Technological solutions and operational measures to prevent escapes of Atlantic cod (Gadus morhua) from sea cages

Escapes of cod (Gadus morhua) from sea cages represent an economic problem for farmers and a potential environmental problem. We estimate that 0–6% of cod held in sea‐cage farms in Norway were reported to have escaped each year from 2000 to 2005, which is a high proportion compared with salmon. We interviewed employees at 19 coastal sea‐cage cod farms in Norway to investigate both how and why cultured cod escape and to document cage handling and management strategies that were effective in minimizing escapes. Based on the interviews, we describe five working hypotheses that may explain why a greater proportion of cod than salmon escape: (1) cod are more willing to escape than salmon; (2) cod bite the net cage and create wear and tear; (3) net cages have insufficient technical standards for cod culture; (4) cod are placed in sea cages at considerably smaller sizes than salmon; and (5) cod are more popular feed for predators. Preliminary testing of the hypothesis that cod bite netting and create holes was done by placing pre‐damaged net panels with cut twines and control panels inside sea cages. Holes in the pre‐damaged net panels increased in size over a period of 3 months. The type of damage indicated that biting of netting twines was the likely cause.     

Movements and spatiotemporal distribution of escaped farmed and local wild Atlantic cod (Gadus morhua L.)

Commercial farming of Atlantic cod (Gadus morhua L.) is now being developed in several countries. The ecological consequences of cod culture are poorly understood, but recent research suggests that Atlantic cod are more prone to escape from net pens than Atlantic salmon. Here, we describe the movements and the spatiotemporal distribution of farmed cod after escape relative to wild cod, both during and outside the natural spawning season. The experimental design included simulating escape incidents of farmed cod tagged with acoustic transmitters and using an array of automatic listening stations to monitor their dispersal and distribution. For comparison, local wild cod were monitored using the same array of receivers. The farmed cod dispersed rapidly after a simulated escape, they randomly distributed over large areas and their distribution overlapped with local wild cod. Moreover, escaped farmed fish were found at local cod spawning areas during the spawning season. The study also indicated that the recapture rate of escaped farmed cod was high compared with that of escaped farmed salmon. Thus, while our results showed that there is a considerable potential for ecosystem effects caused by escaped farmed cod, mitigating actions such as an efficient recapture fishery for escapees may be possible.     

Occurrence of Francisella piscicida in farmed and wild Atlantic cod, Gadus morhua L., in Norway

Francisellosis, caused by the bacterium Francisella piscicida , has become one of the most serious diseases in Atlantic cod production in Norway. The major aim of this study was to determine the distribution of F. piscicida in farmed and wild fish in areas with cod farming along the Norwegian coast, and its occurrence in cod from areas without cod farming. Two real-time PCR assays, targeting the 16S rRNA gene and the FopA gene of F. piscicida , were developed since sensitive and specific diagnostic tools are required for detecting asymptomatic carriers of the bacterium. A total of 422 wild cod from 13 sampling areas and 955 farmed cod from 10 areas along the coast of Norway were examined. Using the real-time polymerase chain reaction (PCR) assays, F. piscicida was detected in wild populations of cod from all counties examined south of Sogn og Fjordane in southern Norway (overall prevalence 13%, n  = 221). Wild cod north of Sogn og Fjordane were negative for the bacterium ( n  = 201). Farmed cod from most parts of Norway were F. piscicida positive. The apparent absence of the bacterium in wild populations of cod in the northern parts of Norway and its widespread occurrence in wild cod from southern parts of Norway is believed to relate to differences in seawater temperatures.     

Yersiniosis in Atlantic cod,Gadus morhua (L.), characterization of the infective strain and host reactions

A disease outbreak in farmed Atlantic cod caused by Yersinia ruckeri is reported. Mortality started following vaccination of cod reared in two tanks (A and B). The accumulated mortality reached 1.9% in A and 4.8% in B in the following 30 days when treatment with oxytetracycline was applied. Biochemical and molecular analysis of Y. ruckeri isolates from the cod and other fish species from fresh and marine waters in Iceland revealed a high salinity‐tolerant subgroup of Y. ruckeri serotype O1. Infected fish showed clinical signs comparable with those of Y. ruckeri ‐infected salmonids, with the exception of granuloma formations in infected cod tissues, which is a known response of cod to bacterial infections. Immunohistological examination showed Y. ruckeri antigens in the core of granulomas and the involvement of immune parameters that indicates a strong association between complement and lysozyme killing of bacteria. Experimental infection of cod with a cod isolate induced disease, and the calculated LD₅₀ was 1.7 × 10⁴ CFU per fish. The results suggest that yersiniosis can be spread between populations of freshwater and marine fish. Treatment of infected cod with antibiotic did not eliminate the infection, which can be explained by the immune response of cod producing prolonged granulomatous infection.     

