Allometric growth in juvenile cod (Gadus morhua) and possible effects on cannibalism

Cod juveniles of about 20 mm had the lowest potential predator: prey length ratio based on mouth gape of the predator and the body height of the prey. This implies that the cannibalistic potential is maximized in this size range.     

The effect of energetic condition on growth dynamics and health of Atlantic cod (Gadus morhua)

Prolonged starvation resulting in sublethal condition factor values was hypothesized to have a detrimental effect on short‐term growth capacity upon refeeding. Cod (Gadus morhua) were food‐deprived and their length and mass measured before refeeding and after 3, 6, 9 and 12 weeks of ad libitum feeding. Total mass increase during the first 3 weeks of feeding was greatest in fish with a higher initial condition factor. The reverse situation was observed during the last 3 weeks of feeding. Specific growth rate peaked in the period from week 4 to week 6, except in cod with the highest condition factor for which a steady decline in specific growth rate was observed, and was not influenced by the condition factor at the start of the feeding period. Total mass increase over 12 weeks was also not influenced by initial condition factor. Thus by the end of the experiment, condition factors were lowest in fish with initially low condition factors. The hepatosomatic index and gonadosomatic index did not differ at the end of the experiment, but the proportion of mature cod increased with increasing initial condition factor. A disease outbreak caused significant mortalities among fish shortly after the start of the feeding period. Forty‐one percent of the fish had died after 84 days. No mortality was observed among fish that had started the experiment with the highest condition factor. Mortality was inversely related to initial condition factor. Growth was examined for survivors exclusively. Poor condition in wild fish may increase vulnerability to diseases and compensatory growth may not allow cod with low condition factors to fully recover unless food availability remains high over a long period of time.     

Minisatellite DNA fingerprints of Atlantic cod (Gadus morhua)

The human minisatellite probes 33.6 and 33.15 cross–hybridized to Hae III and Hinf I digested cod DNA, revealing complex fragment patterns in both Arctic and coastal cod. The DNA fingerprints were highly polymorphic, individual specific and stable in the germline. The potential applications of multi locus probes in cod research are discussed.     

Substrate selection by juvenile Atlantic cod (Gadus morhua): effects of predation risk

Although predator avoidance has been proposed as one possible factor influencing the distribution of fish among substrate types, no study has addressed this question directly. Groups of juvenile Atlantic cod were offered a choice between pairs of the following three substrates: sand, gravel-pebble and cobble. Their distribution on these substrates was compared prior to, during and following exposure to a predator (i.e. a larger conspecific). With no apparent risk of predation, juvenile cod preferred sand or gravel-pebble. When cobble was present, juveniles hid in the interstitial spaces of this substrate in the presence of a predator. With no cobble present, juveniles showed no preference between sand and gravel-pebble, and did not seek refuge from predation in association with these substrates. Following exposure to a predator (i.e. 2.5 h later) larger juvenile cod again showed a preference for the finer-grained substrates, but smaller individuals continued to associated with the cobble. The presence of cobble resulted in fewer juveniles being captured and a significant increase in the latency until the first juvenile was captured by the predator. Results are discussed with respect to the effects of predation on the distribution and survival of fishes among substrate types.     

Countergradient variation in body shape between two populations of Atlantic cod (Gadus morhua)

Variation in morphological traits is generally thought to be cogradient, with environmental effects on phenotypic expression reinforcing genetic differences between populations. We compared body shape between two populations of Atlantic cod (Gadus morhua). Striking shape differences occurred between juveniles from the two populations when reared in a common laboratory environment. However, no difference in body shape occurred between wild-reared juveniles from the two populations, suggesting that the genetic differences between populations were obscured by opposing effects of the environmental differences experienced in the wild. We suggest that much of the genetic diversity in body shape of fishes may be cryptic, with stabilizing selection for the same optimal phenotype resulting in genetic divergence between populations subject to contrasting environmental influences.     

