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.     

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.     

Thermal shock induction of triploidy in Atlantic cod (Gadus morhua L.)

The effects of thermal treatments on induction of triploidy in Atlantic cod have been investigated. Cold shock [−1.7±0.1°C at 20 min post fertilization (PF) for 2 h] was based on a previously developed protocol, and heat shocks, below the lethal threshold of 24°C, were at 16, 18 or 20°C applied 20, 30 or 40 min PF for 20 min. Cold shock did not affect larval survival and was ineffective for producing triploids (range 0–4%). A heat shock of 20°C at 20 min PF generated the highest percentages (range 66–100%) of triploid larvae at hatching, with survival ranging from 10% to 20% relative to the controls. Lower heat shock temperatures or delayed shocks increased survival but decreased the number of triploids, providing no net gain in triploid yield (range 1–9%). Heat shocks applied later than 20 min PF produced 2–4% tetraploid larvae at hatching. A thermal shock of 20°C initiated at 20 min PF and lasting 20 min proved to be the most generally efficient treatment for induction of triploidy in Atlantic cod.     

Tracking escaped Atlantic cod (Gadus morhua L.) aggregated around Norwegian farms: Considerations for management strategies

To examine site fidelity and potential for ecosystem impacts, 20 escapee cod (Gadus morhua L.) from fjord‐based cod farms were tagged with acoustic transmitters, released in the vicinity of the same farm, and their subsequent behaviour and survival monitored using a fixed array of acoustic receivers. Most individuals demonstrated high fidelity to their farm where tagged, and this increased during the polar night. Only five individuals (25%) were occasionally detected at another farm (4 km distance), and only three cod (15%) migrated out of the fjord during the 6‐month study period. Vertical movements were consistently greater in the hours around noon than at night, a behaviour inconsistent with that observed for wild fish. A high mortality rate (40%) was observed during the first 4 months post‐release, and actual mortality may have been even higher, limiting the potential for negative ecosystem effects from cod escapees. Findings from this study are used to discuss the development of recapture strategies for minimising potential impacts from cod escapees to ensure sustainable farming activity.     

Spawning‐related mortality in captive Atlantic cod (Gadus morhua L.)

Two year‐classes of Atlantic cod were reared in indoor tanks from November 2002 to November 2004. The average annual mortality was 21% and 36% for males and females respectively. The highest monthly mortality rates in females were observed around and during the spawning periods from March to July, maximally 7% and 20% for year‐class 2001 and 18% and 34% for year‐class 2000. The mortality in males was less dependent on season, reaching a maximum of 7% and 5% month^?1 for year‐class 2001 and 2000 respectively. The maximum mortality during spawning increased with age from: 7%, 19% and 34% per month for 2‐, 3‐ and 4‐year‐old females respectively. The average gonadosomatic indices in 3‐ and 4‐year‐old mature females that died during the spawning periods were 22% and 26% respectively and most of them were spawning when they died. The results suggest that egg‐retention may have been the main cause of death in females during spawning.     

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.     

Vertical distribution and sexual maturation in cage‐farming of Atlantic cod (Gadus morhua L.) exposed to natural or continuous light

The vertical distribution and seasonal timing of sexual maturation were investigated in Atlantic cod in four large commercial sea cages in Northern Norway during their second year of on‐growth. Replicate cages were exposed to either natural light (NL) or continuous light (LL) from May 2007 to January 2008. Hydro‐acoustic monitoring revealed that the cod in the NL cages were mainly distributed from 5 to 13 m depth at day while they were more dispersed and swam deeper at night. A clear ascent was observed prior to feeding events. The cod in the LL cages displayed a similar seasonal and daily pattern, but with a more dispersed vertical distribution range at both day and night. It is hypothesized that surface feeding motivation resulted in cod preferentially occupying the upper reaches of the sea cage, generally at depths < 13 m. This would also reduce the need for excessive alterations in swim bladder volume. Both the LL and the NL groups were exposed to LL the year before the study, resulting in first time sexual maturation (puberty) being delayed until the summer in both groups. However, while cod in the NL cages matured during the autumn 2007, the LL cod did not.     

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.     

Effects of pre-rigour filleting on quality aspects of Atlantic cod (Gadus morhua L.)

