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     

Variability in larval development of the tropical reef fish Pomacentrus amboinensis (Pomacentridae): the parental legacy

The effects of food availability, female size, and social interactions on the quality of Pomacentrus amboinensis larvae at hatching were examined using two field-based experiments. In Experiment 1, food availability and female size significantly influenced size, eye diameter and levels of yolk reserves of larvae at hatching. Small females (47 to 52 mm standard length, SL) whose diets were not supplemented, produced the longest larvae (3.0 ± 0.01 mm total length, TL) with the least yolk reserves (50.1 ± 1.04 μm²). Irrespective of female size, those that received additional food produced larvae with the largest yolk-sacs (large females: 87.60 ± 1.53 μm²; small females: 80.14 ± 1.24 μm²). In Experiment 2, interactions with conspecifics had a greater affect on the somatic development of larvae at hatching than food availability. Increased social interactions resulted in larvae that were ⋍3% longer, with 2% greater head depth, than larvae from females that spawned in isolation on the experimental reefs. Fed females produced larvae with ⋍20% more yolk than larvae from females whose diets were not supplemented. All three factors (food availability, female size, and intensity of social interactions) tested within these experiments vary spatially and temporally among reefs. There is the potential, therefore, for larvae at the onset of the planktonic stage to vary in quality, level of development, and probability of survival. Received: 12 August 1996 / Accepted: 26 August 1996     

Poecilogony in an estuarine opisthobranch: planktotrophy, lecithotrophy, and mixed clutches in a population of the ascoglossan Alderia modesta

A San Diego population of the opisthobranch mollusc Alderia modesta (Lovén, 1844) exhibits poecilogony, the presence of two development modes within a single species. In spring, half of the adults spawned masses containing ∼300 eggs with a mean diameter of 68 μm. After 3 d, these egg masses hatched planktotrophic veligers with a maximum shell dimension of 116 μm. The remaining adults spawned masses containing ∼30 eggs with a mean diameter of 105 μm. These egg masses hatched after 5 to 6 d, releasing lecithotrophic larvae with a maximum shell dimension of 186 μm. About 1% of field-collected adults produced mixed clutches containing a continuum of larval sizes, spanning the size extremes of planktotrophy and lecithotrophy and hatching larvae with a mean maximum shell dimension of 152 μm. Adults producing planktotrophic and lecithotrophic larvae were interfertile, and no hybrid breakdown was observed through the F3 generation. When starved, adults which previously produced only lecithotrophic larvae switched to producing planktotrophic larvae or mixed clutches with both planktotrophic and lecithotrophic larvae. Sequence-polymorphisms from a portion of the mitochondrial cytochrome c oxidase I gene support the conclusion that the two reproductive morphs represent a single species. Most of the lecithotrophic larvae and a small percentage of the larvae from mixed clutches were metamorphically competent within 3 d of hatching. A. modesta is the only molluscan species as yet known to have both planktotrophic and pelagic lecithotrophic development within a single natural population. Received: 14 August 1997 / Accepted: 11 April 1998     

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     

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     

Osmoregulation, nutritional effects and buoyancy of marine larval fish: a bioassay for assessing density changes during the earliest life-history stages

It has been hypothesized that marine fish larvae in the advanced stages of starvation would show increased density (ρ = mass volume⁻¹) from water loss due to osmoregulation failure. Changes in larval buoyancy are currently attributed to swim bladder regulation and protein synthesis or catabolism. Osmoregulation-related changes in density is an alternative mechanism, the importance of which remains untested in the laboratory and the influence of which on vertical distributions is unknown. We provide evidence that loss of osmotic control is a plausible mechanism for increased density of larval cod (Gadus morhua L.). Furthermore, our results show that this mechanism is not restricted to larvae in the advanced stages of starvation. “Relative” larval densities are estimated using a modified density gradient. We use a gravimetric method to separate the effects of nutrition from osmoregulation failure. We assessed the importance of sampling strata on estimates of larval density. Proportional sampling within three depth strata (stratified sample) produced the least biased method for determining the “average” density of a population of larvae in laboratory culture. Larvae sampled from the bottom third of the culture tank were significantly more dense then those sampled from the surface. This was true for larvae of all ages. The average change in density from hatching till death from starvation for larvae sampled in the surface stratum was nominal (Δρ = 5.0 × 10⁻⁴ g cm⁻³), while the change for those sampled from the bottom stratum was large (Δρ = 3.8 × 10⁻³ g cm⁻³). These large density differences suggest that larvae sampled from the bottom stratum were either osmotically stressed or were facultatively changing their density via regulatory pathways. Preliminary observations suggest that vitality is lower amongst those larvae which are sampled near the bottom. The small change in average density of larvae sampled from the surface stratum was due to starvation. The density differences we observed between “osmotically stressed” and “starving” larvae could readily have been misconstrued as differences in feeding and growth experienced by individual larvae. The potential bias of increased density from osmoregulation failure must be considered as a factor in experimental designs developed to assess the effect of fed and starved treatments on buoyancy for larvae of all ages. The simple bioassay we describe may prove useful both as a means of assessing larval condition and as a mechanism for evaluating factors affecting larval vertical distributions in the field. Received: 13 January 1997 / Accepted: 3 February 1997     

