Development and evaluation of microparticulate diets for early weaning of Atlantic cod Gadus morhua larvae

Early weaning trials were conducted with cod larvae to investigate the effectiveness of microparticulate diets (microbound and microcoated) with and without lipid-walled capsules (LWCs). The microparticulate diets were evaluated by measuring physical parameters of the diet in the water column (leaching and settling rate), palatability (intestinal fullness), performance of the diet (survival and growth), and examination of the diet in the larval intestine (histological analysis). A feeding trial was conducted using four experimental diets (carrageenan microbound diet, carrageenan microbound diet with LWCs, zein microcoated diet and zein microcoated diet with LWC), one commercial diet (BioKyowa: A-250) and a live feed control (rotifers and Artemia ). Survival of cod larvae to 39 days post-hatch ranged from 5 to 10% with the experimental diets, 22.9% with the BioKyowa diet, and 36.5% with live prey. There was evidence of food absorption with all diets in the form of lipid vacuoles in the midgut and supranuclear vacuoles in the hindgut. Large vacuoles in the midgut were more abundant in the enterocytes of larvae fed the experimental diets compared with larvae on the BioKyowa diet and the live feed control. Based on observations of intestinal fullness, the experimental diets appeared to be less palatable than the BioKyowa diet. As a result, it took longer to wean the larvae and higher mortality was experienced during weaning. Once successfully weaned, the experimental diets yielded growth rates equivalent to larvae feeding on the commercial diet for the remainder of the experiment.     

Effects of protein hydrolysate in weaning diets for Atlantic cod (Gadus morhua L.) and Atlantic halibut (Hippoglossus hippoglossus L.)

The study aims to test whether predigested dietary protein enhances the utilization of formulated diets at weaning, and also whether it stimulates intestinal maturation. In this study, Atlantic cod [ Gadus morhua L.; 41 days posthatch (dph)] and Atlantic halibut [ Hippoglossus hippoglossus L.; 63 days postfirst feeding (dpff)] were weaned onto diets with graded levels of protein hydrolysate. By increasing the inclusion of dietary protein as hydrolysate from 0 to 400 g kg⁻¹, cod increased the rate of survival from 7 ± 1% to 18 ± 2% (82 dph; regression, P =  4*10⁻⁷). In halibut, the survival rate decreased from 57 ± 9% to 22 ± 7% as the inclusion of protein in the form of hydrolysate increased from 0 to 450 g kg⁻¹ (119 dpff; regression, P =  8*10⁻⁵). Growth was not affected in any of the species. Results in specific activities of the intestinal enzymes leucine aminopeptidase (LAP) and alkaline phospatase (AP) supported the results in survival in halibut and partly also in cod, by showing increased activities in groups with increased survival ( anova , P <  0.05). The lower optimal level of hydrolysed protein in halibut than in cod is suggested mainly because of a slower feeding practice in halibut, which allows more extensive nutrient leaching before ingestion.     

Dietary carbohydrate utilization in cod (Gadus morhua): metabolic responses to feeding and fasting

Moist diets with increasing amounts of carbohydrate (0.5%, 10% and 21% on a dry weight basis) were each fed to duplicate groups of cod (initial weight 370 g) for 8 weeks, after which all groups were fasted for 4 weeks. Protein energy was high and accounted for more than 70% of the gross energy content in all feeds, and the diets were maintained isocaloric by substituting lipid energy for carbohydrate energy. No indigestible binder was added. Excellent growth and feed conversion were obtained in all groups. After 4 weeks of fasting, fish previously fed diets with either 10% or 21% carbohydrate showed significantly higher weight loss than fish fed the diet without carbohydrate. Liver glycogen reached 10% of liver wet weight in fish fed diets containing 10% or 21% carbohydrate and 5% in fish receiving 0.5% carbohydrate after 8 weeks. Following 4 weeks of fasting, liver glycogen was reduced to similar levels in all fish. Plasma glucose levels 4 h after feeding were higher in fish fed the diets with 10% or 21% carbohydrate and plasma free amino acid levels (FAA) were lower, than in fish fed the diet containing 0.5% carbohydrate. Blood lactate concentrations were unaffected during the first 24 h after feeding. After 4 weeks of food deprivation, the levels were significantly reduced only in the 21% carbohydrate group. A link between glucagon and protein metabolism is suggested because plasma glucagon concentration followed the same pattern as the concentrations of plasma FAA throughout the study. Insulin and glucagon-like peptide (GLP) showed a covariation throughout the experiment. Reduced plasma insulin levels were seen after fasting concomitant with reduction in the levels of FAA and glucose. It is suggested that insulin secretion in cod is affected both by plasma FAA and glucose and that cod meets food deprivation by slowing down metabolism.     

