Strong phenotypic and genetic correlation between size and first maturity in Atlantic cod Gadus morhua L. reared in commercial conditions

Three‐year classes of Atlantic cod Gadus morhua were studied throughout their rearing in the eastern coast of Iceland from 2004 to 2011. The growth and status of maturity were recorded during the rearing. For one of the year classes, genetic parameters for body weight and maturity status were estimated from 757 individuals, which were the offspring of 40 dams and 20 sires. The estimate for heritability of body weight was h² = 0.34 at the average weight of 630 g, and heritability for proportion of maturity was h² = 0.17 given the same weight. The relationship between body weight and the proportion of mature individuals at first winter revealed a strong genetic correlation of r_G = 0.90. The phenotypic relationship between body weight and proportion of maturity was estimated with a Bayesian logistic regression as P(yi = 1(mature)) = Φ(β0 + β1_weight + β2_sex). The best fit yielded β0 = ?2.9320 with a 95% interval between ?3.2807 and ?2.5394, β1_weight = 0.0041 with a 95% interval ranging from 0.0035 to 0.0046 and β1_sex = ?0.0201 with a 95% interval from ?0.2003 to 0.1445. The gender had no notable effect. This strong phenotypic and genetic correlation in body weight and maturation suggests that an increased growth rate will consequently lead to a higher proportion of mature individuals in the population. As a consequence, genetic manipulations to simultaneously increase growth and delay maturation may present a challenge.     

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.     

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.     

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.     

Benchmarking growth of farmed Atlantic cod,Gadus morhua: a case study in Ireland

The growth performance of different stocks of farmed North Atlantic cod over several years in Irish rearing conditions is described. The growth trajectories for all stocks reared were similar. From all growth data, it was found that weight at 180 and 365 days post hatch (dph), with mean values of 17.7 and 158.7 g, respectively, are accessible and useful indicators of hatchery growth performance. The importance of using standard benchmarking points to facilitate concerted and improved research efforts is highlighted here. Growth performance was evaluated against existing predictive models and these were found to be quite robust and very useful within certain constraints. Comparative growth modelling using prevailing ambient sea surface temperature regimes for five different countries (Ireland, Norway, Canada, Iceland and the Faeroes) was also undertaken. Harvest size of 2.5 kg was estimated to be reached in approximately 36 months post hatch in the Faeroes, 38 months in Ireland, 37 months in Norway, 40 months for Canada and up to 42 months in Iceland.     

Experimental mycobacteriosis in Atlantic cod, Gadus morhua L

Piscine mycobacteriosis causes losses in a number of fish species both in the wild and in aquaculture worldwide. Mycobacterium salmoniphilum infections have on several occasions been reported in farmed Atlantic salmon, Salmo salar L. The present study tested and confirmed the susceptibility of Atlantic cod, Gadus morhua L., an important yet relatively novel aquaculture species, to infection with M. salmoniphilum. Atlantic cod injected intraperitoneally with a suspension of this bacterium were maintained together with cohabitant (COH) fish in a flow-through marine water system at 10-11 °C. The fish were supervised daily and samples taken at 2, 7, 14, 23, 34 and 53 weeks post-infection and examined pathologically, bacteriologically and using molecular biology. Injected mycobacteria were re-isolated in high concentrations from both injected and COH fish groups. Death attributable to mycobacterial infection was observed in both injected (47%) and COH (28%) fish groups. Extensive development of granuloma in visceral organs, mainly the mesenteries, spleen, kidney and liver (lesser extent) and at later stages of the infection in heart tissues and gills, was observed in both injected and COH fish. Granulomas underwent a temporal progression of distinct morphological stages, culminating in well-circumscribed lesions surrounded by normal or healing tissue. Acid-fast bacilli were detected in both granulomas and non-granulomatous tissues. This study confirms that Atlantic cod is highly susceptible to M. salmoniphilum infection and that this bacterial species may be a threat to cod both in the wild and in the aquaculture.     

