respiration and ammonium excretion rates of freshly collected specimens of all species ranged from 4 to 16% d^-1, the mean atomic O:N ratios were 10 to 16, and ammonium averaged 60 to 90% of the total dissolved nitrogen excreted. For adult ctenophores, the carbon content ranged from 0.6% carbon (as percent of dry weight) for Bolinopsis vitrea to 3.7% carbon for Beroë ovata. There was a marked increase in the organic content (% carbon of dry weight) of small Bolinopsis vitrea with tentacles compared to fully lobate adults. B. vitrea had increasingly higher metabolic rates when held at food concentrations up to 100 copepods 1^-1 (about 250 g C 1^-1). The overall range between starved and well-fed B. vitrea was about two times for respiration and a factor of three for ammonium excretion. B. vitrea decreased from well-fed to a starved metabolic rate in about a day after removal from food. The metabolic rate of Eurhamphaea vexilligera was not measurably affected by short-term starvation or feeding (maximum 25 copepods 1^-1). In feeding experiments, E. vexilligera of 20 to 56 mm length fed at rates equivalent to clearance rates of 250 to 1 800 ml h^-1.     

Oxygen consumption of pelagic juveniles and demersal adults of the deep-sea fish Sebastolobus altivelis,measured at depth

Oxygen consumption rates of the deep-sea fish Sebastolobus altivelis were measured in situ on pelagic juveniles at mesopelagic depths (608 m) and on demersal adults at bathyal depths (1 300 m) in the Santa Catalina Basin in March 1982. Two pelagic juveniles were individually collected, and respiration was measured continuously for approximately 2 d with a slurp gun respirometer manipulated from the submersible “Alvin”. Oxygen consumption rates of these juveniles were highly variable and were 1.5 to 1.8 times higher during the night than during the day. Gut contents of the juveniles were mainly euphausiids (Euphausia pacifica and Nematoscelis difficilis). Four demersal adults were collected by “Alvin” and individually placed in fish-trap respirometers on the bottom where respiration was measured continuously for approximately 1 d. Weight-specific O₂ consumption rates for adults decreased with increasing body weight and were consistent in magnitude throughout the incubation period. Population O₂ consumption for demersal S. altivelis (calculated from abundance, size-frequency distribution, and O₂ consumption regression equation) was 11.01 μl O₂ m^-2 h^-1, which is two orders of magnitude less than the O₂ consumption rate for the population of the most abundant epibenthic megafaunal species in the Santa Catalina Basin, the ophiuroid Ophiophthalmus normani. O. normani is a principal prey for adults of S. altivelis based on gutcontent analysis. Given the population O₂ consumption rate as an estimate of food energy demand, the demersal population of S. altivelis would assimilate only 0.007% of the standing crop of O. normani per day.     

Effects of feeding frequency and symbiosis with zooxanthellae on nitrogen metabolism and respiration of the coral Astrangia danae

Colonies of the temperate coral Astrangia danae occur naturally with and without zooxanthellae. Basal nitrogen excretion rates of nonsymbiotic colonies increased with increasing feeding frequency [average excretion rate was 635 ng-at N (mg-at tissue-N)^-1 h^-1]. Reduced excretion rates of symbiotic colonies were attributed to N uptake by the zooxanthellae. Nitrogen uptake rates of the zooxanthellae averaged 8 ng-at N (10⁶ cells)^-1 h^-1 in the dark and 21 ng-at N (10⁶ cells)^-1 h^-1 at 200 Ein m^-2 s^-1. At these rates the zooxanthellae could provide 54% of the daily basal N requirement of the coral if all of the recycled N was translocated. Basal respiration rates were 172 nmol O₂ cm^-2 h^-1 for starved colonies and 447 nmol O₂ cm^-2 h^-1 for colonies fed three times per week. There were no significant differences between respiration rates of symbiotic and nonsymbiotic colonies. N excretion and respiration rates of fed (symbiotic and nonsymbiotic) colonies increased greatly soon after feeding. N absorption efficiencies decreased with increasing feeding frequency. A N mass balance, constructed for hypothetical situations of nonsymbiotic and symbiotic (3×10⁶ zooxanthellae cm^-2) colonies, starved and fed 15 g-at N cm^-2wk^-1, showed that the presence of symbionts could double the N growth rate of feeding colonies, and reduce the turnover-time of starved ones, but could not provide all of the N requirements of starved colonies. Rates of secondary production, estimated from rates of photosynthesis and respiration were similar to those estimated for reef corals.     

