Production, Quality, and Low Temperature Incubation of Eggs of Atlantic Cod Gadus morhua and Haddock Melanogrammus aeglefinus in Captivity

Atlantic cod Gadus morhua and haddock Melanogrammus aeglefinus broodstock maintained under altered regimens of temperature and photoperiod spawned up to 8 mo per year. The cod broodstock produced viable embryos from October through June. The haddock broodstock produced viable embryos from December through May. Egg diameters were largest during the middle of the spawning season when water temperature was at a minimum, resulting in an inverse relationship between egg diameter and water temperature in both species. Egg quality was high, as evaluated by buoyancy, fertilization rate, regularity of early cleavage, and percent viable hatch. Low temperature incubation of cod and haddock eggs extended the embryonic period. Cod embryos tolerated a wider range of temperatures than haddock. High mortality (1 90%) was observed before hatching in haddock embryos incubated at 1 C. Atlantic cod embryos hatched at temperatures as low as —1 C, extending the embryonic period to 59 d. At 8 C Atlantic cod and haddock embryos hatched in 11–12 d. To determine if extending the embryo incubation time by using low temperatures had a detrimental effect, embryos were incubated through hatch at either 1 C or 6 C, and the larvae from both groups reared at 6 C. Growth and early survival of larvae were comparable in both treatments.     

Optimum temperature and food-limited growth of larval Atlantic cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) on Georges Bank

We estimated recent growth of Atlantic cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) larvae collected on the southern flank of Georges Bank in May 1992–94 from the ratio of RNA to DNA (R/D) and water temperature. Growth of both species increased with water temperature to about 7°C and then decreased. The highest growth rates were observed in May 1993 at water temperatures around 7°C. These data confirm an earlier observation of comparable temperature optima for growth of Atlantic cod and haddock larvae in the north‐west Atlantic. Comparisons of field growth rates and temperature optima with data for larvae cultured at high temperatures and prey densities in the laboratory suggest that growth may have been food‐limited at higher temperatures on Georges Bank. Given that 7°C is the long‐term mean water temperature on the southern flank in May and that climate models predict a possible 2–4°C rise in water temperatures for the western North Atlantic, our findings point to a possible adverse effect of global warming on Atlantic cod and haddock.     

Proteome profile comparison of two differently fed groups of Atlantic cod (Gadus morhua) larvae

Proteome analysis was used to study the effects of feeding early Atlantic cod (Gadus morhua) larvae with a saithe (Pollachius virens) protein hydrolysate (SPH). Protein hydrolysates have previously been shown to beneficially affect fish larval development. Feeding was initiated on day 2 post hatch (ph) or as soon as the larvae opened their mouth and the protein expression was monitored 4 days later or in 6‐dph cod larvae. The results demonstrated changes in the abundance of 13 protein spots in the cod larvae fed SPH. Of these, seven protein spots were up‐regulated and six protein spots showed down‐regulation. Five of the up‐regulated proteins in cod larvae are known to be involved in energy metabolism. A few early larval specific proteins were down‐regulated in the SPH‐fed cod larvae possibly because of an enhanced development in this group relative to the control group. Two trypsin isoforms were detected within the cod larval proteome. The detection of the trypsin spots was made possible by co‐electrophoresis of known cod trypsins with the cod larval protein extract. Surprisingly, no difference in trypsin content was observed between the SPH‐fed and the control larval groups.     

Efficacy of egg surface disinfectants in captive spawning Atlantic cod Gadus morhua L. and haddock Melanogrammus aeglefinus L.

In an effort to optimize the efficiency of high‐density incubation of Atlantic cod Gadus morhua and haddock Melanogrammus aeglefinus eggs, the per cent hatch of eggs treated with four disinfectants (3% hydrogen peroxide, 1% polyvinylpyrrolidone iodine, 0.1% sodium hypochlorite and a 0.005% antibiotic solution – penicillin/streptomycin) was compared in both species. The per cent hatch of eggs of each species was greatest after a 24 h treatment with the antibiotic solution. The hatching success of eggs treated within the different disinfectant treatments depended upon the embryonic developmental stage in both species. The sodium hypochlorite treatment had the lowest % coverage of colony growth after disinfected haddock eggs were plated onto sterile agar media, highest survival rates to the end of the embryonic period, but the lowest per cent hatch.     