Aquaculture is promising in increasing the amount of fresh Atlantic cod available all year around. The aim of this study was to investigate the effects of pre‐rigour filleting on fillet quality of wild and farmed cod. Pre‐rigour filleting resulted in significantly less fillet gaping than in fillets produced post rigour. The pre‐rigour fillets were, however, 12–13% shorter post rigour. In spite of the significantly lower water content of the muscle 6 days post mortem, a significantly higher liquid loss was seen in the pre‐rigour group. Differences in gaping score and muscle pH were dependent on whether the fish were wild or fed. Fillet gaping increased with storage time and was particularly pronounced for well‐fed cod. Fed cod had a significantly higher hydroxyproline (Hyp) content and significantly lower amount of glycosaminoglycans (GAGs) than wild cod. The ratio between GAGs and Hyp was thus significantly lower in fed cod. A connection between connective tissue components and fillet gaping may exist. However, further investigations are required.     

Effect of Antarctic krillmeal on quality of farmed Atlantic cod (Gadus morhua L.)

Farmed Atlantic cod with a mean weight of 4.8 kg were maintained for 9 weeks in sea cages and fed diets where the dietary fishmeal component was substituted with increasing proportions (0%, 22%, 63% and 100%) of meal from Antarctic krill, Euphausia superba. Wild‐caught cod were included in the study as external control. At termination of feeding, all fish were slaughtered and muscle pH was immediately recorded. The fish were then stored on ice for three days and assessed for muscle pH and objective (skin and fillet colour, and fillet texture) and subjective (sensory evaluation) quality criteria. Replacement of fishmeal with krillmeal in the diets resulted in the skin colour above and below the lateral line to be more red than the control group without krillmeal substitutions, even though this difference was not significant, and with more yellow hue. Additions of krillmeal increased the muscle whiteness and yellow hue compared with the control group and wild fish. There was no difference in red hue between the groups. Muscle pH, texture or sensory attributes were unaffected by dietary inclusion level of krillmeal. Wild‐caught cod deviated in several aspects from the farmed cod. It is concluded that the replacement of fishmeal with Antarctic krillmeal in the diets two months before slaughter did not move the sensory attributes more towards wild fish.     

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.     

Effects of filleting method,stress, storage and season on the quality of farmed Atlantic cod (Gadus morhua L.)

The effects of filleting method, pre‐slaughter stress, storage and season on drip loss, water content, water‐holding capacity, rigour contraction and colour in Atlantic cod, Gadus morhua L., were examined. A total of 182 fish were sampled under commercial conditions. To test the effects of filleting method and stress, stressed (S) and unstressed (C) fish were filleted either pre (Pr) or post rigour (Po) at 1 and 48 h post mortem, receptively. The muscle pH significantly decreased from 0 to 144 h of storage in all groups. The onset of the rigour contraction was more pronounced in the Pr‐S group as compared with the Pr‐C group, but after 144 h of storage, no difference in degree of contraction was observed. Filleting method, stress and storage time influenced the colour of the fillets. Post‐rigour filleting caused a significantly increased in measured Lightness (L*) during storage. Stress caused a significant increase in measured redness (a*). No significant difference was found for water‐holding capacity. The water content was influenced by filleting method, where the Pr‐S group had a significantly lower water content compared with the Pr‐C groups; water content changed also according to season. The findings of this study show that stress, filleting method, time of storage and season had a significant effect on the drip loss, where a combination of all factors will determine the total loss.     

Application of prospective probionts at early stages of Atlantic cod (Gadus morhua L.) rearing

This work aimed at validating the use of two prospective probionts (Arthrobacter sp. and Enterococcus sp.) at early stages of cod (Gadus morhua L.) rearing. Ova at late post‐fertilized stage and larvae during their first 4 weeks of life were bathed with both probionts, isolated previously from the cod‐rearing environment. This treatment was compared with groups fed rotifers supplemented with a commercial probiotic (Remus^®) and those untreated. Microbiological analyses (total viable counts, presumptive Vibrio and lactic acid bacteria) were performed in rearing systems and larval survival, growth and development were assessed. Larval development was evaluated by proteolytic activity of larval lysates and immunological analysis of important proteins: apolipoprotein A‐I, haemoglobin, C‐reactive protein, C3 and cod serum proteins. Bacterial bathing led to a significantly higher larval weight, length and culturable microbial load in larval gastrointestinal (GI) tract when compared with the control and Remus groups. Development occurred earlier in bathed larvae. However, their survival was negatively affected compared with the control group, but was significantly higher than for the Remus group. The non‐pathogenicity of both probionts was demonstrated by intraperitoneal injection of 13 g cod juveniles. The results suggest that Arthrobacter and Enterococcus probionts affected the larval GI microbiota and contributed to growth, development and digestion, either directly or indirectly.     