Ammonia excretion by the benthic estuarine shrimp Crangon franciscorum (Crustacea: Crangonidae) in relation to diet

Ammonia excretion of individual Crangon franciscorum Stimpson was monitored in response to ingestion of single meals. The three experimental diets were tubificids, mysids and fish. Ammonia excretion was also monitored for individual shrimp which had been starved. The rate of ammonia excretion was higher for fed than for starved individuals in all cases. Ammonia excretion rates were higher for shrimp which were fed tubificids than those fed the other diets. The rate of excretion was influenced by both weight of the individual and the amount ingested of each diet. Ammonia excretion was influenced by dietary factors other than nitrogen content of the diet or the quantity ingested. The data suggest that field estimates of ammonia excretion based on the excretion rates of starved animals may be underestimates. The recent feeding history of an organism influences the rate of ammonia excretion as well as the relationship between the rate of excretion and weight.     

Influence of temperature changes on oxygen uptake and ammonia and phosphate excretion,in relation to body size and weight,in Oikopleura dioica (Appendicularia)

The relationship between the rates of oxygen consumption, ammonia and phosphate excretion of a pelagic tunicate, the larvacean Oikopleura dioica Fol, 1872 were assessed as a function of size, dry weight and ash-free dry weight at 15°, 20° and 24°C. O. dioica has higher respiration and excretion rates than copepods of similar weight, but the weight exponent of the allometric power function: Y=aX ^b is similar to that of other poikilotherms. Temperatures above 20°C have a depressing effect on respiration and ammonia excretion. 90% of the variance in metabolic rates is explainable by body mass and temperatures Q₁₀ values for oxygen consumption, ammonia and phosphate excretion, respectively, are 2.45, 1.86 and 1.75 between 15° and 20°C, and 3.75, 2.90 and 3.60 between 20° and 24°C. Metabolic quotients (O:N, O:P, N:P) indicate a protein-oriented diet. The results of this study suggest weak metabolic regulation in O. dioica, an energetic strategy which allows an immediate response to favourable changes in feeding conditions.     

Physiological and behavioural responses of the intertidal scavenging gastropod Nassarius festivus to salinity changes

Physiology (oxygen consumption and ammonia excretion) and behaviour (feeding and activity) of the intertidal gastropod Nassarius festivus (Powys) at five different salinities [15, 20, 25 (control), 30 and 35‰] were studied for 4 weeks. Oxygen consumption and ammonia excretion rates were reduced immediately after salinity was either elevated or reduced. Subsequently, both rates were increased while the O:N ratio was decreased at all salinities, including the control, in the first 2 weeks and then levelled off. Such changes were probably attributed to osmotic adjustment and reproductive activity. Activity and feeding were reduced at low salinity, particularly in the first week. Reproductive output, in terms of the total number of egg capsules and the mean number of eggs per capsule, was also lowered at reduced salinities. Nevertheless, individuals at all salinities are able to maintain a positive energy balance. Results are discussed with respect to the distribution of N. festivus in Hong Kong waters. Received: 14 April 1997 / Accepted: 9 May 1997     

Ammonia excretion of Artemia sp. (crustacea: Branchiopoda) under axenic conditions)

Ammonia excretion of the Utah strain of Artemia sp. grown in axenic culture media was followed throughout the different stages of larval development. The ammonia accumulated in the culture medium does not affect ammonia excretion rates under our laboratory conditions. Growth rate, density and starvation all affect ammonia excretion. Ammonia excretion is directly related to the numerical growth index and inversely related to Artemia sp. density. Starved adult Artemia sp. excrete considerably lower amounts of ammonia than brine shrimp grown in nutritive medium. In view of the difficulties often encountered in estimating the excretion rates of small crustaceans, our culture conditions would appear advantageous for further studies on the purine metabolism of Artemia sp.     

Respiration and nitrogen excretion of zooplankton. IV. The influence of starvation on the metabolism and the biochemical composition of some species

Changes in the respiration, ammonia excretion and biochemical composition were studied for three species of starving zooplankton (Calanus finmarchicus, Sagitta elegans, and Acartia clausi). Over the period of starvation, the respiration rate of all three species followed the same pattern of an initial decrease followed by a more or less constant level. A similar pattern was observed for the ammonia excretion rate of S. elegans and A. clausi, whereas C. finmarchicus excretion appeared to oscillate between high and low levels of protein catabolism. Study of the biochemical changes showed that C. finmarchicus consumed primarily lipids, and at times proteins, to meet its energy requirement whereas S. elegans and A. clausi primarily used protein. Variations in the elemental composition as well as the O:N ratio confirmed that C. finmarchicus alternated between periods of protein-dominant catabolism and lipid-dominant catabolism during starvation. No similar change in catabolism was observed in the two other species. The results are discussed in terms of physiological mechanisms of resistance to starvation and were used to calculate the energy budget of S. elegans and C. finmarchicus during the period of total starvation. The significance of such budgets is discussed and some of the sources of error examined.Bedford Institute of Oceanography Contribution.     