Rotifers enriched with iodine,copper and manganese had no effect on larval cod (Gadus morhua) growth,mineral status or redox system gene mRNA levels

It currently remains unclear if rotifers contain sufficient mineral levels to meet larval fish requirements. In this study, rotifers were enriched with a commercial enrichment (control), or with additional iodine, iodine and copper, or iodine, copper and manganese, and the effects of feeding these rotifers to Atlantic cod (Gadus morhua) larvae from 3 to 18 days post hatch were investigated. Rotifer enrichment with minerals was successful, but Mn enrichment also increased rotifer zinc levels. No differences were observed between treatments in larval growth or survival, or in the mRNA levels in the majority of the redox system genes analysed. Only Zn levels increased in cod larvae in response to mineral enrichment of rotifers. Apart from Zn, little evidence was found to suggest that cod larvae require increased concentrations above the control rotifer levels of the essential elements studied here.     

Improved performance of Atlantic cod (Gadus morhua L.) larvae following enhancement of live feed using a fish protein hydrolysate

High larval mortalities and anatomical deformities are among the major obstacles restricting the development of Atlantic cod (Gadus morhua) aquaculture. The immune system of cod larvae is poorly developed at hatch, and innate immune parameters are therefore of importance for defence against environmental microorganisms. Two separate experiments were conducted with bioencapsulation of the live feed of cod larvae using a pollock (Pollachius virens) protein hydrolysate. Offering peptide enhanced live feed to larvae during the first weeks of exogenous feeding promoted larval development, with reduced incidence of severe deformities to 3.0% as compared with 9.6% deformities observed in the control group at 160 days posthatch. The production and distribution of IgM and lysozyme were furthermore increased in larvae fed peptide enhanced feed compared with control larvae. IgM was predominantly detected in the foregut and the epithelial lining of the digestive tract as well as in the epidermal mucus of the skin. Lysozyme was mainly detected in the epidermal mucus of the skin and in the foregut. Overall, the results indicate that live feed enhancement using a protein hydrolysate derived from pollock may reduce deformities and promote normal development during early production stages of cod larvae.     

The settlement and reproductive success of Lepeophtheirus salmonis (Krøyer 1837; Copepoda: Caligidae) on atypical hosts

The reproductive success of Lepeophtheirus salmonis settled on host and non‐host fish has been compared. Triplicate single species tanks of Atlantic salmon, marine three‐spined sticklebacks, saithe and Atlantic cod were exposed to 10 adult female L. salmonis per tank (n=30 lice per species). Adult female L. salmonis settlement and egg string production occurred only on salmon and cod, with no egg production occurring on saithe and three‐spined sticklebacks. The number of eggs in egg strings, hatching success of eggs and the survival of all larval stages to the copepodid stage were severely affected by the species of fish on which female L. salmonis had settled. L. salmonis settled on cod produced significantly fewer eggs, lower hatching rates and lower survival rates of larvae than females on Atlantic salmon. The production of egg strings by L. salmonis females infecting cod, which successfully hatch and moult through to the infective copepodid stage, albeit in small numbers, is discussed in terms of the implications to aquaculture and salmon and cod farming scenarios.     

The growth of larval cod and haddock in the Irish Sea: a model with temperature,prey size and turbulence forcing

We applied a physiological individual‐based model for the foraging and growth of cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) larvae, using observed temperature and prey fields data from the Irish Sea, collected during the 2006 spawning season. We used the model to estimate larval growth and survival and explore the different productivities of the cod and haddock stocks encountered in the Irish Sea. The larvae of both species showed similar responses to changes in environmental conditions (temperature, wind, prey availability, daylight hours) and better survival was predicted in the western Irish Sea, covering the spawning ground for haddock and about half of that for cod. Larval growth was predicted to be mostly prey‐limited, but exploration of stock recruitment data suggests that other factors are important to ensure successful recruitment. We suggest that the presence of a cyclonic gyre in the western Irish Sea, influencing the retention and/or dispersal of larvae from their spawning grounds, and the increasing abundance of clupeids adding predatory pressure on the eggs and larvae; both may play a key role. These two processes deserve more attention if we want to understand the mechanisms behind the recruitment of cod and haddock in the Irish Sea. For the ecosystem‐based management approach, there is a need to achieve a greater understanding of the interactions between species on the scale a fish stock is managed, and to work toward integrated fisheries management in particular when considering the effects of advection from spawning grounds and prey–predator reversal on the recovery of depleted stocks.     