The effect of food availability,age or size on the RNA/DNA ratio of individually measured herring larvae: laboratory calibration

RNA/DNA ratios in individual herring (Clupea harengus) larvae (collected from Kiel Bay, Baltic Sea, in 1989) were measured and proved suitable for determining nutritional status. Significant differences between fed and starving larvae appeared after 3 to 4 d of food deprivation in larvae older than 10 d after hatching. The RNA/DNA ratio showed an increase with age or length of the larvae and was less pronounced in starving larvae compared to fed larvae. The individual variability of RNA/DNA ratios in relation to larval length of fed larvae and of larvae deprived of food for intervals of 6 to 9 d is presented. Based on the length dependency and the individual variability found within the RNA/DNA ratios, a laboratory calibration is given to determine whether a larva caught in the field has been starving or not. An example for a field application is shown.     

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.     

Facultative lecithotrophy during larval development of the burrowing shrimp Callianassa tyrrhena (Decapoda: Callianassidae)

Survival, developmental and consumption rate (Artemia nauplii ingested per day) as well as predation efficiency (ingested per available Artemia nauplii) were studied during the larval development of the shallow-water burrowing thalassinid Callianassa tyrrhena (Petagna, 1792), which exhibits an abbreviated type of development with only two zoeal stages and a megalopa. The larvae, hatched from berried females from S. Euboikos Bay (Aegean Sea, Greece), were reared at 10 temperature–food density combinations (19 and 24 °C; 0, 2, 4, 8 and 16 Artemia nauplii d⁻¹). Enhanced starvation resistance was evident: 92 and 58% of starved zoeas I molted to zoea II, while metamorphosis to megalopa was achieved by 76 and 42% of the hatched zoeas at 19 and 24 °C, respectively. The duration of both zoeal stages was affected by temperature, food density and their interaction. Nevertheless, starvation showed different effects at the two temperatures: compared to the fed shrimp, the starved zoeae exhibited accelerated development at 19 °C (8.4 d) but delayed metamorphosis at 24 °C (5.9 d). On the other hand, both zoeal stages were able to consume food at an increased rate as food density and temperature increased. Predation efficiency also increased with temperature, but never exceeded 0.6. Facultative lecithotrophy, more pronounced during the first zoeal stage of C.tyrrhena, can be regarded as an adaptation of a species whose larvae can respond physiologically to the different temperature–food density combinations encountered in the wide geographical range of their natural habitat. Received: 28 February 1998 / Accepted: 21 October 1998     

Free amino acids and energy metabolism in eggs and larvae of seabass, Lates calcarifer

Contents of free amino acids (FAA), protein and ammonium ions together with rates of ammonia excretion and oxygen consumption were measured in order to study the role of FAA as an energy substrate in developing eggs and larvae of seabass (Lates calcarifer) maintained in seawater (30 ppt) at 28 °C without feeding. Initially eggs contained 25.3 nmol ind⁻¹ of FAA of which 21.5 nmol was rapidly utilised by the developing eggs and larvae during the period up to 40 h post spawning (PS) when nearly all the yolk had been resorbed. During the same period, a net increase in protein content of 1.7 μg ind⁻¹ was observed, indicating that the major part of the amino acids lost from the free pool had been polymerised into body proteins. Assuming that the balance of the FAA after protein synthesis was used entirely for energy metabolism, FAA appeared to be an important energy substrate during the embryonic stages (2 to 16 h PS); after hatching, the contribution of FAA to energy metabolism was less significant. From 50 h PS until the end of the study period at 100 h PS, amino acids derived from somatic protein were used for energy metabolism. For the overall period from just after spawning up to 100 h PS, the data indicate that ca. 14% of the total aerobic energy metabolism was derived from amino acid catabolism. Received: 26 September 1997 / Accepted: 1 April 1998     