Measurements of larval Atlantic cod (Gadus morhua) routine metabolism: temperature effects, diel differences and individual-based modeling

In the present study, dry mass (M_D, μg) and routine respiration rate (R_R) (μl O₂ ind^?1 h^?1) were measured for larval cod, Gadus morhua (L.) that were reared and tested at 5.0, 7.5, and 10.0°C. Bi‐hourly measurements of R_R were made on groups of larvae using a closed‐circuit respirometer over a 24‐h period (14L : 10D light regime) to test temperature and body size effects and whether unfed larvae exhibited diel differences in metabolism. At 10°C, the relationship between mean R_R and mean M_D was: ln R_R = 1.16·lnM_D ? 6.57 (n = 31, r² = 0.883, P < 0.001). The exponential increase in R_R with temperature was described by a Q₁₀ of 3.00. Diel differences in unfed larvae were only apparent in groups of the largest larvae. A comparison of Q₁₀ estimates from this and other studies suggest a linear decrease in the effect of temperature on cod R_R with increasing log M_D for sizes encompassing larvae to large juveniles. The trend may explain, in part, observations of cod juveniles exploiting a wider range of in situ temperatures than larvae. Finally, the two most comprehensive data sets on larval cod R_R compare poorly (approximately five‐fold differences) and our results support the assertion that daily metabolic energy loss in many larval cod individual‐based models were based upon measurements that over‐estimated hourly metabolic rates by a factor of approximately four.     

A reappraisal of activity metabolism in Atlantic cod (Gadus morhua)

Atlantic cod ( Gadus morhua ) were forced to swim in a swim tunnel respirometer until fatigued; oxygen consumption rate (O₂) was measured during swimming at incremental speeds until the fish was exhausted and during recovery from exhaustion. Maximal oxygen consumption (O_2max) occurred during maximal activity as has been found for other fish species, but at odds with the current paradigm for Atlantic cod. Earlier experiments had drawn the conclusion that O_2max in Atlantic cod occurs during recovery from exhaustive exercise. We found no support for this paradigm in our experiments and we propose that the respiratory physiology of Atlantic cod is not unlike that of other fishes.     

Phenotypic flexibility of digestive system in Atlantic cod (Gadus morhua)

This study examined the restoration of the digestive capacity of Atlantic cod (Gadus morhua Linnaeus) following a long period of food deprivation. Fifty cod (48 cm, 1 kg) were food-deprived for 68 days and then fed in excess with capelin (Mallotus villosus Müller) on alternate days. Ten fish were sampled after 0, 2, 6, 14 and 28 days and the mass of the pyloric caeca, intestine and carcass determined. Two metabolic enzymes (cytochrome c oxidase and citrate synthase) were assayed in white muscle, pyloric caeca and intestine, and trypsin activity was measured in the pyloric caeca. A delay of 14 days was required before body mass started to increase markedly, whereas most of the increase in mass of both the pyloric caeca and intestine relative to fish length occurred earlier in the experiment. By day 14, the activities of trypsin and citrate synthase in the pyloric caeca as well as citrate synthase in the intestine had reached maxima. The growth of the digestive tissues and restoration of their metabolic capacities thus occur early upon refeeding and are likely required for recovery growth to take place. The phenotypic flexibility of the cod digestive system is therefore remarkable: increases in trypsin activity and size of pyloric caeca resulted in a combined 29-fold increase in digestive capacity of the fish during the refeeding period. Our study suggests that Atlantic cod are able to cope with marked fluctuations in food availability in their environment by making a rapid adjustment of their digestive capacity as soon as food availability increases.     

Diclidophora merlangi (Monogenea: Diclidophoridae) on Atlantic cod, Gadus morhua

Diclidophora merlangi (Kuhn, in Nordmann, 1832), as species specific to whiting, Merlangius merlangus (Linnaeus, 1758), is reported and described for the first time on the gills of Atlantic cod Gadus morhua (Linnaeus, 1758). Of the about 1,200 cod examined in this article, only 2 specimens of D. merlangi occurred on 2 fish, suggesting that they represented accidental infections. The 2 specimens showed specific traits of D. merlangi, but their size was considerably smaller than that of D. merlangi on whiting. Principal-component analyses showed that the morphology of the specimens on cod was more similar to that of D. merlangi than those of other congeneric species from the North Atlantic, additionally supporting their assignment to D. merlangi. Quantitative data documenting the potential reproductive consequences of D. merlangi occurring on cod is also provided. The testes and germaria of the specimens on cod were noticeably smaller than those of D. merlangi on whiting, but the number of testes was similar and spermatozoa were observed in the sperm duct. By contrast, the aspect and smaller size of the oocytes suggested that D. merlangi on cod could not produce viable ova, although 1 specimen exhibited an egg with normal sized and shaped capsule.     