Vertebral deformity in cultured Atlantic cod larvae: ontogeny and effects on mortality

The effects of different egg incubation densities on the incidences of vertebral deformities in Atlantic cod larvae were investigated. Cod eggs were incubated at four different densities, 3, 6, 12 and 48 mL eggs L⁻¹, of water.  When all the eggs hatched, larvae were reared in 30 L glass aquaria. Larval samples were taken at 0, 14, 42 and 56 days post hatch (dph) for deformity analysis. Larval samples were stained using bone and cartilage staining methods to determine vertebral deformity. Incubation densities did not have any significant effects on vertebral deformities in Atlantic cod larvae. However, the incidence of larval vertebral deformity was high at hatch and decreased as the larvae grew older until 42 dph, indicating selective mortality of deformed larvae during this period. Larvae at 56 dph, however, showed an increase in the incidence of vertebral deformity, indicating a possible nutritional or prey-type effect. To our knowledge, no studies have documented the occurrence of variable patterns in vertebral deformities in cod at various developmental stages. Overall, our results suggest that broodstock husbandry, genetics and/or nutrition could play a major role in causing vertebral deformities in Atlantic cod at hatch; however, nutrition and prey type may play a major role during metamorphosis.     

Effects of different step‐wise temperature increment regimes during egg incubation of Atlantic cod (Gadus morhua L.) on egg viability and newly hatched larval quality

We carried out an experiment to determine how rapidly the early incubation temperature of Atlantic cod eggs can be increased without affecting normal embryonic development and hatching. Atlantic cod eggs were incubated at a constant low temperature (4.5 ± 0.5°C; T5 – control) and four temperature increment treatments where the temperatures were increased stepwise from 4.5°C at zygote stage to 9.5 ± 05°C (T1‐8 h, T2‐32 h, T3‐64 h and T4‐96 h). Embryonic cell symmetry, embryonic mortality, hatching success and larval skeletal abnormalities, length and yolk sac volume were recorded. Larval samples were also taken at hatch for histological analysis. Except for higher egg mortality and lower hatching success in the T1, the differences among experimental groups were minor. Cell asymmetries and embryo mortalities were not significantly different between the control and T2–T4 treatment groups. Control larvae were significantly longer and had smaller yolk reserves at hatch than T1–T4 larvae and larvae from T2 had the largest yolk reserves. Tissue and organ histology of hatched larvae were similar. Considering embryonic cleavage pattern, hatching success and larval morphology and histology, a gradual increment of temperature in 32 h seems to be the better choice for future developmental programming studies in Atlantic cod.     

Larval group differentiation in Atlantic cod (Gadus morhua) inside and outside the Gullmar Fjord

The spatial and temporal occurrence of pelagic fish stages and their biological variability may affect their dispersal and survival, and ultimately fish recruitment. We collected Atlantic cod larvae at one station inside and at one station outside the Gullmar Fjord, eastern Skagerrak, in order to investigate small-scale larval group differentiation. Rectangular midwater trawl hauls were taken every 6 h (during 24 h) from three separate depth intervals between the surface and 70 m depth. About 80% and 20% of all larvae occurred above the halocline at the Fjord station and the Coastal station, respectively. Hatching (at both stations) occurred from the 3rd week in February to the 1st week in May, indicating that cod larvae were present for at least 5 months (from late February to early August). The length and hatch date frequency distributions of larvae from the surface layer were unimodal inside the fjord but bimodal outside the fjord. Analyses of seven microsatellite DNA loci indicated that larvae collected inside the fjord (where local spawning occurs) were genetically distinct from larvae sampled on the outside (F_ST = 0.0026). The two age cohorts outside the fjord were not, however, genetically different, nor were larvae collected at different depths. We conclude that small-scale variability of vertical concentration and larval life history variability should have consequences for interpreting models of larval dispersal and survival, and subpopulation structure analyses.     

The effect of different HUFA enrichment emulsions on the nutritional value of rotifers (Brachionus plicatilis) fed to larval haddock (Melanogrammus aeglefinus).