Diet of 0-group stages of capelin (<Emphasis Type="Italic">Mallotus villosus</Emphasis>), herring (<Emphasis Type="Italic">Clupea harengus</Emphasis>) and cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) during spring and summer in the Barents Sea

Recruitment of capelin in the Barents Sea fail when juvenile herring and cod are abundant and the potential for feeding competition of wild sympatric capelin and herring larvae and small cod juveniles were investigated. The frequency of gut evacuation after capture of capelin larvae were also studied in mesocosms. Small capelin larvae (<35 mm length) fed on small prey including phytoplankton, invertebrate eggs and nauplii, bivalves, other invertebrate larvae and small copepods. Calanus copepodites were only observed in large capelin larvae (>26 mm length). Calanus copepodites were the major food sources for contemporary herring larvae (25–35 mm length) and Calanus and euphausiids were the major prey for small juvenile herring (37–60 mm length) and cod (18–40 mm length). Capelin larvae reared in mesocosms evacuated the guts shortly after capture. Capelin larvae had a smaller mouth and fed on smaller prey than herring and cod of the same length. This implies that the small capelin larvae, in contrast to sympatric small herring and cod, are not tightly linked to the food chain involving Calanus and euphausiids. Thus, exploitative competition between capelin larvae and planktivorous fish that rely on Calanus and euphausiids in the Barents Sea may be relaxed.     

Asymptotic growth trajectories of larval sardine (Sardinops melanostictus ) in the coastal waters off western Japan

Growth trajectories of individual larvae of Japanese sardine, Sardinops melanostictus, caught in the coastal waters off western Japan were back-calculated from the first feeding stage up to date of capture (approximate size of 20 to 35 mm total length; TL) based on individually determined allometric relationships between otolith daily ring radii and fish total lengths. The larvae in January-, February-, and March-hatched cohorts in the coastal waters grew faster and more uniformly than those in the oceanic waters offshore of the Kuroshio current. Growth trajectories of the three hatch-month cohorts were similar and could be expressed by the Gompertz model. The inflection points of the growth curves were reached at 9 to 11 d after hatching, when larvae were 10.8 to 11.8 mm TL. Maximum growth rates at these points were 0.80 to 0.85 mm d⁻¹. Growth rates gradually declined after the inflection points, and larval TLs converged into the infinite length of 29 to 32 mm, the sizes at which metamorphosis from larvae to juveniles is initiated. This asymptotic growth pattern in the larval stage resulted in the narrow ranges in TLs in spite of the wide range of ages of the larvae caught by boat seiners in the coastal waters. Slow growth and therefore long duration of the metamorphosing stage could be influential in determining the cumulative total mortality in the early life stages of the Japanese sardine. Received: 14 July 1996 / Accepted: 20 August 1996     

Influence of cadmium accumulation and dietary status on fatty acid composition in two colour forms of shore crabs, Carcinus maenas