Purification and characterization of pancreatic elastase from Atlantic cod (Gadus morhua)

1. A pancreatic elastase from Atlantic cod (Gadus morhua) has been purified and characterized. 2. The enzyme is a very basic protein with an approximate mol. wt of 28,000. 3. The cod elastase has higher elastin specificity than porcine elastase, and it is inhibited by soybean trypsin inhibitor, which has no effect on porcine elastase. 4. The cod elastase expresses a higher turnover number (kcat) and catalytic efficiency (kcat/Km) than porcine elastase, but it is less thermostable.     

Correlations between hemoglobin type and temperature preference of juvenile Atlantic cod Gadus morhua

Atlantic cod (Gadus morhua L.) exhibits polymorphic hemoglobin variants with the HbI locus showing a strong North-South geographic cline in frequency distribution of three main types (1/1, 1/2 and 2/2). This may indicate selective advantages of the different HbI types under various temperature regimes. Despite this only one study has directly examined the temperature preference of the two homozygous types, HbI-1/1 and HbI-2/2, whereas the preference of the heterozygote (HbI-1/2) has never previously been addressed. By exposing fish to a 4–19 °C temperature gradient in an annular preference chamber we recorded the preferred temperature of wild juvenile G. morhua of all three main Hbl types originating from an area where they co-exist. HbI-2/2 G. morhua preferred significantly cooler water (8.9 ± 0.2 °C) compared to the HbI-1/1 group (11 ± 0.6 °C), this difference, however, not being as distinct as previously reported. There was pronounced inter-individual variation in the temperature preference of the HbI-1/2 G. morhua ranging between 6.7 and 13.8 °C, and their overall preference (10.5 ± 0.9 °C) did not differ significantly from either of the homozygous HbI types. Notably, the mean range of utilized temperature (temperature span between 1st and 3rd quartile) was very similar between all 3 Hbl types with 3.2–3.5 °C. Considering the complexity of a trait like temperature preference, there are clearly many other factors besides HbI type that influence the thermal biology of cod, and therefore we also investigated possible associations between genotype and temperature preference for 12 variable candidate gene single nucleotide polymorphisms (SNPs) a priori expected to be related to growth and reproduction. There were, however, no significant correlations between temperature preference and any of the candidate gene SNPs indicating that none of these polymorphisms strongly associates with thermal behavior. Considering however the high-throughput genotyping methods becoming increasingly accessible there is great potential for association studies involving many more genetic markers to identify additional genetic polymorphisms that are important for temperature preference in G. morhua. In conclusion, we support the notion of a ‘warm’ (HbI-2/2) and a ‘cold’ (HbI-1/1) Hb type, although we suggest the difference to be more subtle than previously reported. Furthermore HbI-1/2 G. morhua shows rather inconsistent thermoregulatory behavior. To obtain a more definitive picture of the extent to which thermal niches are realized under natural conditions field observations in areas where the 3 HbI types co-exist should be performed.     

Neutrophils and B-cells in Atlantic cod (Gadus morhua L.)

Proportions of leucocytes from head kidney, blood and spleen were identified as B-cells and neutrophils using a polyclonal antibody to cod IgM and a monoclonal antibody which previously has been shown to bind specifically to salmon and trout neutrophils. The cell specific binding of the antibodies was supported by double immunostaining. The morphology of isolated leucocytes was examined on Diff Quick stained slide preparations, and myeloperoxidase positive neutrophils were identified by diaminobenzidine staining. The antibodies clearly identified distinct cell populations. Using flow cytometry, high proportions of neutrophils were observed in peripheral blood leucocytes and high proportions of B-cells were found in head kidney leucocytes when compared to proportions of these cells in Atlantic salmon (Salmo salar L.). The spleen contained the highest proportion of B-cells. Cytoplasmic staining of immunoglobulin positive cells in slide preparations indicated that plasma cells were present, but not strikingly abundant, in head kidney, spleen and peripheral blood. Staining for myeloperoxidase identified, in accordance with the flow cytometry results, a large number of neutrophils, especially in peripheral blood leucocytes. The neutrophil nucleus was not clearly segmented, but appeared more irregular than rounded. The findings of high proportions of neutrophils in peripheral blood suggest that these cells of the innate immune system might have a central role in defence and protection against infections in cod.     