Muscle growth of triploid Atlantic cod juveniles (Gadus morhua)

This study has investigated the muscle growth of diploid and triploid Atlantic cod (Gadus morhua) juveniles raised in replicate tanks over a period of 29 weeks and analysed at three sampling points (February, June and September). Data for weight, length, condition factor (K), muscle fibre growth and myogenic progenitor cells (MPCs) number were collected and results were analysed in relation to body growth and ploidy status. Diploids were significantly heavier than triploids throughout the trial (~10–20%) and had K in June and September samplings. Over the whole period, the rate of muscle fibres' recruitment was 318 fibres per day and 252 fibres per day for diploid and triploid cod respectively. The larger body weight of diploids resulted in a total number of fast fibre number of 114 979 compared to 91 086 in triploids. The average diameter of the 2.5% of the smallest fibres (2.5th percentile) was higher in diploids than triploids at the start of the trial, with a reversed picture for the average of the upper 2.5% (97.5th percentile) at the end of the trial. The probability density function of the estimated muscle fibre diameters showed similar fibre size distribution between size‐matched diploids and triploids at all sample points. The peak fibre diameter was approximately 25 μm in February and increased to approximately 50 μm in June and September, irrespectively of ploidy. Pax 7 were used as molecular markers for MPCs. A positive correlation between Pax 7⁺ cells and total body length was observed only among triploid fish at the onset of the experiment.     

Modeling the effects of temperature on the survival and growth of North Sea cod (Gadus morhua) through the first year of life

Temperature and body size are widely agreed to be the primary factors influencing vital rates (e.g., growth, mortality) in marine fishes. We created a biophysical individual‐based model which included the effects of body size and temperature on development, growth and mortality rates of eggs, larvae and juveniles of Atlantic cod (Gadus morhua L.) in the North Sea. Temperature‐dependent mortality rates in our model were based on the consumption rate of predators of cod early‐life stages. The model predicted 35%, 53% and 12% of the total mortality to occur during the egg, larval and juvenile stages, respectively. A comparison of modeled and observed body size suggested that the growth of survivors through their first year of life is high and close to the growth rates in ad libitum feeding laboratory experiments. Furthermore, our model indicates that experiencing warmer temperatures during early life only benefits young cod (or theoretically any organism) if a high ratio exists between the temperature coefficients for the rate of growth and the rate of mortality. During the egg stage of cod, any benefit of developing more rapidly at warmer temperatures is largely counteracted by temperature‐dependent increases in predation pressure. In contrast, juvenile (age‐0) cod experiences a higher cumulative mortality at warmer temperatures in the North Sea. Thus, our study adds a new aspect to the ‘growth–survival’ hypothesis: faster growth is not always profitable for early‐life stages particularly if it is caused by warmer temperatures.     

Changes in selected stress and immune‐related genes in Atlantic cod,Gadus morhua,following overcrowding

Using Atlantic cod, Gadus morhua, as a model for the stress response in gadoid fish, the changes in the expression of some stress and immune genes as well as the profiles of plasma cortisol were examined. Adult fish were kept at a density of ca. 100 kg m^?3 by lowering the water level in the rearing tank for 1 h and this short‐term crowding stress was repeated thrice over a 12‐h interval period. Blood samples were collected before exposure and at 2, 24 and 72 h post crowding. Plasma cortisol level significantly increased at 2 h post crowding but returned to pre‐crowding levels 24 h after exposure. The relative expression of the stress response genes, glucose transporter‐3 and a putative heat shock protein 70 significantly increased at 2 and 24 h post crowding respectively. Significant up‐regulation of the pro‐inflammatory cytokines, interleukin (IL)‐1β and IL‐8, as well as anti‐bacterial genes, g‐type lysozyme and bactericidal permeability‐increasing protein/lipopolysaccharide‐binding protein (BPI/LBP) was also observed at 2 h and the levels were maintained until 72 h post exposure, except for BPI/LBP which had maximum up‐regulation at 24 h. The present observations have implications with respect to fish welfare and assessment of the health status of the farmed fish.     

