Oxygen uptake and comparative energetics among eggs and larvae of three subtropical marine fishes

Oxygen uptake was measured and energy budgets were determined for eggs and larvae of bay anchovy (Anchoa mitchilli), sea bream (Archosargus rhomboidalis) and lined sole (Achirus lineatus). Using an energetics model, both minimum rations and prey levels were estimated. The Q_{O 2} [μl (mg dry wt)^-1 h^-1] increased approximately five-fold for all species during development from egg to feedingstage larvae. The weight exponents in power functions, R=aW^b, relating oxygen uptake (R) to dry weight (W) did not differ significantly among species. The exponents were: bay anchovy, 0.9770; sea bream, 0.8382; lined sole, 0.9416. Oxygen uptake was generally lower for bay achovy than for the other species. In the energy budgets issimilation efficiencies of all species were low (24–75%). Gross growth efficiencies ranged from 11–41%, and net growth efficiencies from 38–57%. An estimated 32–83% of ingested energy was excreted in feces and urine. The lavae used a relatively small proportion of ingested energy in metabolism (6–31%). The anchovy had the lowest assimilation and gross growth efficiencies, and the highest excretion rate. Consistent with the better growth and survival observed in previous experiments, sea bream were the most efficient assimilators and had the lowest required minimum rations. Required minimum prey levels for first-feeding larvae ranged from 400 copepod nauplii l^-1 for sea bream and bay anchovy to 1 000 l^-1 for lined sole. The estimated required prey levels were higher than levels at which significant survival had been observed.     

Food concentration and stocking density effects on survival and growth of laboratory-reared larvae of bay anchovy Anchoa mitchilli and lined sole Achirus lineatus

Bay anchovy (Anchoa mitchilli) eggs were stocked at densities from 0.5 to 32.0 l^-1 and larvae were fed on wild plankton (copepod nauplii) in concentrations that ranged from 50 to 5000 prey l^-1. Lined sole (Achirus lineatus) eggs were stocked at 0.5 to 16.0 l^-1 and larvae were fed wild plankton at concentrations from 50 to 1000 prey l^-1. Some larvae of each species survived at all stock and food levels to the transformation stage at 16 days after hatching. Survival rates for both species exceeded 40% when food concentration was 1000 l^-1 or higher. Growth and dry weight yields also increased significantly at the higher food concentrations. Effects of initial stocking density were not well defined, but both survival and growth decreased at the highest stocking rates. Standardized culture of bay anchovy and lined sole larvae can be based on a food concentration of 1000 copepod nauplii l^-1 to routinely produce healthy larvae.     

Yolk resorption,onset of feeding and survival potential of larvae of three tropical marine fish species reared in the hatchery

This paper provides basic early life-history information on milkfish (Chanos chanos), seabass (Lates calcarifer) and rabbitfish (Siganus guttatus) which may explain in part the observed differences in their survival performance in the hatchery. Egg size, larval size, amount of yolk and oil reserves and mouth size are all greater in milkfish than in seabass, and greater in the latter than in rabbitfish. During the first 24 h after hatching, rabbitfish larvae grow much faster than milkfish and seabass larvae at similar ambient temperatures (range 26°–30°C, mean about 28°C). The eyes become fully pigmented and the mouths open earlier in seabass and rabbitfish (32–36 h from hatching) than in milkfish (54 h). Seabass larvae learn to feed the earliest. Yolk is completely resorbed at 120 h from hatching in milkfish, and yolk plus oil at 120 h in seabass and 72 h in rabbitfish at 26° to 30°C. Milkfish and seabass larvae have more time than rabbitfish to initiate external feeding before the endogenous reserves are completely resorbed. Delayed feeding experiments showed that 50% of unfed milkfish larvae die at 78 h and all die at 150 h from hatching. Milkfish larvae fed within 54 to 78 h after hatching had improved survival times: 50% mortality occurred at 96 to 120 h, and 10 to 13% survived beyond 150 h. Unfed seabass larvae all died at 144 h, while 6 to 13% of those fed within 32 to 56 h after hatching survived beyond 144 h and well into the subsequent weeks. Unfed rabbitfish larvae all died at 88 h, while 7 to 12% of those fed within 32 to 56 h after hatching survived beyond 88 h. A delay in initial feeding of more than 24 h after eye pigmentation and opening of the mouth may be fatal for all three species.Contribution No. 167 from the SEAFDEC Aquaculture Department     