Optimized biophysical model for Icelandic cod (Gadus morhua) larvae

A characteristic of biophysical models (BPM) is that they contain a number of parameters that are poorly known or only known within a range of possible values. This paper describes an efficient optimized BPM developed to simulate the Icelandic cod pelagic 0‐group survey data. The method is based on presenting the results as a probability density function (PDF) that a particle released from a given spawning drifts downstream to a given grid location some time later. The model determines egg production model parameters (peak spawning time, spawning duration, number of eggs released) for each of 15 spawning grounds as the solution of a bound constrained optimization problem that minimizes model‐data misfits in abundance and age distributions. The model is applied to simulating the 2002 and 2003 summer survey data. The model does a reasonable job of simulating the observed inshore / offshore abundance gradient and spatial age gradient for each year. Problem areas are explained from the point of view of model limitations. We caution that the results from an optimized model should always be assessed with the model’s limitations in mind and with respect to whatever biological data are available.     

Digestive capacity and compensatory growth in Atlantic cod (Gadus morhua)

The objective of this study was to examine whether digestive capacity correlates with growth rate in Atlantic cod (Gadus morhua). Ninety fish (1035±240 g; 478±36 mm) were assigned to one of three treatments: deprived of food for 5 weeks and then re-fed for 24 days, deprived for 10 weeks and then re-fed for 24 days, or fed for 24 days without deprivation (controls). Three times a week during feeding, the fish were provided with meals of capelin (Mallotus villosus) in excess. Within each treatment, ten fish were sacrificed before and twenty after the feeding period. The relative masses of the pyloric caeca, intestine, and white muscle were determined. Cytochrome c oxidase (CCO), citrate synthase (CS), and nucleoside diphosphate kinase (NDPK) activities were assayed in the pyloric caeca, intestine, and muscle as was trypsin activity in the pyloric caeca. During the re-feeding period, fish that had been deprived of food for 10 weeks showed compensatory growth, growing 1.9 times faster than the controls (0.94±0.26 versus 0.50±0.31% body mass ⋅ day⁻¹). Fish that displayed compensatory growth had a lower relative white muscle mass than controls (66.5±4.1% vs. 69.6±7.4%) while the relative masses of the pyloric caeca (2.21±0.49% vs. 1.78±0.31%) and intestine (0.95±0.20% vs. 0.77±0.15%) were greater than the controls, suggesting that the sizes of these digestive tissues might correlate with compensatory growth capacity. Citrate synthase in the pyloric caeca was the only enzyme that showed higher activity during compensatory growth (8.57±0.94 U ⋅ g tissue⁻¹ compared to 7.13±1.03 U ⋅ g tissue⁻¹ in the control group). This suggests that aerobic catabolic capacity of pyloric caeca could be related to growth capacity during recovery, possibly via the energetic cost of digestive enzyme synthesis. This revised version was published online in July 2006 with corrections to the Cover Date.     

Inclusion of camelina meal as a protein source in diets for farmed Atlantic cod Gadus morhua

Camelina meal Camelina sativa (CM) is a potential protein source in aquaculture feeds, because of its crude protein level (39%) and essential amino acids. Two feeding experiments were conducted with Atlantic cod Gadus morhua. Cod in Experiment I (19.4 g fish^?1) were fed diets with 0%, 12% or 24% CM for 9.5 weeks at 10°C; and cod in Experiment II (14.4 g fish^?1) were fed diets with 0%, 15%, 30% or 40% CM for 13 weeks at 10°C. Growth, lipid and amino acid tissue composition were compared amongst cod fed varying levels of CM. In Experiment I, cod could tolerate the highest level of CM inclusion (24%) without affecting growth compared to cod fed the control diet. In Experiment II, growth performance was significantly affected at 30% CM inclusion compared to the control treatment, and cod fed 15% CM displayed some signs of depressed growth (reduced feed intake and weight gain). Both treatment and duration were interacting factors (P = 0.015) that determined growth performance when comparing both experiments. Muscle tissue composition was relatively unaltered with less than 30% CM inclusion; however, multivariate statistics revealed significant differences in muscle tissue fatty acid composition between cod fed 40% CM and the control diet. The tissue amino acid profile was generally unaltered because the dietary amino acid profile was consistent after CM inclusion. A few antinutritive compounds in CM may have affected palatability in diets with greater than 30% CM inclusion, which may have resulted in reduced growth performance.     