Effect of feed composition and feeding frequency on growth, feed utilization and nutrient retention in juvenile Atlantic cod, Gadus morhua L.

Juvenile Atlantic cod (Gadus morhua) were fed extruded feeds formulated to contain 360–660 g kg^?1 protein, 80–280 g kg^?1 lipid and 80–180 g kg^?1 starch at feeding frequencies of either once per day or every second day to satiation. The trial was conducted at 8 °C and lasted for 28 weeks during which fish were weighed five times at regular intervals. Sampling for proximate analysis was performed at the start, after 12 weeks and at the end of the trial. Fish grew from an average weight of 192 g to between 750 and 866 g, with growth being negatively affected by low dietary protein concentration. High dietary starch concentrations had some negative effects on growth, whereas changes in dietary fat concentration had no significant effect on growth. Liver indices (at the end of the experiment) varied between 80 and 170 g kg^?1, and there was a negative correlation between the ratio of protein to fat and liver index. Feed conversion ratio (FCR) ranged between 0.74 and 0.88, and feed utilization improved with increasing concentrations of dietary protein and fat. Increasing dietary starch concentrations resulted in poorer feed utilization. To achieve good growth and protein retention, and avoid excessive liver size in juvenile cod, feeds should contain 500–600 g kg^?1 crude protein, 130–200 g kg^?1 lipid and <150 g kg^?1 starch.     

Growth and gut morphology of diploid and triploid juvenile Atlantic cod (Gadus morhua)

The objective of this paper was to compare the growth and gut morphology of juvenile diploid and triploid Atlantic cod (Gadus morhua) reared under similar conditions. Individually tagged 36‐week‐old diploid (mean weight 49.3 ± 13.8 g) and triploid (mean weight 43.6 ± 11.2) juvenile cod were measured at intervals during a 29‐week growth trial. Data for weight, length, condition factor (K), hepato‐somatic index (HSI), gonado‐somatic index (GSI), Relative Gut Length (RGL) and pyloric caeca number were collected and results were analysed in relation to ploidy status, gender and family contribution. At the end of the experiment, only one family (M2xF3) had many representatives with a relatively even distribution of sexes and ploidies. Diploid females were significantly heavier and had higher K than triploid females in the M2xF3 family (body weight 371.2 ± 120.2 vs. 298.4 ± 100.7 g; K 1.1 ± 0.1 vs. 0.93 ± 0.1), but no differences were found between diploid and triploid males. In the other families (pooled data), no differences in body weight were found between the ploidy groups. In general, triploids had a shorter intestine (RGL) and fewer pyloric caeca than their diploid siblings regardless of gender suggesting possible impairments in nutrient utilization and growth.     

Development of a continuous cell line from larval Atlantic cod (Gadus morhua) and its use in the study of the microsporidian,Loma morhua

In vitro cell culture methods are crucial for the isolation, purification and mass propagation of intracellular pathogens of aquatic organisms. Cell culture infection models can yield insights into infection mechanisms, aid in developing methods for disease mitigation and prevention, and inform commercial‐scale cultivation approaches. This study details the establishment of a larval cell line (GML‐5) from the Atlantic cod (Gadus morhua) and its use in the study of microsporidia. GML‐5 has survived over 100 passages in 8 years of culture. The line remains active and viable between 8 and 21°C in Leibovitz‐15 (L‐15) media with 10% foetal bovine serum and exhibits a myofibroblast phenotype as indicated by immuno‐positive results for vimentin, α‐smooth muscle actin, collagen I and S‐100 proteins, while being desmin‐negative. GML‐5 supports the infection and development of two microsporidian parasites, an opportunistic generalist (Anncaliia algerae) and cod‐specific Loma morhua. Using GML‐5, spore germination and proliferation of L. morhua was found to require exposure to basic pH and cool incubation temperatures (8°C), in contrast to A. algerae, which required no cultural modifications. Loma morhua‐associated xenoma‐like structures were observed 2 weeks postexposure. This in vitro infection model may serve as a valuable tool for cod parasitology and aquaculture research.     