The influence of temperature on the development of Baltic Sea sprat (Sprattus sprattus) eggs and yolk sac larvae

In spring 2004 and 2005 we performed two sets of experiments with Baltic sprat (Sprattus sprattus balticus Schneider) eggs and larvae from the Bornholm Basin simulating ten different temperature scenarios. The goal of the present study was to analyse and parameterise temperature effects on the duration of developmental stages, on the timing of important ontogenetic transitions, growth during the yolk sac phase as well as on the survival success of eggs and early larval stages. Egg development and hatching showed exponential temperature dependence. No hatching was observed above 14.7°C and hatching success was significantly reduced below 3.4°C. Time to eye pigmentation, as a proxy for mouth gape opening, decreased with increasing temperatures from 17 days post hatch at 3.4°C to 7 days at 13°C whereas the larval yolk sac phase was shortened from 20 to 10 days at 3.8 and 10°C respectively. Maximum survival duration of non-fed larvae was 25 days at 6.8°C. Comparing the experimental results of Baltic sprat with existing information on sprat from the English Channel and North Sea differences were detected in egg development rate, thermal adaptation and in yolk sac depletion rate (YSDR). Sprat eggs from the English Channel showed significantly faster development and the potential to develop at temperatures higher than 14.7°C. North Sea sprat larvae were found to have a lower YSDR compared to larvae from the Baltic Sea. In light of the predictions for global warming, Baltic sprat stocks could experience improved conditions for egg development and survival.     

Effects of temperature on oxygen consumption,growth, and development of embryos and yolk-sac larvae of Siganus randalli (Pisces: Siganidae)

Eggs from laboratory spawnings of the coralreef fish Siganus randalli Woodland were incubated at two temperatures (27 and 30 °C). Eggs and larvae were sampled until larval starvation, while changes in oxygen consumption, growth, yolk utilization, and development were monitored. Oxygen consumption, which peaked at hatching, was higher for embryos incubated at 30 °C than at 27 °C. Rates of oxygen consumption (nl h^-1 individual^-1) at hatching were similar to those for other temperate and tropical species. Rates of oxygen consumption by yolk-sac larvae were highly variable, and these data suggest that larval oxygen consumption prior to yolk-sac absorption may not be significantly influenced by temperature. Rates of yolk depletion were higher for larvae at the higher temperature. After an initial rapid increase in length, length of larvae at 30 °C decreased with age. Egg size, egg weight, and maximum notochord length of larvae differed significantly between spawns. Age-specific oxygen consumption rates by the embryos varied between spawns, but regressions describing oxygen consumption as a function of age did not differ significantly. The initiation and completion of eye pigmentation were used as developmental markers to calculate the amount of yolk remaining for larvae at the different temperatures. Larvae maintained at 30 °C completed eye pigmentation approximately 3 h sooner than those maintained at 27 °C, but had less endogenous reserves. This finding indicates a trade-off between rapid development and efficient utilization of the endogenous reserves. The completion of eye pigmentation in larvae incubated at the higher temperature occurred at midnight and, depending on the amount of time that the larvae have to initiate feeding prior to the point-of-no-return, the timing of completion of eye pigmentation could influence larval survival.     

