Spermatocrit and spermatozoa density in Atlantic cod (Gadus morhua): correlation and variation during the spawning season

The utility of spermatocrit (the proportion of solid packed material in semen after centrifugation) as an indicator of spermatozoa density and male spawning stage was tested in Atlantic cod (Gadus morhua). Semen was collected from captive male cod over three spawning seasons. Spermatocrit was positively and significantly correlated with spermatozoa density measured with a Coulter counter (Multisizer), but not with spermatozoa counts in a haemacytometer. Spermatocrit increased significantly as the spawning season progressed. However, day of season explained only 35% of the variation because spermatocrit varied among individual males. Spermatozoa size remained unchanged throughout the sampling period and was not correlated with spermatocrit, indicating that variation in spermatocrit was due to variation in spermatozoa number and not their size. Spermatocrit is a good estimator of sperm density but is not reliable as indicator of spawning stage because of the variation among individual males.     

Discrimination between Icelandic cod (Gadus morhua L.) populations from adjacent spawning areas based on otolith growth and shape

Studies on Atlantic cod (Gadus morhua L.) have demonstrated extensive variation in life history characteristics among populations exposed to different environmental conditions. In Iceland, cod sampled from adjacent spawning areas, within the main spawning area at the southwest coast, exhibit variation in vital life history characteristics. In this study, we investigated the stock structure of Icelandic cod on the main spawning grounds to discriminate between these adjacent spawning groups by using otolith growth and shape. Otolith growth was based on annual increment width estimated for the major age groups (6–7 year olds) of the spawning stock. Otolith shape was determined using Fourier analysis and compared among cod sampled from the different spawning areas with stepwise canonical discriminant analysis (CDA). We found significant differences in growth and otolith shape between adjacent spawning groups of cod where those sampled close to the coast differed from those that were sampled further out on the bank and continental shelf. Our results suggest that these cod may belong to different populations and the large and fast growing cod spawning in the coastal area may need special protection given their significance to the overall productivity of the stock.     

Larval group differentiation in Atlantic cod (Gadus morhua) inside and outside the Gullmar Fjord

The spatial and temporal occurrence of pelagic fish stages and their biological variability may affect their dispersal and survival, and ultimately fish recruitment. We collected Atlantic cod larvae at one station inside and at one station outside the Gullmar Fjord, eastern Skagerrak, in order to investigate small-scale larval group differentiation. Rectangular midwater trawl hauls were taken every 6 h (during 24 h) from three separate depth intervals between the surface and 70 m depth. About 80% and 20% of all larvae occurred above the halocline at the Fjord station and the Coastal station, respectively. Hatching (at both stations) occurred from the 3rd week in February to the 1st week in May, indicating that cod larvae were present for at least 5 months (from late February to early August). The length and hatch date frequency distributions of larvae from the surface layer were unimodal inside the fjord but bimodal outside the fjord. Analyses of seven microsatellite DNA loci indicated that larvae collected inside the fjord (where local spawning occurs) were genetically distinct from larvae sampled on the outside (F_ST = 0.0026). The two age cohorts outside the fjord were not, however, genetically different, nor were larvae collected at different depths. We conclude that small-scale variability of vertical concentration and larval life history variability should have consequences for interpreting models of larval dispersal and survival, and subpopulation structure analyses.     

The spatial distribution of cod (Gadus morhua L.) spawning grounds in the Kattegat, eastern North Sea

Similar to many other commercial marine fish species, Atlantic cod (Gadus morhua L.) migrate towards specific sites at spawning. These temporal aggregations are generally the most targeted by the commercial fishery. The Kattegat cod has undergone a substantial reduction over the past 25 years and both stock size and spawning stock biomass have remained at very low levels since the end of the 1990s. There is, therefore, an urgent need to map and document spawning grounds still in use. In the present study, spawning sites of Atlantic cod were identified in the Kattegat using a combination of commercial and fishery independent data from 1996 to 2004. Moreover, putative spawning and non-spawning areas were also sampled before and during the reproductive season between 2002 and 2006, and the proportion of mature females together with the individual physiological status were used to validate and strengthen the spatial analyses.