Residency of adult Atlantic cod (Gadus morhua) in the western Gulf of Maine

A mark and recapture study of cod in the western Gulf of Maine was conducted to study the seasonal movements of fish, particularly as they related to areas closed to commercial fishing. A total of 27,772 cod were tagged, and 1334 (4.8%) were recaptured with sufficiently detailed recapture location and date to be included in the study. Results indicated that the group is resident to the area and sedentary. Although there were a small percentage of fish (2.5%) that traveled long distances (>100 km), most were recaptured near their release location. There was no linear relationship between fish length and linear distances traveled, and linear distance traveled was only weakly related to days-at-large. For all groups of cod tagged and released in particular areas and months, mean distances traveled were small (<65 km), rates of travel were slow (<0.2 km/day), and rates of group dispersion were <50 km²/day. There were no recognizable spatial or temporal patterns in the mean angles of travel for groups released in various month/area combinations, and the angular deviations associated with the mean angles were quite large. Although there seems to be little pattern in the movement of cod in our study area, temporal and spatial changes in abundance indicate that movements are occurring.

The general pattern was a concentration of large cod into one small, inshore area (Area 133) in both the spring and winter, and dispersion from this area in the ensuing months. Monthly percentages of ripe cod in Area 133 provide evidence for two spawning groups; a winter group that spawns from November through January, and a spring group that spawns from April through July. Thus it is likely that the observed spring and winter concentrations of fish in Area 133 were associated with spawning. Fish in the spring group were the largest encountered in the study. The study also provided some evidence of natal homing, i.e. a return to the same spawning grounds year after year, for the spring spawning group. Although there is some exchange of fish between most of the closed areas, the timing of the closures appears to protect the largest aggregations of cod.     

Experimental susceptibility of Atlantic cod, Gadus morhua (L.), and Atlantic halibut, Hippoglossus hippoglossus (L.), to different genotypes of viral haemorrhagic septicaemia virus

Viral haemorrhagic septicaemia (VHS) is a well-characterized disease of rainbow trout, Oncorhynchus mykiss, which has also caused economic losses in marine turbot farms in the British Isles. We have previously demonstrated that turbot, Scophthalmus maximus, are susceptible to isolates of viral haemorrhagic septicaemia virus (VHSV) that are endemic in the marine environment, highlighting a potential risk to marine aquaculture. Given the increasing interest in the intensive rearing of additional aquaculture species such as Atlantic cod, Gadus morhua, and Atlantic halibut, Hippoglossus hippoglossus, this study aimed at investigating the susceptibility of these species to VHSV. Both species were found to be largely resistant to VHS following immersion challenge with a selection of 18 isolates, representing the known marine VHSV genotypes. Only one and two VHSV-associated mortalities occurred out of a total of 1710 and 1254 halibut and cod, respectively. These findings suggest that there is a low direct risk to the development of commercial cod and halibut aquaculture from the existing endemic reservoir of VHSV. This study, coupled to field observations has, however, highlighted the fact that both species can become infected with VHSV. The known adaptability of RNA viruses, together with the selection pressures associated with intensive aquaculture would thus advocate a cautious approach to VHSV surveillance within these emerging industries.     

Egg quality determinants in cod (Gadus morhua L.): egg performance and lipids in eggs from farmed and wild broodstock

Lipids and essential fatty acids, particularly the highly unsaturated fatty acids, 20:5n‐3 (eicosapentaenoic acid; EPA), 22:6n‐3 (docosahexaenoic acid; DHA) and 20:4n‐6 (arachidonic acid, AA) have been shown to be crucial determinants of marine fish reproduction directly affecting fecundity, egg quality, hatching success, larval malformation and pigmentation. In Atlantic cod (Gadus morhua L.) culture, eggs from farmed broodstock can have much lower fertilization and hatching rates than eggs from wild broodstock. The present study aimed to test the hypothesis that potential quality and performance differences between eggs from different cod broodstock would be reflected in differences in lipid and fatty acid composition. Thus eggs were obtained from three broodstock, farmed, wild/fed and wild/unfed, and lipid content, lipid class composition, fatty acid composition and pigment content were determined and related to performance parameters including fertilization rate, symmetry of cell division and survival to hatching. Eggs from farmed broodstock showed significantly lower fertilization rates, cell symmetry and survival to hatching rates than eggs from wild broodstock. There were no differences in total lipid content or the proportions of the major lipid classes between eggs from the different broodstock. However, eggs from farmed broodstock were characterized by having significantly lower levels of some quantitatively minor phospholipid classes, particularly phosphatidylinositol. There were no differences between eggs from farmed and wild broodstock in the proportions of saturated, monounsaturated and total polyunsaturated fatty acids. The DHA content was also similar. However, eggs from farmed broodstock had significantly lower levels of AA, and consequently significantly higher EPA/AA ratios than eggs from wild broodstock. Total pigment and astaxanthin levels were significantly higher in eggs from wild broodstock. Therefore, the levels of AA and phosphatidylinositol, the predominant AA‐containing lipid class, and egg pigment content were positively related to egg quality or performance parameters such as fertilization and hatching success rates, and cell symmetry.     