Effect of temperature on viability and axial muscle development in embryos and yolk sac larvae of the Northeast Arctic cod (Gadus morhua)

Cod (Gadus morhua L.) eggs may develop and hatch within temperatures of −1.5 to 12 °C, but little is known about the effects of very low temperatures on larval characteristics. Eggs of the Northeast Arctic cod (Gadus morhua) were incubated at 1, 5 or 8 °C from Day 1 after fertilisation until hatching, and transferred to 5 °C after hatching. Histological samples of the axial musculature were taken at hatching and 5 d after hatching, and the data on muscle cellularity from these samples were related to survival and hatching, size, developmental data and viability of the yolk sac larvae. All larvae hatched at the same developmental stage. Incubation of eggs at 1 °C produced shorter larvae with a larger yolk sac and more, small deep fibres at hatching than larvae from eggs incubated at 5 or 8 °C. The larval size difference was still present 5 d after hatching, a time at which the larvae from 1 °C-incubated eggs were less developed and less resistant to an acute viability stress test (65 ppt salinity). Although there were no differences between temperature groups in number and size of muscle fibres 5 d after hatching, the deep fibres of the 1 °C-group contained less myofibrils than the two other groups. The phenotype of the larvae at hatching was thus affected within these incubation temperatures. Although all groups were transferred to the same temperature after hatching, the lowest egg incubation temperature (1 °C) still had a negative effect 5 d after hatching, as these larvae were both smaller, less resistant to stress and had less functional muscles at the time of first feeding. Our conclusion is therefore that 1 °C is close to, or below, the lower thermal tolerance limit for normal functional development of Northeast Arctic cod. The results are discussed in relation to larval viability and recruitment of this species in the wild. Received: 4 February 1998 / Accepted: 10 July 1998     

Effects of temperature on yolk utilization,initial growth,and behaviour of unfed marine fish-larvae

The effect of temperature on yolk utilization, initial growth and behaviour of larvae of four species of marine fishes, i.e.,Acanthopagrus schlegeli, Engraulis japonica, Pagrus major andParalichthys olivaceus, was investigated under laboratory conditions at Hiroshima in 1989. The yolk sac was absorbed earlier with increasing temperature for all species. Morphological characters such as pectoral fin appearance, mouth opening and eye pigmentation differed from species to species over the range of experimental temperatures, and the sequence of development of these characters varied with temperature even within the same species. Temperature did not have a large influence on the maximum growth of unfed larvae, but had a clear effect on the time to starvation, occurring earlier at higher temperatures. Larval behaviour as indicated by time spent moving was also influenced by temperature; behavioural activity was greater at higher temperatures for all species examined. The effect of temperature on the early stage of larval life is discussed in terms of behavioural and morphological developments, as well as on yolk utilization, and its influence on larval survival in nature and under rearing conditions is evaluated.     

Comparison of somatic growth and otolith increment growth in laboratory-reared larvae of Pacific saury, Cololabis saira, under different temperature conditions

The relationship between somatic growth and incremental growth of otoliths of Pacific saury, Cololabis saira (Brevoort), larvae under different temperature conditions was studied in the laboratory for three age groups (0 to 9, 10 to 20 and 20 to 30 d posthatch). Larvae were incubated from hatching to 9 d at 24, 20, and 16 °C. Further, larvae initially reared at an ambient temperature of 21.7 °C were transferred to experimental temperatures of 22, 18, and 14 °C on Day 10 and reared to Day 20 and similarly from Day 20 and reared to Day 30 posthatch. Growth trajectories of larvae sampled at the end of the three experiments were back-calculated using the biological intercept method and compared to the measured values 0 and 5 d after the start of each experiment. Back-calculated knob length at the different temperatures indicated no significant difference to the measured knob lengths except for the cases at 20 °C from hatching to 9-d-old larvae and at 14 °C from 20- to 30-d-old larvae. The close correlation between somatic and otolith growth shown in this study indicated the feasibility of estimating the growth history of Pacific saury larvae using otolith readings. Received: 14 April 1999 / Accepted: 27 October 1999     

Energy metabolism during development of eggs and larvae of gilthead sea bream (Sparus aurata)

Developing eggs and larvae of laboratory-reared gilthead sea bream (Sparus aurata) maintained in filtered seawater (40 ppt) at 18°C, were measured for oxygen uptake, ammonia excretion, contents of free amino acids (FAA), protein, fatty acids (FA) accumulated ammonia, and volumes of yolk-sac and oil globule. Absorption of the yolk coincided with the consumption of FAA and was complete ca. 100 h post-fertilisation. Amino acids from protein were mobilised for energy in the last part of the yolk-sac stage. Absorption of the oil globule occurred primarily after hatching following yolk absorption, and correlated with catabolism of the FA neutral lipids. Overall, FAA appear to be a significant energy substrate during the egg stage (60 to 70%) while FA from neutral lipids derived from the oil globule are the main metabolic fuel after hatching (80 to 90%).     