A feeding study was conducted in the winter 2001 to determine the effects of feeding rotifers (Brachionus plicatilis) enriched with various levels of essential fatty acids on the growth and survival of haddock larvae (Melanogrammus aeglefinus). Rotifer enrichment treatments were: 1) mixed algae, 2) high DHA (docosahexaenoic acid, 22:6n-3), 3) high DHA and EPA (eicosapentaenoic acid, 20:5n-3), and 4) DHA, EPA, and AA (arachidonic acid, 20:4n-6). Larvae were fed rotifers enriched with the different treatments from days 1 to 16 post-hatch. From day 17 until 25 all treatment groups were fed rotifers reared on mixed algae and then weaned onto the International Council for Exploration of the SEA (ICES) Standard Reference Weaning diet (http://allserv.rug.ac.t/aquaculture/rend/rend.htm) over a five day period. The experiment was terminated on day 41 post-hatch. The enrichment treatments affected the fatty acid composition of the rotifers and correlated with the accumulation of these fatty acids in the haddock larvae. However, no significant differences in larval growth or survival to 40 days post hatch were detected, suggesting that all treatments provided the minimal essential fatty acid requirements for haddock.     

A synthesis of large-scale patterns in the planktonic prey of larval and juvenile cod (Gadus morhua)

Data from 40 published studies of the diet composition of larval and juvenile cod (Gadus morhua) from around the northern North Atlantic were summarized to assess generic patterns in ontogenetic and regional variability in the key prey. The results showed that larvae at the northern edge of the latitudinal range of cod depend primarily on development stages of the copepod Calanus finmarchicus, whilst those at the southern edge depend on Para‐ and Pseudocalanus species. Juvenile cod preyed on a wider range of taxa than larvae, but euphausiids were the main target prey. Analysis of regional variations in the relative abundances of C. finmarchicus and Para/Pseudocalanus spp. in the plankton, as estimated by the continuous plankton recorder (CPR) surveys, showed a similar geographical pattern to the larval cod stomach contents. Comparison of CPR data from the 1960s and 70s with data from the 1990s showed that the boundary between C. finmarchicus and Para/Pseudocalanus spp. dominance has shifted northwards on both sides of the Atlantic, whilst the abundance of euphausiids in the southern cod stock regions has declined. The results are discussed in relation to regional differences in the response of cod stocks to climate variability.     

Selective haddock (Melanogrammus aeglefinus) trawling: Avoiding cod (Gadus morhua) bycatch

The critical condition of the North Sea cod stocks has resulted in restrictions on not only cod, but also haddock and other species that are caught together with cod. Thus full exploitation of the haddock stock is unachievable unless cod can be excluded from the haddock catch. We designed a selective trawl based on the behavioral differences between haddock and cod as they enter a trawl, i.e., cod stay close to the seabed whereas haddock rise above it. The trawl's fishing line is raised ～60 cm above the seabed to allow cod to escape beneath the trawl while haddock are retained. To collect the escapees, three sampling bags were attached beneath the raised fishing line. The selective haddock trawl reduced the total catch of cod by 55% during the day and 82% at night, and 99% of the marketable haddock was caught during the day and 89% at night. Cod escape rates were highly length dependent: smaller cod escaped the trawl in greater numbers than did larger individuals. Whiting, saithe, lemon sole, and plaice were included in the analysis.     

Behavioural responses to visual environment in early stage Atlantic cod Gadus morhua L. larvae

Marine fish larvae exhibit species‐specific sensitivity to light, which changes during ontogeny. Our aim was to explore how visual environment, characterized by light intensity, microalgae addition and wall colour, affects spatial distribution of Atlantic cod larvae. We applied two different small‐scale factorial designs, each of them as a series of independent short‐term trials at fixed larval ages from 5 to 35 days post hatch (DPH). Positive phototaxis increased in larvae from 5 to ≈24 DPH, followed by reduced response at later ages. Effects from the experimental factors on larval behaviour changed with age, and significant interactions between factors were present. Algae significantly affected larvae only at 5 DPH, increasing phototactic response. White wall colour induced distribution of larvae close to the tank walls at all ages. The experimental approach demonstrated clear effects from visual environment on larval distribution, and revealed both relative importance of individual factors, interactive effects and temporal changes due to larval ontogeny. Our results suggest that by fine‐tuning the visual environment in tanks, the larval distribution may be managed to improve overall rearing conditions through a more even distribution with reduced closeness to tank walls.     

