Reconstructing migrations of individual cod (Gadus morhua L.) in the Baltic Sea by using electronic data storage tags

A new methodology is presented to reconstruct migration pathways of individual fish inhabiting ecosystems with moderate‐to‐strong gradients in temperature or salinity. The method uses measurements of ambient pressure, temperature and salinity obtained from electronic data storage tags attached to individual fish and is particularly applicable in areas with negligible tides. We demonstrate the method with Baltic cod. Hydrographic fields obtained from hydrodynamic modelling were used as a geolocation database to identify daily positions of Baltic cod by comparison with the environmental data collected by the tags. Using randomly distributed individual parameter perturbations in the range of the instrument precision of the tag we simulated a cod migrating through the Baltic Sea. The distance between the prescribed and geolocated positions of this artificial cod was on average 2.9 ± 4.7 (SD) km. Subsequently, the method was used to reconstruct migration routes of 10 real cod tagged in the Bornholm Basin of the Baltic Sea in early spring 2003. Differences were compared between the tag data and the geolocation database. The uncertainty in geolocation at recapture day was on average 75 ± 23 (SD) km, as shown by comparison between geolocated position and recapture position.     

Chemical surface disinfection of eggs of Baltic cod,Gadus morhua L.

The effect of two disinfectants on eggs and larvae of Baltic cod, Gadus morhua, was investigated. The eggs were disinfected for 10 min using various concentrations of either glutaraldehyde (100, 200, 400, 600 and 800 mg L^?1) or iodophor (10, 50, 100 and 150 mg L^?1), 1–4‐days post‐fertilization. Bactericidal effect of disinfection, survival to hatching, hatching success and larval abnormalities were assessed. Larval survival was recorded at 5‐, 10‐ and 15‐days post‐hatch (dph). Although Baltic cod eggs have an unusually thin chorion, they could tolerate surface disinfection. A reduction in bacterial growth was observed with increased concentrations of disinfectant (3.0 × 10⁷–1.6 × 10¹ CFU mL^?1). Abnormalities in newly hatched larvae were not related to disinfection. Survival of the yolk sac larvae was significantly better for eggs treated with 400 mg L^?1 glutaraldehyde for 10 min at 10 and 15 dph. Effective disinfection was also recorded using 100 mg L^?1 Actomar K30. Egg batch effect rather than initial bacterial concentration, disinfectant type or incubation method determined the survival of the eggs to hatching and survival of larvae. Because of the carcinogenic effect of glutaraldehyde, iodophor is recommended for routine disinfection of cod eggs.     

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.     

Variations in the drift of larval cod (Gadus morhua L.) in the Baltic Sea: combining field observations and modelling

Coupled three-dimensional (3-D) physical oceanographic modelling and field sampling programmes were carried out in May 1988 and August 1991 to investigate the potential drift of larval cod ( Gadus morhua L.) in the Bornholm Basin of the Baltic Sea. The goals were to predict the transport of cod larvae, thus aiding the identification of physical processes influencing larval retention/dispersal. Numerical simulations were performed using a 3-D eddy-resolving baroclinic model based on the Bryan–Cox–Semtner code adapted for the Baltic Sea. Within the Bornholm Basin, the model was initialized with ground truth data of physical parameters obtained on the research cruises, and all simulations were forced with actual wind data. Outside the basin, generalized hydrographic features of the Baltic Sea were utilized by incorporation of simulated hydrographic fields from previous model runs typical for the time periods considered. Larval drift was simulated either by incorporation of passive drifters, or as the initial horizontal distribution of larvae implemented into the model. Drift model simulations of larval transport agreed relatively well with field observations. The influence of variations in the vertical distribution on a smaller scale, i.e. vertical deviations of ± 6 m from the observed mean centre of mass, on the drift was examined, revealing no significant differences in the drift of larvae depending on their vertical distribution. The different wind forcing during the investigated time periods was linked to a retention situation in May 1988 and to a dispersal situation in August 1991. Finally, observed spatial distribution patterns of 1-group cod based on Baltic Young Fish Surveys (BYFS) were compared with their predicted transport in the larval phase and examined with respect to recruitment.     

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

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

Process-based model for direct and indirect effects of hydrographic conditions on Central Baltic cod (Gadus morhua) egg mortality

A process‐oriented model for the mortality of eggs of cod Gadus morhua in the Central Baltic Sea is developed, based on a synthesis of existing knowledge of the effects of salinity, oxygen and predation by sprat Sprattus sprattus. The models show the importance of the vertical and temporal overlap between eggs and predations. Effects related to the changing buoyancy of the eggs due to age and size of the mother fish, batch number and stock structure are not of major importance for the egg survival of this stock. It is demonstrated that under the present high sprat predation pressure, the observed delay in spawning time has increased egg survival.     

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.     

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.     