Influence of Lernaeocera branchialis (Crustacea: Copepoda) on growth rate of Atlantic cod, Gadus morhua

Influence of Lernaeocera branchialis, a hematophagous copepod, on the growth rate of Atlantic cod, Gadus morhua, was investigated in fish of various sizes. Initially, it appeared that growth, over a 16-mo period, was reduced when single or multiple infections were present. We then examined growth rate by comparing food consumption, weight gain, food conversion efficiency, and condition (k) factor over a 2-mo period and subsequently over a 4 wk in control and experimentally infected fish. Total body weight and length of each fish, controls and infected were determined before and after feeding known weights of food. Adults infected with young parasites consumed more food and gained more weight than controls, especially in the first month, but food conversion efficiency and k-factor were greater than those of controls after 2 mo, but food efficiency was still lower than in controls. In 2 groups of young infected fish, all values were consistently lower than those of uninfected fish of comparable size during the 2-mo period. Three additional trials on immature cod confirmed that infected fish consumed less food and exhibited less growth than infected controls. Our results suggest that only adult cod infected with immature parasites compensate for the infections by increasing food consumption, thus resulting in a transitory weight gain. Additionally, parasites that have completed their reproductive activities still produce subtle effects in the form of low food conversion efficiency in their hosts.     

New haemoglobin genotypes in Atlantic cod, Gadus morhua: possible relation with growth

In a preliminary study, 121 individually tagged juvenile Atlantic cod (Gadus morhua) were classified according to their haemoglobin genotypes into four groups, i.e., two main haemoglobin genotypes [Hb-I(1/2), Hb-I(2/2)] and two sub-types [Hb-I(1/2b), Hb-I(2/2b)], and reared for 3 months at 10 degrees C, 13 degrees C and T-step (fish reared at 16 degrees C and then subsequently moved to 13 and later to 10 degrees C). Overall growth rates across temperatures were 10% and 19% higher in the Hb-I(2/2b), Hb-I(1/2b) sub-types compared to corresponding Hb-I(2/2) and Hb-I(1/2) main types, respectively. Individual growth rate trajectories varied between the genotypes at all temperatures studied. Our study indicates that under certain environmental conditions higher growth in the two sub-types compared to the main genotypes could be expected. This may indicate difference in other physiological characters not studied here, but seen in previous studies, i.e., oxygen affinity and competitive performance.     

The acute phase response of Atlantic cod (Gadus morhua): humoral and cellular response

Intra-muscular injection of turpentine oil was used to induce acute phase response (APR) in Atlantic cod (Gadus morhua L.). The effects on the serum cortisol, total protein, IgM and pentraxin concentration were examined as well as the effects on natural antibody, anti-trypsin and leukocyte respiratory burst activity. The turpentine injection resulted in a 26 fold increase in the cortisol level after 72 h. Slightly reduced serum protein level in both groups was attributed to the restricted feeding during the experimental period. The IgM serum concentration was significantly reduced after 168 h in the turpentine treated fish while the natural antibody activity was not affected. The anti-trypsin activity was initially suppressed but recovered to normal levels at the end of the experiment. The turpentine injection had little effect on the serum level of the pentraxins, CRP-PI and CRP-PII. The respiratory burst activity was significantly suppressed after 72 h. It is concluded that 1) cod shows a relatively slow humoral and cellular response to APR induction, 2) the increase in serum cortisol level may be the key modulator of the mainly suppressive effects on the immune parameters and 3) pentraxins are not typical acute phase proteins in cod.     