Growth and lipid composition of Atlantic cod (Gadus morhua) larvae in response to differently enriched Artemia franciscana

Considerable progress has been achieved in the intensive culture of Atlantic cod (Gadus morhua). However, there is little information concerning optimum live-feed enrichments for cod larvae, since many of the techniques used during the larviculture have been borrowed from other fish species and adapted for the production of Atlantic cod. The present study compared four different protocols for the enrichment of Artemia to be used as live feed for cod larvae. The protocols tested were: (1) AlgaMac 2000, (2) AquaGrow Advantage, (3) Pavlova sp. + AlgaMac 2000, and (4) DC DHA Selco + AlgaMac 2000. Larvae were fed differently enriched Artemia between 37 and 59 days post hatch. At the end of the experiment, larvae from treatment 1 [specific growth rate (SGR) = 10.4 ± 0.4% day⁻¹] grew faster than larvae from treatments 3 (SGR = 6.9 ± 0.2% day⁻¹) and 4 (SGR = 4.9 ± 0.4% day⁻¹, P < 0.0001). However, treatments 3 and 4 resulted in better larval survival at the end of the experimental period, estimated to be 3 on a scale from 1 to 5, whereas the survival estimates for the two other groups were 2. The treatments affected the fatty-acid composition of Artemia and of cod larvae. Larvae from treatment 1 had a higher percentage of AA (20:4ω6, P < 0.0001) and ω6DPA (22:5ω6, P < 0.0001) than the other larvae. Levels of DHA (22:6ω3) were similar in larvae from treatments 1 and 4, and higher than in the other larvae (P < 0.0001). Our results suggest that Artemia containing a DHA/EPA/AA ratio of 7/2/1 result in good larval performance. Joseph A. Brown—Deceased September 2005.     

A method to determine protein digestibility of microdiets for larval and early juvenile fish

A method to evaluate protein quality using in vivo methods was developed for larval fish. FluoSpheres^® fluorescent microspheres (10 μm) were incorporated into two test diets, our standard zein microdiet (ZMD) and a microdiet with identical ingredients except for the replacement of high quality fish meal with the same product cooked for 24 h at 80 °C (ZMD-CF). Several trials were performed to design a reliable method to test digestibility using FluoSpheres^® as a marker. The developed in vivo technique was tested on 35 days posthatch (dph) larval Atlantic cod ( Gadus morhua L.) and two tropical fish species in the early juvenile stage. The method took into account loss of total protein to the faecal pellet and water column. Apparent digestibility of protein in larval cod fed ZMD was significantly higher than that of larvae fed ZMD-CF ( P  < 0.05). A growth study to validate differences between the two diets showed significant differences in growth and survival of larvae fed ZMD versus ZMD-CF ( P  < 0.05). Further validation of our results was indicated through the use of a pH-stat method using enzymes extracted from 35 dph larval cod guts. This novel technique will be advantageous for researchers to evaluate feed ingredients for larval marine fish and is adaptable to many different areas of larval fish nutrition.     

Effect of acclimation temperature on the acute stress response in juvenile Atlantic cod, Gadus morhua L., and haddock, Melanogrammus aeglefinus L.

Juvenile Atlantic cod (Gadus morhua L.) and haddock (Melanogrammus aeglefinus L.) were subjected to 30 s air exposure stressors following acclimation to 4, 10 and 14 and 4, 8 and 14°C respectively. Both species responded to the stressor with increases in plasma cortisol at all temperatures tested. At 14°C cortisol levels peaked within 1 h post‐stressor, and returned to pre‐stressor levels within 24 h. In contrast, at 4°C, peak cortisol levels were not attained until 6 h post‐stressor in haddock and remained elevated beyond 24 h in both species. The rate of plasma glucose accumulation was greater at higher temperatures in both species and no increase was seen at 4°C. Lysozyme activity in cod, in response to the stressor, was lower than the values reported for some other species and increased slightly at 14°C. The results show that cod and haddock acclimated to different temperatures respond to common, acute stressors in a manner similar to other teleosts.