Territorial and agonistic interactions between farmed and wild cod (Gadus morhua)

Studies of contest competition between wild and farmed fish have mostly focused on fish with strongly territorial behaviour. Little is known about species with more plastic social behaviour, such as Atlantic cod (Gadus morhua L.), a species that can either aggressively defend territories or shoal. There is also concern that cod that escape from farms will compete with wild populations. We examined dyadic contest competition between wild and farmed juvenile cod using an intruder–resident experimental set‐up in the laboratory. No prior residency advantage was observed, but the differences between farmed and wild cod were clear. Farmed cod were more submissive than wild cod and fled earlier during contests, which suggests that wild fish often out‐compete farmed intruders. Both fish types initiated aggression earlier against fish of the same background. A multivariate analysis of 11 different behavioural traits indicated that a group of 59% of farmed fish were behaviourally very similar to the 55% most submissive wild fish. These results suggest that wild juvenile cod may be quite robust towards competition for food and shelter from juvenile farmed cod, but further research is needed to verify this pattern.     

Differences in growth and chemical composition between male and female farmed cod (Gadus morhua) throughout a maturation cycle

The growth of farmed cod fed to satiation under natural conditions in the sea in North Norway (67°N) was followed over 1 year. At the start of the experiment in September 2003, the average weight was 1300 g. After 1 year, in September 2004, the average weight was 2800 g. From the end of November to June, the fish matured and spawned and there was little or no weight gain in the fish during this time. The male cod developed gonads earlier than the females. In February, the male gonad index was 19% and decreased to 7% by the end of April, whereas the female gonad index increased from 14% in February to 25% in April. The male liver index decreased by 2% in February and by another 3% in April, when it only decreased by 1% in the female cod. The muscle protein in the female cod was reduced by more than 20 g kg^?1 in April, and the difference between the muscle protein in the male and the female cod was still significant in September. No differences were found in the post‐rigour muscle pH due to the sex of the fish.     

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 general biophysical model of larval cod (Gadus morhua) growth applied to populations on Georges Bank

Field-derived growth rates (RNA-DNA based) of cod (Gadus morhua) larvae collected on the southern flank of Georges Bank were higher on average in May 1993 than May 1994, despite the apparent higher abundance of potential prey in 1994. A biophysical modeling study is presented here in which factors are examined that may have led to the difference in population mean growth. A one-dimensional physical model, forced by winds and tides, was used to simulate the vertical structure (of currents, temperature field, and turbulent kinetic energy dissipation rate) following a column of water in a Lagrangian sense at a site on the southern flank of Georges Bank over 5-day periods in late May of 1993 and 1994. The biophysical model and observed zooplankton abundance allowed us to explore the vertical structure and temporal (hourly) evolution of feeding and growth for cod larvae in relation to environmental conditions. Our trophodynamic model is improved over previous versions and now includes the effect of light on larval feeding response, as well as the effect of temperature on larval metabolic costs, ingestion, and digestion. Larval prey profiles, comprising four copepod species, were used from a time series of 1/4-m² MOCNESS tows to define the prey field. Data from a collateral time-series of larval gut contents (1-m² MOCNESS tows) was used to define maximum ingestion (satiation level) and prey selection. Model outputs provide depth-dependent estimates of growth, prey biomass ingested, larval length, and larval weight. Water-column growth-rate profiles were made for four size classes of larvae (5, 6, 7 and 9 mm) under the environmental conditions observed in May 1993 and 1994. A weighted-mean growth rate based on the mean vertical distribution of larvae was estimated for each size class. In all cases, when using all available potential prey, the model-derived 1994 growth rates were higher (by 3–6% day⁻¹) than those for 1993. However, simulations in which 7-mm larvae followed the field-derived weighted mean depth over the sampling period, and were limited to their preferred Pseudocalanus prey, resulted in average growth of 12.2% day⁻¹ for 1993 and 9.7% day⁻¹ for 1994. These compared closely to the field growth means of 11.3% day⁻¹ in 1993 and 9.8% day⁻¹ in 1994. Thus, the lower observed growth in May 1994 may have resulted from depth-dependent food limitation and prey-selectivity coupled with the greater metabolic costs induced by the higher temperature that year.     