Food selection by laboratory-reared larvae of the scaled sardine Harengula pensacolae (Pisces,Clupeidae) and the bay anchovy Anchoa mitchilli (Pisces,Engraulidae)

Food selection by laboratory-reared larvae of scaled sardines Harengula pensacolae, and bay anchovies Anchoa mitchilli, was compared. Natural plankton was fed to the larvae during the 22 days following hatching. Food levels in the rearing tanks were maintained at an average of 1,600 to 1,800 potential food organisms per liter. Larvae of both species selected as food copepod nauplii, copepodites, and copepods; initial feeding was on organisms of 50 to 75 body width. Larvae of H. pensacolae averaged 4.2 mm in total length at hatching and those of A. mitchilli about 2.0 mm. H. pensacolae larvae grew about 1.0 mm per day and A. mitchilli 0.70 mm per day. The mean number of food organisms in each digestive tract was greater in H. pensacolae than in A. mitchilli, and the difference in number increased as the larvae grew. Average size of food organisms eaten increased for both species with growth, because of selection by the larvae; the average size of copepodites and copepods in digestive tracts increased at a faste rate in H. pensacolae than A. mitchilli. A. mitchilli longer than 8 mm did not eat copepod nauplii.Contribution No. 170, Bureau of Commercial Fisheries Tropical Atlantic Biological laboratory, Miami, Florida 33149, USA.     

Feeding of laboratory-reared larvae of the sea bream Archosargus rhomboidalis (Sparidae)

Feeding by larvae of the sea bream Archosargus rhomboidalis (Linnaeus) was investigated from late September, 1972 to early May, 1973 using laboratory-reared larvae. Fertilized eggs were collected from plankton tows in Biscayne Bay, and the larvae were reared on zooplankton also collected in plankton nets. Techniques were developed to estimate feeding rate, food selection, gross growth efficiency, and daily ration. Daily estimates of these were obtained through 16 days after hatching at rearing temperatures of 23°, 26°, and 29°C. Feeding rate increased exponentially as the larvae grew, and increased as temperature was raised. At 23°C larvae began feeding on Day 3, at 26° and 29°C larvae began feeding on Day 2. Feeding rates at initiation of feeding and on Day 16 were, respectively: 23°C, 7.16 food organisms per larva per hour (flh) and 53.78 flh; 26°C, 7.90 flh and 168.80 flh; 29°C, 17.62 flh and 142.07 flh. Sea bream larvae selected food organisms by size. At initiation of feeding they selected organisms less than 100 m in width. As larvae grew they selected larger organisms and rejected smaller ones. The major food (more than85% of the organisms ingested) was copepod nauplii, copepodites, and copepod adults. Minor food items were barnacle nauplii, tintinnids, invertebrate eggs, and polychaete larvae. Mean values for gross growth efficiency of sea bream larvae ranged from 30.6% at 23°C to 23.9% at 29°C. Mean values for daily ration, expressed as a percentage of larval weight, ranged from 84% at 23°C to 151% at 29°C and tended to decline as the larvae grew.This paper is a contribution from the Rosenstiel School of Marine and Atmospheric Science, University of Miami, Miami, Florida, USA     

Growth, carbon, nitrogen and caloric content of Solea senegalensis (Pisces: Soleidae) from egg fertilization to metamorphosis