A follow‐up study of selected biomarkers of health in cod Gadus morhua L. collected from the southern Baltic off the Polish coast

Selected biomarkers of health were examined in 50 post‐spawning cod Gadus morhua collected in November 2015 from the southern Baltic. The biomarkers included condition factor (CF), macroscopic lesions, histopathology of spleen, liver and gonads, and morphometry of follicular atresia and hepatic and splenic melanomacrophage cells (MMC). All fish appeared in good body condition. One fish had a dermal ulcer, and in seven, macroscopic nematodes were noted within body cavity. Microscopic lesions in the liver included biliary myxozoanosis, microsporidial and necrocentric granulomas, parasitic hepatitis, multifocal necrosis, foci of cellular alterations, spongiosis, peliosis and cytoplasmic fibrillar inclusions. The spleen and gonads had microsporidial and/or necrocentric granulomas. Some of the biomarkers showed differences as compared to spawning cod collected in May from the same location in 2012, most importantly values an order of magnitude lower for splenic MMC in post‐spawning fish. In post‐spawning fish, there were statistically significant correlations between MMC, CF, follicular atresia, parasitic hepatitis and microsporidiosis. This is the first comparison of biomarkers of health in post‐spawning and spawning Baltic cod. Future studies need to examine the relationships of biomarkers to levels of pollutants in the environment and in tissues of cod.     

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.     

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.     

Evaluating spatio‐temporal variability in the habitat quality of Atlantic cod (Gadus morhua) in the Gulf of Maine

Reduced abundance and contracted spatial distribution of Atlantic cod, Gadus morhua, in the Gulf of Maine (GOM) may indicate large spatio‐temporal variation in their habitat quality. Season‐specific Habitat Suitability Index (HSI) models were developed to quantify such variation in the offshore GOM management area. Data used were non‐zero cod catch rates with calibrations from the Northeast Fisheries Science Center (NEFSC) spring and fall bottom trawl surveys over the period 1982–2013 and key physical environmental variables including depth, bottom temperature, bottom salinity and sediment types. Significant declines were found in the average HSI across the study area in the springs of early 2000s and 2010s. These low average HSI values coincide with reduced age‐1 recruitment of GOM cod stock after the mid‐1990s. Moreover, the western coastal areas of the GOM generally exhibited higher average HSI values than the eastern coastal areas, whereas the offshore areas always had the lowest average HSI. Relatively higher cod survey catch rates in the western GOM may imply positive influences of environmental controls on the distribution of GOM cod.     

Changes in Atlantic cod (Gadus morhua L.) sperm quality during the spawning season

The biology of cod reproduction is well described in the scientific literature. However, sperm biology and spermatozoa management are poorly studied in this species. Because of its recent farming expansion, a better knowledge of cod gametes is becoming especially useful. This work aimed at establishing tools to study sperm biology in cod, and also investigated the existence of changes in cod sperm quality during the spawning period. We showed that sperm concentration could be assessed using spectrophotometry at 260 nm. Sperm motility significantly decreased after a 168‐h storage at 4 °C. A 1:9 dilution of sperm in a non‐activating medium (1/3 seawater and 2/3 freshwater, osmotic pressure: 360 mOsm kg^?1) improved sperm storage. Sperm concentration, sperm velocity and storage capacity at 4 °C peaked during the medium period of the spawning season and then decreased to values close to those observed at the beginning of the reproductive period. The measured values of osmotic pressure, pH, protein, Na⁺, Cl^? and Ca²⁺ concentrations of the seminal fluid were modified along the spawning period. Cell damage was noted at the end of the spawning period: local blebs were observed on the flagellum but also loops at its distal part. On the other hand, spermatocrit did not vary with the sampling date. In conclusion, cod sperm quality is modified during the spawning period, the highest‐quality samples being collected during the medium part of this season.     

The effect of dietary chitin on the autochthonous gut bacteria of Atlantic cod ( Gadus morhua L.)

In the present study the impact on autochthonous (adherent) bacteria in proximal intestine (PI) and distal intestine (DI) of Atlantic cod ( Gadus morhua L.) was evaluated following feeding of a control diet and a diet supplemented with 5% chitin. The autochthonous gut bacteria were investigated using denaturing gradient gel electrophoresis (DGGE). Analysis of the microbiota associated with PI and DI of Atlantic cod indicate that dietary chitin modulate the intestinal bacterial community. For example, band 25 ( Escherichia coli–like), band 14 ( Anaerorhabdus furcosa–like) and band 29 (uncultured bacterium–like) in PI were depressed by dietary chitin (P < 0.05). The number of bands (23.7 ± 5.4) in DI of fish fed chitin was marginally higher than the control fish (16.7 ± 2.1) (P = 0.065), and the relative abundances of band 6 (swine faecal bacterium–like) were marginally stimulated by dietary chitin (P = 0.095). Furthermore, the present study reports several novel sequences not previously reported in the gastrointestinal tract of Atlantic cod. Whether the dietary effect of chitin on gut bacterial community has any positive effect of fish health merits further investigations.     