Nitrogen and energy loss via nonfaecal and faecal excretion in the marine teleost Lithognathus lithognathus

Nitrogen excretion and assimilation efficiencies of individual Lithognathus lithognathus (Cuvier 1830), a marine teleost from high energy surf zones in Algoa Bay, South Africa, were determined under laboratory conditions in 1985. Nonfaecal-nitrogen excreted by starved and fed L. lithognathus consists mainly of ammonia with urea and amino acids as secondary excretory products. Ammonia excretion rates were temperature dependant with the excretion rate of starved fish significantly lower than those of fed fish, at all three experimental temperatures. The mass component b of the mass/ammonia excretion equation was temperature independent and ranged from 0.651 to 0.700 and 0.589 to 0.635 for starved and fed fish respectively. The mean percentage of food energy lost via dissolved nonfaecal excretory products (exogenous plus endogenous) was 6.11±6.07%. Assimilation efficiencies ranged from 70.75 to 99.29% for dry matter and from 95.72 to 99.58% on an energy basis. The combined nonfaecal and faecal energy loss was calculated at 11.87% of the ingested energy. The benthic feeding ichthyofauna recycle 255 g total nitrogen per metre strip per year which constitutes 2% of total phytoplankton nitrogen requirements of the surf zone.     

Physiological and biochemical aspects of embryonic and larval development of the winter flounder Pseudopleuronectes americanus

Eggs and larvae of the winter flounder Pseudopleuronectes americanus Walbaum were hatched and raised in the laboratory under controlled conditions. Biochemical composition was measured during development and found to be similar to that of other species: 65 to 80 percent protein, 15 to 30 percent fat, and 0 to 5 percent carbohydrate. Ash content was 7 to 10 percent of dry weight. The chorion comprised more than half of the weight of an egg and the data suggested that it was possibly a source of nutrition to the developing embryo. The sequence of utilization appeared to be carbohydrate and then protein to hatching, lipid, mixed lipid and protein, the predominantly protein until feeding began. Carbohydrate was accumulated at first feeding and depleted when growth began. Protein and lipid were deposited in approximately constant proportions. Respiration rates of eggs were low, 0.002–0.015 l O₂ egg^-1 h^-1, but rose gradually from fertilization to hatching. Respiration rates of early larvae were from two to eight times that of eggs (0.033–0.131 l O₂ larva^-1 h^-1). Variation in larval respiration rates indicated a three-fold difference in rate according to level of activity. Eggs excreted ammonia at an increasing rate from fertilization to hatching. Larvae excreted ammonia, primary amines, and other unidentified organic nitrogenous substances. Rates of excretion and proportions of excretory products varied with stage of development. Primary amine excretion was variable and a major component in early stages. Ammonia-N excreted was two to 20 times primary amine N excreted. Unidentified substances were the predominant form of N excretion during early feeding. Ammonia accounted for most of the N excreted in older larvae. Early specific growth rates were 2.1 and 5.5%. Net caloric conversion and net and gross nitrogen efficiencies were low in first feeding larvae compared to adult fishes (32.2, 27.7, and 10.7% respectively).Contribution no. 5071 from the Woods Hole Oceanographic Institution     

温度、体重对皱纹盘鲍耗氧量和排氨量的影响

皱纹盘鲍的耗氧量和排氨量的实验结果表明：温度和体重对鲍的耗氧量和排氨量有明显的影响，且两固子间存在着交互作用，皱纹盘鲍的个体耗氧量「Ｑ（Ｏ）／ｍｇｈ＾－１」和排氧量「Ｑ（Ｎ）／μｍｏｌｈ＾－１」与温度（θ／℃）和湿重（ｍ／ｇ）存在下列关系：Ｑ（Ｏ）＝０．０１２４ｍ＾０．８５５６１．０８５３＾θ，Ｑ（Ｎ）＝０．０２０７ｍ＾０．８５５６１．０８５３＾θ。２０℃皱纹盘鲍存在代谢的转变点。     

Effects of feeding history on mobilisation and deposition of body constituents and on growth in juvenile Ambassis vachelli (Pisces: Chandidae)

Young Ambassis vachelli (Richardson) 40 to 50 d old, were used in a laboratory experiment to test the effects of starvation and subsequent re-feeding on body constituents and growth. Fish in three laboratory treatments (fed continuously; fed for 9 d and then starved for 15 d; starved for 9 d and fed for 15 d), were compared to fish from a local field population. Different body stores were mobilised at different times and rates during starvation. Carbohydrates were mobilised from the onset of starvation and were depleted after 3 d. Lipid and protein were mobilised at an increasing rate from the onset of starvation. The mortality in starved fish was relatively high (up to 70%) until re-feeding. Upon feeding, all body stores were restored rapidly in fish that were starved, with carbohydrate levels displaying an overshoot (carbohydrate level exceeding normal levels) in comparison to the levels in continuously fed fish. After 15 d of feeding, the starved fish had levels of constituents similar to those in continuously fed fish. Fish that were fed and subsequently starved were able to maintain themselves for at least 15 d prior to death, indicating a better degree of starvation resistance than fish without a history of feeding. This implies that feeding history in the early life of a fish is important in growth and survival but that young fish may have growth regimes flexible enough to survive relatively long periods of starvation.