The spatial analyses identified several spawning areas in the region and two areas in the southeastern part of the Kattegat which appeared to be the most important. The results showed the presence of cod spawning aggregations, although reduced in size, in areas that have been utilized for more than 25 years according to historical information. Some local spawning grounds may have also disappeared. The proportion of mature females was higher in putative spawning than in non-spawning areas (p < 0.001) and females from spawning areas had higher gonadosomatic (p < 0.05) and hepatosomatic (p < 0.001) indices than those from non-spawning areas.

Knowledge of stock spatial and temporal distribution is essential in designing recovery strategies for depleted fish populations. The unambiguous stability of the locations of spawning aggregations over time, as shown in this study, represents a useful aid in order to efficiently implement a recovery plan for the collapsing cod stock in this area.     

Involvement of trypsin and chymotrypsin activities in Atlantic cod (Gadus morhua) embryogenesis

Proteases play a key role in yolk formation and degradation during embryogenesis of marine fish. This study presents the first clear data on the involvement of trypsin and chymotrypsin activities in the embryogenesis of Atlantic cod (Gadus morhua). Both enzyme activities were shown to be present in unfertilized eggs followed by a significant decline (P < 0.01) in trypsin activity during the first 4 days post fertilization (dpf). Thereafter, the trypsin- and chymotrypsin activities increased to a maximum around day 9 pf. A decline in trypsin and chymotrypsin activities was observed from day 10 pf with minimal activity just prior to first feeding (day 15 pf). Western blot analysis, using polyclonal antibodies raised to Atlantic cod trypsins I and Y, mostly coincided with the trypsin activity profile. The novel trypsin Y was previously shown to have both trypsin- and chymotrypsin-like activities. Thus, some of the chymotrypsin activity observed in the samples may be originated from trypsin Y. The low trypsin and chymotrypsin activities just prior to first feeding (13–15 dpf) may indicate insufficient digestive function as trypsin has been shown to be a suitable short-term indicator reflecting the nutritional quality of marine fish larvae.     

Short-term crowding stress in Atlantic cod, Gadus morhua L. modulates the humoral immune response

Considering overcrowding as a stressor, the effects of a short-term exposure on the immune responses of Atlantic cod, Gadus morhua L. were determined at 2, 24 and 72 h post-crowding. While plasma glucose and total antioxidant capacity were quantified as stress responses, immune responses were based on humoral indices. Bactericidal capacity of plasma was also assessed. There was a significant increase in plasma glucose and total antioxidant capacity at 2 h post-crowding, however the former returned to pre-stress levels 24 h later and the latter remained elevated until 72 h. The plasma protein did not reveal any significant change, but there was a transient upregulation of globulin at 24 h post-crowding. The myeloperoxidase activity increased at 2 h after the stress, but returned to its pre-stress levels at 24 h. The lysozyme activity was upregulated even from the first time point after stress, although the differences were significant only at 24 and 72 h compared to the activity prior to stress. At 72 h post-crowding there was a significant increase in alkaline phosphatase activity while haemolytic activity showed a significant decrease. An increase in the proliferation of the bacterial pathogens, Vibrio anguillarum and Aeromonas salmonicida was seen from 24 h in the plasma of the stressed fish. Thus, our finding suggests that short-term crowding stress modulates the immune responses in Atlantic cod and therefore husbandry procedures should consider these effects while addressing better management strategies.     

Variation in fish condition between Atlantic cod (Gadus morhua) stocks, the effect on their productivity and management implications

A comparative analysis of the fish condition (Fulton’s K) of 10 cod stocks in the north Atlantic in relation to the temperature of their habitat, growth rates and their reproductive potential is presented. It is shown that the cod stocks in the north Atlantic display different levels of mean condition, which is partly due to the different temperature regimes of their habitats. Cod living in colder waters, e.g. Greenland, Labrador and Grand Bank stocks, were found to be in poorer condition than cod living in warmer waters, e.g. North Sea and Irish Sea stocks.