The pathology of the early stages of the crustacean parasite, Lernaeocera branchialis (L.), on Atlantic cod, Gadus morhua L

Lernaeocera branchialis is a penellid copepod, the adults of which are commonly found on gadoids in the North Atlantic and North Sea. L. branchialis has a severe negative impact on wild gadoids and is a potentially serious pathogen of farmed cod, Gadus morhua. This paper describes an experimental infection by cohabiting the intermediate host, the flounder, Platichthys flesus, heavily infected with juvenile stages of L. branchialis, with hatchery-reared cod in order to study the pathology of the early stages of infection of the final host. Early stages of L. branchialis induce extensive local gill hyperplasia, large intravascular thrombus formation and a moderate cellular immune response in the cardiac and branchial tissues of G. morhua. The parasite appears to migrate within the lumen of the afferent branchial artery, ventral aorta and in turn, that of the bulbus arteriosus. These findings are consistent with the presence of a haematophagous, voluminous and rapidly invasive parasite.     

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.     

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.     

Oceanographic influences on the distribution of Baltic cod, Gadus morhua, during spawning in the Bornholm Basin of the Baltic Sea

The Baltic Sea is a stratified, semienclosed sea typified by a low-salinity surface layer and a deep saline layer of varying volume, salinity, temperature and oxygen concentration. The relationships between these oceanographic factors and the distribution of Baltic cod are presented, utilizing results from a survey carried out during the 1995 spawning period in the Bornholm Basin, at present the main spawning area of this stock. Cod distribution, abundance and population structure were estimated from hydroacoustic and trawl data and related to hydrographic parameters as well as to bottom depth. In the central basin, cod were aggregated in an intermediate layer about 15 m thick. This area of peak abundance was defined at its upper limit by the halocline and at the lower limit by oxygen content. The majority of individuals caught in the basin centre were in spawning or pre-spawning condition with a high proportion of males to females. On the basin slopes, aggregations of cod were found near the bottom. These individuals were mainly immature and maturing stages with an increasing proportion of females to males with size. Salinity and oxygen conditions were found to be the major factors influencing the vertical and horizontal distribution of adult cod. Abundance of immature cod was also positively related to decreasing bottom depths. The effect of temperature was minor. The observed size- and sex-dependent spawning aggregation patterns, in association with habitat volume and stock size, may influence cod catchability and thereby the assessment and exploitation patterns of this stock.     

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.     

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.     

The influence of temperature on advective loss of Atlantic cod (Gadus morhua) eggs from the inshore environment

We use a simple model of temperature-dependent egg development and mortality to develop several hypotheses concerning the effect of temperature on the occurrence of eggs of Atlantic cod, Gadus morhua , in Placentia Bay, Newfoundland. Predictions from this exploratory model were tested through a series of synoptic ichthyoplankton surveys throughout spawning and postspawning periods during 1997 and 1998. Although several egg mortality relationships were explored through the simulation, a constant mortality rate best represented the pattern observed in the two years of data. Peaks in late stage egg densities occurred in August of both 1997 and 1998 and were apparently decoupled from egg production peaks in April. We observed a decrease in mortality and the distance dispersed during egg development with increases in water temperature. We suggest that the effects of predation are small relative to the advective effects within this system, and that the interaction between advection and temperature-dependent vital rates of eggs may have dramatic consequences for coastal retention of propagules produced by inshore spawning events.     

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.     

The fate of Lernaeocera branchialis (L.) (Crustacea; Copepoda) in Atlantic cod, Gadus morhua L

Lernaeocera branchialis, a copepod crustacean parasite of gadoids, represents a potential threat to both wild and farmed cod, Gadus morhua. The pathological changes associated with the early stages of experimental infection have previously been reported in detail, and this article describes the lesions associated with later chronic stages of experimental infection. Chronic infection is characterised by extravascular granuloma formation and proliferation of fibrovascular tissue around intact and fragmented, degenerate parasites within both the gill arch and cardiac region. The majority of parasite granulomas are located within connective tissues of the gill arch or pericardium; however, low numbers are present within the wall of large vessels. The intraluminal parasites and thrombi of early stage infection are largely absent in these later lesions. We propose that organisation and incorporation of the parasite thrombus into the vessel wall with subsequent granuloma formation and extrusion into the surrounding connective tissue leads to the elimination of the parasite from the vascular system. Thus, rather than being a negative consequence of infection thrombosis is protective, allowing the host to survive the substantial initial vascular insult.     

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.     

Environmental influences on melatonin secretion in Atlantic cod (Gadus morhua L.) and their relevance to commercial culture

This study investigated the effects of photoperiod and temperature on plasma melatonin secretion in Atlantic cod (Gadus morhua L.). Initial work confirmed the presence of a diel profile of melatonin synthesis, with elevated levels during the dark phase. Unusually for fish, the peak in plasma melatonin occurred towards the end of the dark phase, which is indicative of a type `A' melatonin profile. When exposed to 60 hours of continuous darkness a clear endogenous rhythm of melatonin synthesis was observed, which continued for 4 cycles with a periodicity which, approximated to 24 h. When acclimated to varying temperatures (4, 8, 12 or 16 °C) no variation in melatonin production was seen, however, body size appeared to be an important influence, with the smallest fish exhibiting significantly higher levels of dark phase melatonin. Finally, the application of additional night-time illumination to cod maintained in sea cages i.e. without blackout, did not significantly reduce dark phase plasma melatonin, suggesting that cod are less sensitive to photoperiod manipulation in cages than salmonids.