 The effects of cadmium exposure and dietary status on cadmium accumulation, fatty acid (FA) content and profiles were investigated in two colour forms of the shore crab Carcinus maenas. Groups of shore crabs were either starved or fed with blue mussels, Mytilus edulis, during a 40 d exposure period to 2 or 6 μM Cd²⁺ (as CdCl₂). Starved green individuals accumulated more cadmium in haemolymph and hepatopancreas than did red crabs and green crabs fed during the experiments. In the red colour form, no difference in cadmium accumulation was observed between starved and fed individuals. In both colour forms, hepatopancreas contained more FA than gills and muscle. The FAs often present in the largest amounts in the tissues were 16:0, 16:1ω7, 18:1ω7, 18:1ω9, 20:4ω6, 20:5ω3 and 22:6ω3. However, saturated (SAFA) and mono-unsaturated fatty acids (MUFAs) were dominant in hepatopancreas, whereas poly-unsaturated fatty acids (PUFAs) were dominant in gills and muscles. At the beginning of the experiment, the total FA content in the hepatopancreas was 111.6 mg g⁻¹ (dry weight) for red crabs and 78.4 mg g⁻¹ for green shore crabs. During the experiment, however, the FA content decreased in red crabs. This decrease was more pronounced for starved individuals than for fed individuals. Also, the decrease in FA content was more pronounced in crabs exposed to 6 μM cadmium compared to crabs exposed to 2 μM or crabs not exposed to cadmium. No change in FA content was observed in green shore crabs, irrespective of diet and cadmium exposure. For both colour forms, no change in FA content was observed for gills and muscle. In red crabs, a decrease was observed for all FAs in the hepatopancreas. This decrease, however, was more pronounced for SAFAs and MUFAs than for PUFAs, indicating that the metabolism of FAs during starvation and cadmium exposure is selective. The experiments indicate that green colour forms of shore crabs are more tolerant of natural stress such as starvation and anthropogenic stress, e.g. cadmium exposure, than are red colour forms of shore crabs. Received: 23 September 1999 / Accepted: 29 April 2000     

Foraging behavior in early Atlantic cod larvae (Gadus morhua) feeding on a protozoan (Balanionsp.) and a copepod nauplius (Pseudodiaptomussp.)

Food limitation is likely to be a source of mortality for fish larvae in the first few weeks after hatching. In the laboratory, we analyzed all aspects of foraging in cod larvae (Gadus morhua Linnaeus) from 5 to 20 d post-hatching using protozoa (Balanion sp.) and copepod nauplii (Pseudodiaptomus sp.) as prey. A camera acquisition system with two orthogonal cameras and a digital image analysis program was used to observe patterns of foraging. Digitization provided three-dimensional speeds, distances, and angles for each foraging event, and determined prey and fish larval head and tail positions. Larval cod swimming speeds, perception distances, angles, and volumes increased with larval fish size. Larval cod swam in a series of short intense bursts interspersed with slower gliding sequences. In 94% of all foraging events prey items were perceived during glides. Larval cod foraging has three possible outcomes: unsuccessful attacks, aborted attacks, and successful attacks. The percentage of successful attacks increased with fish size. In all larval fish size classes, successful attacks had smaller attack distances and faster attack speeds than unsuccessful attacks. Among prey items slowly swimming protozoans were the preferred food of first-feeding cod larvae; larger larvae had higher swimming speeds and captured larger, faster copepod nauplii. Protozoans may be an important prey item for first-feeding larvae providing essential resources for growth to a size at which copepod nauplii are captured. Received: 20 April 1999 / Accepted: 12 January 2000     

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     

Longevity of subitaneous and diapause eggs of Centropages hamatus (Copepoda: Calanoida) from the northern Gulf of Mexico

Copepod resting eggs are abundant in the seabed of many bays and estuaries where they provide a potential source of recruits for growth of planktonic populations. In the northeastern Gulf of Mexico the copepod Centropages hamatus (Lillejeborg) occurs in the water column only during the late fall, winter and early spring. The species produces subitaneous and diapause eggs, and both egg types have been found in the seabed. We determined the longevity of these two egg types to ascertain their potential for contributing to the growth of the planktonic population and for sustaining a persistent egg bank. Eggs were collected from females and incubated in the laboratory under temperature and oxygen conditions chosen to simulate field conditions. The diapause eggs were also exposed to sulfide. The total hatching success of subitaneous eggs in two experiments declined from highs of 78 and 97% to zero after 60 and 90 d of exposure to anoxia. The total hatching success of diapause eggs that were exposed to anoxia for 90 d however was typically greater than 80%. Some diapause eggs hatched after being incubated under anoxia for 437 d. Diapause eggs survived longer at ambient field temperatures when incubated under anoxia (437 d) compared to normoxia (118 d). Exposure to sulfide did not result in greater mortality of diapause eggs compared to anoxia alone. Diapause eggs that were incubated at ambient field temperatures did not hatch when exposed to normoxia until the temperature dropped to <20 °C. The results of this study suggest that C.␣hamatus sustain a short-term reserve of subitaneous eggs in the seabed that provides recruits for the current year's population. The greater longevity of diapause eggs suggests that they sustain the seasonal reappearance of the species year after year in the northeastern Gulf of Mexico. However, the contribution of diapause eggs of C. hamatus from the Gulf of Mexico to a persistent egg bank is questionable since hatching ceased after 437 d. Received: 30 July 1997 / Accepted: 18 January 1998     