Poor condition causes reduced productivity in terms of slow growth and low recruitment potential. Stocks in better condition display significantly higher weights at age 4 than stocks in poor condition. The a coefficients (function’s slopes) obtained from standardised Ricker’s recruitment–spawning stock biomass (SSB) relationships were defined as indicators for the recruitment potential of stocks. These a coefficients were found to be positively correlated with the mean condition factor of the 10 stocks analysed. This indicates that stocks consisting of individuals in poor condition appear to be very susceptible to reduced recruitment at low SSB, while the stocks that consist of fish in good condition seem to behave more robustly with a higher probability of good recruitment at low SSB. The positive effect of the cod condition on their reproductive potential generally implies that the stocks in good condition in the temperate regions of the northeast and west Atlantic can sustain higher exploitation rates than stocks in poor condition in the colder regimes of the northwest Atlantic (Greenland, Labrador and Grand Bank). This is confirmed by the positive relationship established between the estimated biological management reference points F_med and the mean cod condition factors, as well as by the recent status of these stocks.     

Technological solutions and operational measures to prevent escapes of Atlantic cod (Gadus morhua) from sea cages

Escapes of cod (Gadus morhua) from sea cages represent an economic problem for farmers and a potential environmental problem. We estimate that 0–6% of cod held in sea‐cage farms in Norway were reported to have escaped each year from 2000 to 2005, which is a high proportion compared with salmon. We interviewed employees at 19 coastal sea‐cage cod farms in Norway to investigate both how and why cultured cod escape and to document cage handling and management strategies that were effective in minimizing escapes. Based on the interviews, we describe five working hypotheses that may explain why a greater proportion of cod than salmon escape: (1) cod are more willing to escape than salmon; (2) cod bite the net cage and create wear and tear; (3) net cages have insufficient technical standards for cod culture; (4) cod are placed in sea cages at considerably smaller sizes than salmon; and (5) cod are more popular feed for predators. Preliminary testing of the hypothesis that cod bite netting and create holes was done by placing pre‐damaged net panels with cut twines and control panels inside sea cages. Holes in the pre‐damaged net panels increased in size over a period of 3 months. The type of damage indicated that biting of netting twines was the likely cause.     

Anaesthesia of Atlantic cod (Gadus morhua) — Effect of pre-anaesthetic sedation, and importance of body weight, temperature and stress

The efficacy of the anaesthetic agents benzocaine, metacaine (MS-222), metomidate and 2-phenoxyethanol was studied in Atlantic cod (Gadus morhua) with average body weights of 10 ± 4 g, 99 ± 33 g and 1022 ± 274 g at water temperatures of 8 °C and 16 °C. The agents were tested individually and as combination anaesthesia comprising pre-anaesthetic sedation with a low dosage of metomidate or 2-phenoxyethanol followed by anaesthesia with benzocaine or MS-222. All agents were administered through bath immersion with an exposure time of 5 min. The different treatments resulted in average induction and recovery times ranging from 52 ± 6 s to 182 ± 16 s and 77 ± 26 s to 659 ± 46 s respectively. Induction and recovery times varied in relation to water temperature and were generally shorter at 16 °C for all weight groups and treatments compared to 8 °C. For benzocaine and MS-222 induction and recovery times were found to increase with increasing body weight. For metomidate the recovery time increased with increasing weight whereas there were no weight related differences in induction time. No differences in either induction or recovery times associated to body weight were found for 2-phenoxyethanol. Acute stress prior to anaesthesia with MS-222 resulted in significantly shorter induction time and prolonged recovery time, as well as deeper anaesthetised fish. The dosage of MS-222 had to be reduced in order to avoid mortality in fish subjected to acute stress. Combination anaesthesia allowed a reduction of the dosages used for inducing anaesthesia and produced markedly reduced recovery times compared to agents administered individually.     