Eggs and larvae of the Senegal sole, Solea senegalensis Kaup, were reared from fertilization until the end of metamorphosis, which occurs by Day 17 after hatching at 19.5 °C. Changes in energy content and biomass quality were studied in terms of dry weight and of carbon, nitrogen and energy content. S. senegalensis spawned eggs of about 1 mm diameter which hatched 38 h after fertilization. Average dry weight of individual eggs was 46 μg, the chorion accounting for about 18% of total dry weight. Gross energy of recently fertilized sole eggs was approximately 1 J egg⁻¹. From fertilization to hatching, eggs lost 8% of their total energy (chorion not included). After hatching, larvae lost 14% of their initial energy until the start of feeding which occurred about 48 h afterwards. The principal components catabolized during embryogenesis were carbon-rich compounds that decreased by 26%, while nitrogen-rich compounds decreased by only 10% and were practically unaltered from hatching to the start of feeding. Feeding larvae displayed constant growth during the period studied (specific growth rate on a dry weight basis was 0.26 d⁻¹). The relative proportion of carbon and nitrogen content revealed an accumulation of high energy compounds in the days before metamorphosis. By Day 14, the energy content reached values similar to those of recently hatched embryos, but decreased again during metamorphosis. Received: 10 June 1998 / Accepted: 28 January 1999     

Assimilation of dissolved glucose from sea water by the sea bass Dicentrarchus labrax and the sea bream Sparus aurata

Larvae of sea bass (Dicentrarchus labrax) and sea bream (Sparus aurata) were transferred to normal or glucose-enriched sea water immediately after mouth opening to assess their ability to absorb and assimilate glucose at the beginning of the larval period. Assimilation was monitored by histological and cytochemical analysis of the liver. The results showed that (1) the larvae of both species regularly ingested water, (2) glucose absorption resulted in glycogen accumulation in the hepatocytes (this was more marked in sea bass than in sea bream), and (3) glucose delayed the pathological effects of fasting. Consideration of metabolic derivatives indicates that hepatic glycogen probably arises from neoglucogenesis.     

Elevated temperature impairs onset of symbiosis and reduces survivorship in larvae of the Hawaiian coral, <Emphasis Type="Italic">Fungia scutaria</Emphasis>

Many corals obtain their obligate intracellular dinoflagellate symbionts from the environment as larvae or juveniles. The process of symbiont acquisition remains largely unexplored, especially under stress. This study addressed both the ability of Fungia scutaria (Lamarck 1801) larvae to establish symbiosis with Symbiodinium sp. C1f while exposed to elevated temperature and the survivorship of aposymbiotic and newly symbiotic larvae under these conditions. Larvae were exposed to 27, 29, or 31°C for 1 h prior to infection, throughout a 3-h infection period, and up to 72 h following infection. Exposure to elevated temperatures impaired the ability of coral larvae to establish symbiosis and reduced larval survivorship. At 31°C, the presence of symbionts further reduced larval survivorship. As sea surface temperatures rise, coral larvae exposed to elevated temperatures during symbiosis onset will likely be negatively impacted, which in turn could affect the establishment of future generations of corals.     

The effects of temperature and salinity on egg production and hatching success of Baltic Acartia tonsa (Copepoda: Calanoida): a laboratory investigation

The functional response of the aspects of reproductive success of a southwestern Baltic population of Acartia tonsa (Copepoda: Calanoida) was quantified in the laboratory using wide ranges in temperatures and salinities. Specifically, daily egg production (EP, # female⁻¹ day⁻¹) was determined for 4 or 5 days at 18 different temperatures between 5 and 34°C and the time course and success of hatching were evaluated at 10 different temperatures between 5 and 23°C. The effect of salinity (0 to 34 psu) on egg hatching success was also examined. The highest mean rates of EP were observed between 22 and 23°C (46.8–50.9 eggs female⁻¹ day⁻¹). When studied at 18 psu, hatching success of eggs increased with increasing temperature and was highest (92.2%) at 23°C. No hatching was observed for eggs incubated at low temperatures (≤12°C) that were produced by females acclimated to temperatures ≤10°C indicating a possible thermal threshold between 10.0 and 13.0°C below which only the production of diapause (or low quality) eggs exists in this population. When tested at 18°C, the hatching success of eggs incubated at 15 different salinities increased asymptotically with increasing salinity and was maximal (81.4–84.5%) between 17 and 25 psu. The high reproductive success observed over wide ranges in temperatures and salinities in this Baltic population demonstrates one of the mechanisms responsible for the cosmopolitan distribution of this species within productive, estuarine and marine habitats.     