Time-course of digestive-enzyme acclimation in the cockle Cerastoderma edule

We determined the temporal evolution of amylase, cellulase, laminarinase and protease in the digestive gland and crystalline style of cockles Cerastoderma edule held over 9 to 12 d in the presence and absence of food. Cockles were fed a constant diet of 1.5 mm³ l⁻¹ of Tetraselmis suecica for 9 to 12 d and were then starved for 6 to 8 d in late summer (September 1992) and in winter (January 1993). Feeding increased the dry weight and total cellulase, laminarinase and protease activities of the digestive gland irrespective of season, whereas amylase activity remained unchanged. In winter (i.e. when cockles are metabolically weak) the response was faster and stronger, especially for protease. An additional experiment in September starved cockles for 20 d before resuming feeding. In agreement with the seasonal differences, the presence of food after prolonged starvation induced a rapid and marked increase in protease in the digestive gland of the cockles. In winter, the possible effects of the biochemical composition of food on their enzymatic response were tested by feeding two groups of cockles with the same ration of T. suecica but harvested at different growth phases. A compensatory induction of cellulases occurred in cockles fed on T. suecica with a lower carbohydrate content. In the crystalline style, the protein level and carbohydrase fell during the first day of feeding and increased during the first day of subsequent starvation. These results indicate that the release of enzymes from the style prevails over the incorporation of enzymes during the early stages of feeding, whereas the opposite occurs during starvation. Received: 15 February 1998 / Accepted: 22 February 1999     

Effects of limiting access to prey on development of first zoeal stage of the brachyuran crabs Cancer magister and Hemigrapsus oregonensis

We tested the influence of limiting access to prey on larval development of the crabs Cancer magister and Hemigrapsus oregonensis by raising their Stage 1 larvae in the laboratory on different prey densities and with various periods of access to prey. Experiments were conducted in 1995 and 1996 at the Shannon Point Marine Center in Anacortes, Washington, USA. Our results show that crab larvae do not require continuous access to prey for optimal development nor do they appear to require light for prey capture. Survival and duration of Stage 1 C. magister fed continuously on only one-fourth the amount of the control density of prey and those fed at the control density for only 6 h per day were the same as for larvae fed continuously at the control density (20 ml⁻¹). Larvae with cyclic access to prey at the control density for 24 h and then starved for 72 h showed significantly lower survival and longer instar duration to Stage 2. Experiments on Stage 1 H. oregonensis which investigated a combination of prey density, period of access to prey and light/dark conditions during feeding revealed that survival decreased with decreasing prey density or with decreasing feeding period, but no differences were observed during periods of limited prey availability as a function of light or dark conditions. Stage duration was not affected by reduced prey density nor by the light/dark condition at the time of feeding, but it was prolonged when the period of access to prey was limited. The period of access to prey did not affect the weight of Day 1 Stage 2 larvae. Larvae fed high densities of prey for 4 h followed by 20 h of reduced-density diet exhibited the same survival and stage duration as controls that were continuously fed high-density prey. Our results define sub-optimal diets that can be used experimentally to determine the nutritional contributions made by naturally-occurring prey organisms during larval development in the two species. In nature, larvae may satisfy nutritional requirements through periodic encounters with dense prey patches during vertical migrations by day or night. Received: 12 August 1997 / Accepted: 5 February 1998     

Effects of temperature and delayed feeding on growth and survival of larvae of three species of subtropical marine fishes