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.     

Occurrence of Francisella piscicida in farmed and wild Atlantic cod, Gadus morhua L., in Norway

Francisellosis, caused by the bacterium Francisella piscicida , has become one of the most serious diseases in Atlantic cod production in Norway. The major aim of this study was to determine the distribution of F. piscicida in farmed and wild fish in areas with cod farming along the Norwegian coast, and its occurrence in cod from areas without cod farming. Two real-time PCR assays, targeting the 16S rRNA gene and the FopA gene of F. piscicida , were developed since sensitive and specific diagnostic tools are required for detecting asymptomatic carriers of the bacterium. A total of 422 wild cod from 13 sampling areas and 955 farmed cod from 10 areas along the coast of Norway were examined. Using the real-time polymerase chain reaction (PCR) assays, F. piscicida was detected in wild populations of cod from all counties examined south of Sogn og Fjordane in southern Norway (overall prevalence 13%, n  = 221). Wild cod north of Sogn og Fjordane were negative for the bacterium ( n  = 201). Farmed cod from most parts of Norway were F. piscicida positive. The apparent absence of the bacterium in wild populations of cod in the northern parts of Norway and its widespread occurrence in wild cod from southern parts of Norway is believed to relate to differences in seawater temperatures.     

Resistance of aquaculture net cage materials to biting by Atlantic Cod (Gadus morhua)

Cod bite on aquaculture net cages has resulted in damages like frayed netting and holes, which in part can explain why cultured cod have escaped more frequently than salmon over the last years. We describe damages found on various netting materials subjected to cod bite through field experiments at commercial cod farms. Further, a method to test local cod bite resistance of traditional netting structures is suggested and initial results from a test jig prototype are given. Results from field experiments indicated that cod may have been attracted by types of netting that made it possible to draw filaments out of the twine, while stiff, coated netting structures and thick filaments showed no sign of bite damage during the test period. We concluded that netting materials for cod aquaculture must be resistant to cod bite or be repellent or uninteresting for cod. Based on the present findings, the better choice among the traditional netting materials seemed to be hard-laid netting materials, preferably with a primer that glues the filaments together.     

Is there a genetic basis to growth in Atlantic cod?

There is still much disagreement and debate about whether or not genetically based growth differences occur in Atlantic cod, and there is evidence on both sides. In this review, data on genetically based growth differences in cod will be presented to shed light on this hypothesis. Motivated by the hypothesis that growth patterns may reflect specific genotype adaptations, we review stock‐specific responses on growth. An example of genetically based differences between the population units at two spawning localities off south Iceland is discussed. Here, significant differences in growth performance of the different Syp‐I genotypes were found. Also, the cod sampled at Loftstaðahraun displayed higher mean weight and length compared to the cod from Kantur indicating that these population units may display different life histories. Other studies have shown conflicting results depending on which side of the Atlantic the problem has been investigated. We propose that a common‐garden meta‐analysis with several cod stocks from both sides of the Atlantic is needed to give any reasonable answer to the question of genetically based growth differences. Until such studies have been conducted, it is premature to conclude one way or the other. In this review, we have not tried to quantify how large the environmental part of growth regulation versus the genetic part is, as this information is not available in the published literature on cod. Based on recent research on two flatfish species (turbot and Atlantic halibut), approximately 30% of growth variation is caused by genetical factors, but it remains to be seen if this is similar in cod. A fruitful way to continue this research might be to conduct controlled experiments, where performance (growth, food intake, feed conversion efficiency, feeding behaviour, etc.) and environmental factors (e.g. temperature, oxygen, photoperiod, predation risk, food availability) are studied simultaneously for different genotypes and different stocks.     