Description of the early development of the halibut Hippoglossus hippoglossus and attempts to rear the larvae past first feeding

Rearing experiments on the halibut Hippoglossus hippoglossus (L.) were carried out using gametes from parents caught at a depth of 600 to 700 m off the Norwegian coast in February 1980. After fertilization, the average egg diameter was 3.08 mm, average dry weight 1 038 μg and neutral buoyancy was at 36.5‰ S. The eggs hatched after 20 d at 4.7°C, 18 d at 5°C and 13 d at 7°C. Survival to hatching was better when antibiotics were used. At hatching the larvae were 6.4 mm long, there were no functional eyes or mouth, but prominent neuromast organs were present. Resorption of yolk lasted 50 d at 5.3°C; the eyes and mouth were then functioning and the larva was about 11.5 mm long. The larvae were offered zooplankton as food, but with little success in initial feeding. A few larvae fed and grew in 2 500-litre plastic bags, one reaching a length of 24 mm after 90 d.     

Potential impact of climate warming on the recruitment of an economically and ecologically important species, the European lobster (Homarus gammarus) at Helgoland, North Sea

A laboratory-based study was performed to assess the impact of climate warming on the recruitment of the endangered population of the European lobster (Homarus gammarus) at Helgoland (North Sea, German Bight). Egg-bearing females collected in situ just after spawning in late summer were subjected to various seasonal temperature regimes. Regimes with elevated temperatures (mild winters) resulted in a strong seasonal forward shift of larval hatching. Hatching took place at significantly lower temperatures than under regimes with normal winters. Experiments on larval development across a range of constant temperatures showed that no successful larval development occurred at temperatures below 14°C. Larval survival increased from 9% at 14°C to 80% at 22°C, while duration of larval development decreased correspondingly from 26 to 13 days. We hypothesize that an ongoing warming of the North Sea will strongly affect the recruitment success of the Helgoland lobster, mainly resulting from a decoupling of the seasonal peak appearance of larvae from optimal external conditions (temperature, food availability) for larval development.     

RNA/DNA ratio and expression of 18S ribosomal RNA, actin and myosin heavy chain messenger RNAs in starved and fed larval Atlantic cod (Gadus morhua)

Levels of total RNA, total DNA, 18S ribosomal RNA (rRNA), poly(A) messenger RNA (mRNA), and two mRNAs coding for abundant myofibrillar proteins were estimated in laboratory-reared Atlantic cod larvae (Gadusmorhua Linnaeus) under conditions of feeding and starvation. DNA probes specific for cod 18S rRNA, β-actin mRNA and myosin heavy chain mRNA were developed. In two experiments on newly hatched larvae in fed and starved treatments, changes in 18S rRNA and mRNA were similar to changes in total RNA during the first weeks after hatching. RNA levels in fed and starved larvae in both experiments were stable, or increased, over the first 3 d after hatching, and then decreased to minima at 9 d. RNA levels increased after 9 d, with the degree and timing of the increase varying among the individual classes of RNA. Complete mortality of starved larvae in both experiments was observed shortly after 11 d, corresponding to exhaustion of endogenous yolk reserves. Total RNA content, RNA/DNA ratio, 18S rRNA levels, total mRNA pool, and actin and myosin heavy chain mRNA levels showed significant differences in fed and starved first-feeding larvae after yolk exhaustion. In another experiment with 3- to 4-week-old cod larvae, 18S rRNA levels were significantly lower in starved versus fed larvae after 3 d. Total RNA responded to feeding and starvation within a similar time as 18S rRNA and the mRNAs examined. Analysis of bulk nucleic acids using fluorometric dyes was simpler and faster than analysis of individual RNAs using hybridization probes, and provides valuable information on recent growth and condition of individual larvae. However, analysis of specific RNAs can provide information on expression of the corresponding genes and reveal the changes underlying trends seen in bulk RNA. Received: 9 February 1996 / Accepted: 7 June 1999