In larvae of the bay anchovy Anchoa mitchilli (Valenciennes), the sea bream Archosargus rhomboidalis (Linnaeus), and the lined sole Achirus lineatus (Linnaeus), growth, survival, and starvation times were investigated at temperatures of 22° to 32°C. The rate at which hours after hatching until starvation decreased in relation to temperature for unfed larvae did not differ significantly among the 3 species, ranging from-5.4 to-6.3 h per degree increase in temperature. The total number of hours until starvation did differ for all 3 species: lined soles survived longest, bay anchovies were intermediate, and sea bream survived the least time. At 28°C, unfed sea bream could survive 90.1 h, bay anchovy 102.3 h, and lined sole 119.8 h. The eyes pigmented at nearly the same time after hatching for sea bream and bay anchovy, but took about 20 h longer at all temperatures for lined sole. Quadratic equations best described the relationship between hours after hatching when the eyes became pigmented and temperature. Eye-pigmentation times became nearly constant for all 3 species at temperatures above 28°C. At 28°C, eyes pigmented about 27 h after hatching for bay anchovy and sea bream but not until 47 h for lined sole. Hours after eye pigmentation when unfed larvae starved was a measure of the effective time that larvae had to commence feeding. Bay anchovies and lined soles were nearly alike in this respect, but sea bream starved at tewer hours after eye pigmentation. Slopes of regressions representing decrease in times to staration for increasing temperatures ranged from-3.7 to-4.4 h per degree increase in temperature, and were not significantly different among the 3 species. At 28°C, unfed lined soles starved at 70 a after eye pigmentation, bay anchovies starved at 72.5 h, and sea bream at only 62 h. Yolk absorption was most rapid for all species during the first 20 h after hatching, and was faster at higher temperatures. Amounts of yolk remaining at the time eyes became pigmented were less at higher temperatures for bay anchovy and lined sole, but were greater for sea bream, suggesting that sea bream used yolk more efficiently at higher temperatures. Either no yolk or small traces (>0.20%) remained at 24 h after eye pigmentation in all 3 species. Feeding was delayed for periods of 8, 16, 24, 32, 40 and 48 h after eye pigmentation for all species at a series of experimental temperatures from 24° to 32°C. Growth and survival were affected when food was withheld for more than 24 h at 28°C, but survival did not decrease markedly until food was withheld at least 8 h longer. At lower temperatures food could be withheld longer and at higher temperatures for less time. Feeding can be initiated by most larvae for several hours after all visible yolk reserves have been exhausted. All species tested can survive for 24 to 40 h after eye pigmentation at 24° to 28°C without food and still have relatively good growth and survival when food is offered. If the “critical period” is considered relative to time of hatching, lined soles need not find food for 3 to 3.5 days after hatching, but bay anchovy and sea bream must feed within 2.5 days of hatching.     

Biological weighting of ultraviolet (280–400 nm) induced mortality in marine zooplankton and fish. I. Atlantic cod (Gadus morhua) eggs

In the Gulf of St. Lawrence, Canada, productivity-determining biophysical interactions occur in the upper 0 to 30 m of the water column. The eggs and larvae of several commercially important marine invertebrates and fishes (e.g. Gadus morhua L.) are found in this layer. Measurements of the diffuse attenuation coefficients for ultraviolet-B radiation (280 to 320 nm, UV-B) at various locations in this geographic region indicated maximum 10% depths (the depth to which 10% of the surface energy penetrates at a given wavelength) of 3 to 4 m at a wavelength of 310 nm. This represents a significant percentage of the summer mixed-layer water column: organisms residing in this layer are exposed to UV-B radiation. Laboratory experiments using a Xenon-arc-lamp based solar simulator revealed that cod embryos exposed to UV-B exhibited high wavelength-dependent mortality. The strongest effects occurred under exposures to wavelengths below 312 nm. This susceptibility was also dependent upon developmental stage; mortality was particularly high during gastrulation. At the shorter wavelengths (<305 nm) UV-B-induced mortality was strongly dose-dependent, and not significantly influenced by dose-rate. The biological weighting function (BWF) derived for UV-B-induced mortality in cod eggs is similar to that reported for naked DNA – suggesting that the mortality is a direct result of DNA damage. There was no evidence of a detrimental effect of ultraviolet-A radiation (320 to 400 nm). Calculations based upon the BWF indicate that, under current noon surface irradiance, 50% of cod eggs located at or very near (within 10 cm) the ocean surface will be dead after 42 h of exposure. Under solar spectral irradiance simulating a 20% decrease in ozone layer thickness, this time drops to 32 h. These are first-order estimates based upon surface irradiance taken at a time of day during which the values would be maximal. Nonetheless, they illustrate the relative changes in UV-B impacts that will result from ozone layer depletions expected over the coming decades. It is also important to point out that variability in cloud cover, water quality, and vertical distribution and displacement of cod eggs and larvae within the mixed layer, can all have a greater effect on the flux of UV-B radiation to which fish eggs are exposed than will ozone layer depletion at these latitudes. Received: 2 March 1998 / Accepted: 18 December 1998