Linking cod (Gadus morhua) and climate: investigating variability in Irish Sea cod recruitment

How climatic variability and anthropogenic pressures interact to influence recruitment is a key factor in achieving sustainable resource management. However, the combined effects of these pressures can make it difficult to detect non‐stationary interactions or shifts in the relationships with recruitment. Here we examine the links between climate and Irish Sea cod recruitment during a period of declining spawning stock biomass (SSB). Specifically, we test for a shift in the relationship between recruitment, SSB and climate by comparing an additive (generalized additive model, GAM) and non‐additive threshold model (TGAM). The relationship between recruitment success, SSB and the climatic driver, sea surface temperature, was best described by the TGAM, with a threshold identified between recruitment and SSB at approximately 7900 t. The analysis suggests a threshold shift in the relationship between recruitment and SSB in Irish Sea cod, with cod recruitment being more sensitive to climatic variability during the recent low SSB regime.     

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 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.     

Effects of replacing fish meal and oil with plant resources in on‐growing diets for Atlantic cod Gadus morhua L

The present paper gives an overview on the use of plant protein and plant oils as replacers for fish meal and fish oil in diets for Atlantic cod Gadus morhua L. In focus are effects on growth, feed utilization, digestibility, gut health, muscle and liver uptake and retention of nutrients, and muscle quality. Plant oil can replace fish oil without affecting growth provided that the requirement of marine long chain (LC) n‐3 fatty acids is met, but the altered dietary fatty acid profile in diet will be reflected in both muscle and liver. This can reduce the value of cod liver as an oil source for cod liver oil production. For the fish itself, there are more challenges replacing fish meal than fish oil, due to the amount of fibre and antinutrients in plant protein meals. However, A. cod seems to tolerate a wide range of plant types and their inclusion levels provided that the amino acids requirements are met. It is our view that there is sufficient knowledge to be able to design an A. cod diet based on a mixture of plant and marine ingredients and be able to predict performance such as growth, feed utilization, digestibility, liver size and fish health in general.     

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.     

The effects of feeding frequencies on seasonal changes in growth rate and chemical composition of farmed cod (Gadus morhua)

Atlantic cod (Gadus morhua) were fed with dry feed, in sea net pens from May 2003 to September 2004, under natural conditions occurring in Northern Norway, with a sea temperature varying from 3.3 ° to 16.6 °C. The fish was either fed five times or three times a week, once a day to saturation until December 2003, and after that five times or twice a week, once a day to saturation until September 2004. Fish fed five times a week had a higher FCR 1.47 compared with 1.35 for fish fed three to twice a week for the entire period. No significant differences were found in growth or chemical composition in the cod due to the feeding intervals. The best growth of the fish was in the first summer from May to September 2003 with a specific growth rate (SGR) of 0.81% per day. From June to September 2004, the SGR was 0.42% per day. From December to May, the SGR was negligible caused by low‐water temperature, darkness, and sexual maturation. The liver index increased from 7% to a maximum of 16% during the growth period except during the sexual maturation period from November to April, where it shows a slight decrease. Maturation and seasonal variation are the factors having the largest influence on the growth and chemical composition of the cod.     

Greenland cod (Gadus morhua): modeling recruitment variation during the second half of the 20th century

Two approaches were used to qualify observed variability in Greenland cod (Gadus morhua) recruitment. In the first analysis, we used the linear trend of the Greenland cod recruitment time series and climatic variables, such as air temperatures from the Denmark Strait and wind conditions off East Greenland and Southwest Greenland, to explain the interannual variation in cod recruitment off Greenland. The model resulting from this ‘trend/environmental approach’, explained 79% of the interannual variation in cod recruitment off Greenland. In the second, analytical approach, the ‘regime approach’, multiple linear regression models were used, with the input data being the time series of cod recruitment and spawning stock biomass (SSB) from Iceland and Greenland, sea surface and air temperatures around Greenland, and zonal wind components between Iceland and Greenland. Model results indicated that, during the decades between 1950 and 1990, there were three different cause–effect regimes which significantly influenced the variability of cod recruitment. The three regimes included: (a) the 1950s and 1960s, a regime with favorable sea surface temperatures and a self‐sustaining cod stock off Greenland with high SSB that produced a series of above‐average, strong year classes; (b) the 1970s and 1980s, a regime of declining SSB and recruitment, with recruitment dependent on advection from Iceland; and (c) the 1990s, when the advective potential for recruitment from the Icelandic cod stock was the only available source for replenishment of the Greenland cod stocks, because cod recruitment in Greenland waters was negligible. The three models explained 76–77% of the observed interannual variation in cod recruitment off Greenland. Both approaches suggested that advective factors were the dominant influences for cod recruitment in the ‘Iceland–Greenland System’.     

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.