Susceptibility of juvenile Atlantic cod Gadus morhua to viral haemorrhagic septicaemia virus isolated from wild-caught Atlantic cod

A European strain of viral haemorrhagic septicaemia virus (VHSV) isolated from wild-caught cod Gadus morhua (H16/7/95) was shown to cause clinical disease and mortality in excess of 80% in juvenile Atlantic cod when administered by the intra-peritoneal (i.p.) route. No virus was recovered from cod cohabiting with experimentally infected fish at a ratio of 1:1, and no VHSV-associated mortality was demonstrated following immersion infection. External signs of disease in cod were the presence of exophthalmia and ascites. Virus was identified as VHSV by enzyme-linked immunosorbent assay (ELISA) and was recovered from both brain and organ pools (kidney, liver and spleen) of 100% of i.p. infected cod mortalities. Virus was also detected using an indirect immunofluorescence test on tissue imprints of kidney, liver, spleen and brain taken from moribund fish. The fact that cod were not susceptible to VHSV following waterborne exposure raises important questions surrounding the propagation, maintenance and impact of a naturally occurring reservoir of virus in the marine environment.     

Enhanced protein synthetic capacity in Atlantic cod (Gadus morhua) is associated with temperature-induced compensatory growth

Atlantic cod (Gadus morhua) were held either at seasonal ambient temperatures (-0.3 to 11 degrees C) or at a relatively constant control temperature (8-11 degrees C) to investigate aspects of protein synthesis during a period of compensatory growth. Protein synthesis rate, total RNA, and RNA-specific protein synthesis rate were determined in white muscle and liver when ambient temperatures were -0.3, 4.5, and 11 degrees C in February, June, and July, respectively. To allow for comparisons between treatment temperatures, fish were also acutely transferred to a comparable assay temperature in February and June. Over the transition from 4.5 to 11 degrees C (June to July), the ambient-held cod had a significant increase in size and a substantially higher growth rate relative to control-held fish over the same period, consistent with cold-induced compensatory growth. During the onset of this enhanced growth, in June when ambient temperature was approximately 4.5 degrees C, ambient-held fish elevated their capacity for protein synthesis in the white muscle and liver via elevation of the RNA content. When ambient temperature reached the same point as for the control fish (11 degrees C), the rate of white muscle protein synthesis remained higher in the ambient-held vs. that in the control-held fish, a process facilitated by elevated RNA content and greater RNA-specific rate of protein synthesis. In the liver, all measured characteristics of protein synthesis were the same for ambient and control fish in July. The latter suggests that compensatory growth may be in part explained by improved efficiency of protein synthesis.     

Direct and social genetic parameters for growth and fin damage traits in Atlantic cod (Gadus morhua)

Background

The aim of the study was to estimate genetic parameters for direct and social genetic effects (SGE) for growth and welfare traits in farmed Atlantic cod (Gadus morhua). A SGE refers to the effect of an individual’s genes on trait performance of its social partners. In total, 2100 individually tagged juveniles from 100 families at an average age of 222 days post-hatching were used. Each family was separated into three groups of seven fish, and were randomly assigned to 100 experimental tanks, together with fish from two other families. Body weight and length of the first, second and third dorsal fin and the caudal fin measured by digital image analysis were measured at the start of the experiment, after two weeks, and after six weeks. Fin erosion was scored subjectively after six weeks. Variance components estimated using a conventional animal model were compared to those of an animal model including a SGE.

Results

Heritabilities from the conventional animal model ranged from 0.24 to 0.34 for body weight and 0.05 to 0.80 for fin length. Heritabilities for fin erosion were highest for the first dorsal fin (0.83 ± 0.08, mean ± standard error) and lowest for the third dorsal fin (0.01 ± 0.04). No significant SGE were found for body weight, whereas SGE for fin lengths were significant after two and six weeks. Contributions to the total heritable variance were equal to 21.5% (6.1 ± 2.1) for the direct effect, 33.1% (9.4 ± 3.2) for the direct-social covariance, and 45.4% (12.9 ± 4.1) for the social variance for length of the first dorsal fin. For fin erosion, SGE were only significant for the second and third dorsal fin.

Conclusions

Including SGE for fin length and fin erosion in the animal model increased the estimated heritable variation. However, estimates of total heritable variances were inaccurate and a larger experiment is needed to accurately quantify total heritable variance. Despite this, our results demonstrate that considering social breeding values for fin length or fin erosion when selecting fish will enable us to improve response to selection for welfare traits in Atlantic cod juveniles.