Discrete spatial dynamics in a marine broadcast spawner: Re-evaluating scales of connectivity and habitat associations in Atlantic cod (Gadus morhua L.) in coastal Newfoundland

We examined the spatial dynamics of Atlantic cod, Gadus morhua, through its life history on the northeast and southeast coasts of Newfoundland in order to delineate dispersal and critical habitat requirements as potential determinants of appropriate spatial scales for management. Hydroacoustic surveys across the northeast Newfoundland shelf in 1990, 1995, and 2000 showed the distribution of the remaining large aggregations of cod to be concentrated in inshore embayments, which is contrary to historic patterns. Inshore surveys on the south coast identified local spawning aggregations at specific geographic locations, to which adults homed in subsequent years. Dispersal of eggs and larvae to the offshore occurred during spring, but was localized (within bay) in the warm summer months. Abundance of age-0 cod increased with proximity to spawning location inshore, suggesting self-recruitment within local embayments. Age-2–3 juveniles actively selected structurally complex habitat at scales of less than tens of kilometers, with home ranges typically less than 10 ha. Juveniles were associated with specific habitat types, especially at age 0, in both lab and field experiments. Our results suggest that self-recruitment and critical life-history events (e.g., spawning, settlement, and maturity) may occur within local embayments, which currently assume much greater importance to overall cod stock dynamics than in the past. Given evidence for limited dispersal, and the inshore presence of critical habitat for early life-history stages, spatial management tools that consider local dynamics may provide a greater degree of protection for coastal Newfoundland cod than previously thought.     

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.     

Trophic role of Atlantic cod in the ecosystem

As the world's oceans continue to undergo drastic changes, understanding the role of key species therein will become increasingly important. To explore the role of Atlantic cod ( Gadus morhua Gadidae) in the ecosystem, we reviewed biological interactions between cod and its prey, predators and competitors within six ecosystems taken from a broad geographic range: three are cod-capelin ( Mallotus villosus Osmeridae) systems towards cod's northern Atlantic limit (Barents Sea, Iceland and Newfoundland–Labrador), two are more diverse systems towards the southern end of the range (North Sea and Georges Bank–Gulf of Maine), and one is a species-poor system with an unusual physical and biotic environment (Baltic Sea). We attempt a synthesis of the role of cod in these six ecosystems and speculate on how it might change in response to a variety of influences, particularly climate change, in a fashion that may apply to a wide range of species. We find cod prey, predators and competitors functionally similar in all six ecosystems. Conversely, we estimate different magnitudes for the role of cod in an ecosystem, with consequently different effects on cod, their prey and predator populations. Fishing has generally diminished the ecological role of cod. What remains unclear is how additional climate variability will alter cod stocks, and thus its role in the ecosystem.     

Repeatability of fin length measurements using digital image analysis and studies of fin erosion as indicator of social interactions in Atlantic cod (Gadus morhua)

The aims of this study were to examine fin erosion caused by social interactions between Atlantic cod (Gadus morhua) juveniles, and to evaluate the repeatability of recorded fin length using digital image analysis (DIA). Fin length of 2100 juvenile Atlantic cod was analysed using DIA, during a 6 weeks experiment. The lengths of the three dorsal fins and the caudal fin were measured by three different assessors (M, K, B) three times during the experiment. Repeatability was based on a sub set of data from 42 randomly chosen fish. Assessor M measured the length of the fins of the 42 fish three times (M1, M2, M3). M1 and M2 were done on the same day, while M3 was conducted 2 weeks later. Assessor K measured two times with a 2‐year interval, and assessor B measured the fin length only once. The correlations between replicated measurements within assessor were highest for assessor M measured on the same day indicating an effect of memory. Repeatability of fin length ranged from 0.57 to 0.73 for the four fins. Social interaction with respect to fin erosion was studied using data on the four fins from all 2100 fish. ‘Relative fin length’ (fin length/total body length × 100) was calculated to assess fin erosion. Results indicated that all four fins were eroded due to social interactions, but the caudal fin was most severely eroded. In conclusion, DIA can be used to measured fin length but the method could be improved by using chromatic pictures.