Stress response to anthropogenic noise in Atlantic cod Gadus morhua L.

The potential effects of anthropogenic noise on the physiology of Atlantic cod have not been well described. The aim of the present study was to investigate the impact of anthropogenic noise on Atlantic cod stress response using cortisol as a biomarker as well as on broodstock spawning performance. Results showed that artificial noise consisting of a linear sweep from 100 to 1000 Hz can induce a transient and mild cortisol elevation with a clear noise intensity dose response. In all cases plasma levels returned to baseline levels <1 h post sound exposure. Daily exposure to a similar intensity and frequency noise range applied habitually to a broodstock population during the spawning window resulted in a significant reduction in total egg production and fertilisation rates thus reducing the total production of viable embryos by over 50%. In addition, a significant negative correlation between egg cortisol content and fertilisation rate was observed. These results confirm that cod can perceive noise generated within a frequency range of 100–1000 Hz and display a heightened cortisol plasma level. In addition, anthropogenic noise can have negative impacts on cod spawning performances.     

Dietary plant protein utilization in Atlantic cod, Gadus morhua L.

The present trials aimed to investigate the effects of replacing fish meal with plant proteins in diets for cod, using a regression design where fish meal constituted the control. The plant protein diets were formulated to meet the amino acid requirements according to NRC (1993) and contained corn gluten meal, soybean meal, a mixture of these, or a mixture of wheat gluten meal and soybean concentrate. The plant protein fraction constituted up to 440 g kg⁻¹ of the extruded diet. Two feeding experiments were conducted, one at high (11 °C) and one at low (6.5 °C) temperature. High growth and feed utilization were obtained in all diet groups at both temperatures. However, only in Exp.1 (11 °C), growth and feed utilization were linearly reduced by increases in dietary soybean and corn gluten meal, while no such effect was detected in Exp.2 (6.5 °C). Reductions in protein retention were seen at both temperatures with use of all evaluated plant protein sources. Due to large amounts of plant protein ingredients in diet, differences in diet amino acid composition were seen. These were partly reflected in the muscle free amino acid pool. Dietary plant ingredients did not affect whole body, liver or muscle proximate compositions, or liver indices. In both experiments, blood parameters were within the range of earlier reported normal values and indicated, together with low mortality, good fish health status. The results show that there is high potential to use protein‐rich plant ingredients in diets for Atlantic cod.     

Mercury comparisons between farmed and wild Atlantic salmon (Salmo salar L.) and Atlantic cod (Gadus morhua L.)

Wild and farmed Atlantic salmon ( Salmo salar L.) and Atlantic cod ( Gadus morhua L.) were collected to assess changes in mercury with size in wild vs. farmed fish. Mercury concentrations were compared with Health Canada and United States Environmental Protection Agency consumption guidelines. Lipid dilution of mercury was examined by comparing lipid-extracted (LE) and non-lipid-extracted (NLE) flesh samples in both farmed and wild fish. Mercury concentrations in the flesh and liver of farmed salmon were significantly lower than concentrations in wild salmon of similar fork length ( P <0.001), possibly due to growth dilution in rapidly growing farmed fish. Mercury concentrations were higher in LE tissue compared with NLE ( P <0.05), suggesting lipid dilution of mercury in farmed fish with a high lipid content. Farmed cod, which do not grow more rapidly than wild cod, did not have significantly different flesh and liver concentrations compared with wild cod of similar fork length ( P >0.05). Between species of farmed fish, cod had significantly higher mercury concentrations than salmon ( P <0.05), but neither farmed nor wild salmon mercury concentrations exceeded federal consumption guidelines. These results suggest that rapid growth rates and a high lipid content may play important roles in regulating concentrations of contaminants such as mercury.     

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.