Optimization of Heat Processing of Farmed Atlantic Cod (Gadus morhua) Muscle with Respect to Cook Loss,Water Holding Capacity,Color, and Texture

Convenience products of fish that are heat processed in-pack are often subjected to a compromise between quality and durability within the framework of required safety. The purpose of this study was to provide knowledge for optimizing pasteurization of cod muscle by quantifying changes in water holding capacity (WHC), cook loss, color, and texture within a wide range of processing times and temperatures. A heat load equivalent to 2 min at 70°C may be applied to the cod while maintaining the WHC above 66%, the cook loss below 5.6%, and keeping hardness and whiteness low, provided that the processing temperature is kept in the range 68 ± 4°C.     

Seasonal shift in spawning of Atlantic cod (Gadus morhua L.) by photoperiod manipulation: egg quality in relation to temperature and intensive larval rearing

Commercial intensive fry production of Atlantic cod will be dependent on production of viable eggs independent of season. This can only be done by manipulation of maturation by photoperiod, but little is known about potential effects on egg characteristics and larval viability. In two cod broodstocks, maturation was successfully advanced or delayed 6 months compared with normal spawning season (March–April) by manipulation of photoperiod. The advanced broodstock spawned both in spring and autumn the same year. In two of the spawning tanks during autumn, ambient temperature was reduced after reaching 13.7°C during the first half of the spawning period. Egg quality and viability were monitored, and several egg batches were incubated, hatched and start‐fed for examination of growth and survival. Temperatures above 9.6°C resulted in significant reductions in fertilization and normal egg development. Concurrently, fractions of dead and unfertilized eggs increased with elevated temperature. Actual relative fecundity was not affected by temperature. Egg characteristics improved when temperature was controlled and lowered below 9.6°C. Occurrence of irregular spawners suggests that handling of broodstock fish should be avoided during maturation and spawning. Cod larvae originating from eggs of the advanced or delayed broodstocks were successfully reared beyond metamorphosis. Survival was 9.0–46.6% and 29.3% in green and clear water respectively. Survival correlated with both initial and average feeding conditions, but growth rate did not correlate with either of survival and feeding conditions. Specific growth rates (8.3–13.6% day⁻¹) is comparable with other intensive rearing trials with cod, but were lower than reported from nature‐like systems.     

Natural zooplankton as larval feed in intensive rearing systems for juvenile production of Atlantic cod (Gadus morhua L.)

The growth potential of cod larvae is not fully achieved when rotifers (Brachionus spp.) are used as live feed. In this experiment, we studied the effect of natural zooplankton (mainly copepods) on the growth of cod (Gadus morhua L.) larvae reared in intensive systems. Using a growth model developed for cod larvae, the growth rates observed could be evaluated and compared with growth rates reported previously. The cod larvae showed optimal growth rates until age 19 days post hatch (DPH) when they reached 9.77 ± 0.25 mm standard length (SL). Early weaning (20–25 DPH) resulted in significantly longer larvae at age 30 DPH compared with late weaning (25–32 DPH); however, in this period, the zooplankton concentrations were low. The experimental larvae showed considerably higher growth rates compared with rotifer (Brachionus spp.)‐reared cod larvae in previous experiments. The nutritional composition of cod larvae was analysed and compared with published results on rotifer‐reared larvae. The levels of iodine, manganese, selenium and n‐3 PUFA were considerably higher in larvae fed copepods compared with larvae fed rotifers. The differences in nutritional status may well explain the differences in growth observed between copepod and rotifer‐reared larvae.