Pre-rigor filleting and drip loss from fillets of farmed Atlantic cod (Gadus morhua L.)

Improved slaughtering procedures of farmed fish may provide sufficient time so that filleting can be performed pre‐rigor while the muscle pH is still high. Such filleting not only reduces fillet gaping but also lowers the transportation costs and makes fresh fillets available to the markets at an earlier stage. The aim of our work was to determine the weight reduction of the fillets due to liquid loss and to study the proteins and enzymes in the drip. After 11 days of cold storage, fillets of farmed Atlantic cod produced pre‐rigor had a weight loss of 10% and a fillet contraction of 19% while for fillets produced post‐rigor the values were 5% and 4% respectively. At the same time, approximately twice the amount of proteins had been lost from the pre‐rigor‐produced fillets. Sodium dodecyl sulphate polyacrylamide gel electrophoresis analysis showed that the proteins in the drip were similar to the sarcoplasmatic proteins extracted from the muscle. Specific analysis of proteolytic enzymes indicated that they are less stable in the expelled liquid than in the flesh during storage. The extensive loss of weight and proteins from pre‐rigor‐produced fillets during subsequent storage must be taken into account if such processing is considered for farmed cod.     

Seasonal spawning and wind‐regulated retention‐dispersal of early life stage Atlantic cod (Gadus morhua) in a Newfoundland fjord

Sequential ichthyoplankton surveys were used to determine the spatial and temporal distribution of eggs and larvae over coastal spawning grounds of Atlantic cod (Gadus morhua) in Smith Sound, Trinity Bay, Newfoundland, during the spring and summer of 2006 and 2007. Egg densities showed similar patterns for both years with two peaks in abundance in spring (March–April) and late summer (late July). A clear progression of development stages (1–4) was observed in spring and summer in 2006 and summer in 2007, suggesting retention of eggs within the Sound during these periods. Modelled predictions of vertical egg distributions indicated eggs were broadly distributed near the surface during spring (March–April), but were concentrated below the pycnocline (>10 m) within the inner portions of the Sound during the summer months (July–August). Back‐calculated peaks in spawning based on water temperatures were estimated at 11 and 4 April for 2006 and 2007, respectfully, with late season peaks centred on 21–24 July for both years. Environmental data indicated cooler water temperatures and periods of high wind stress in spring, and warmer, calmer periods late summer, consistent with higher retention and faster development times on the spawning grounds later in the season. We conclude that spring and summer spawning events result in different distributions of early life stages and may lead to different distributions of juvenile and adult fish.     

Investigation of the temporal effects of spawning season and maternal and paternal differences on egg quality in Atlantic cod Gadus morhua L. broodstock

A better understanding of the parameters affecting egg quality and larval survival is of importance for continued development of cod broodstock and efficient husbandry practices. Decision tree analysis (DTA) was applied to analyse 3 years of egg quality data in an effort to extract the most important variables (i.e. predictors) in explaining differences in egg quality. The effect of three predictors (spawning time, maternal and paternal differences) has been studied on early cleavage pattern parameters, egg diameters, fertilization and hatching rates and has shown that females are the dominant variable and that time has a limited and inconsistent impact on the data. When using maternal, paternal differences and batch number (instead of spawning time) as predictors, the results confirm that no particular relationship is found between batch order (i.e. order in time) and egg quality. Moreover, batches with a higher egg quality show a consistency in the parameters assessed (i.e. batches with higher rates of normality in any parameter tend to be normal for other parameters). This is confirmed by the significant correlations found between cleavage parameters. Our results highlight that spawning time is of less importance than female parent contribution in ensuring high rates of fertilization and larval hatch, and maximizing general egg quality.