Abundance and growth of larval and early juvenile cod (Gadus morhua) in relation to variable environmental conditions west of Iceland

Around Iceland, the west- and north-flowing coastal current, induced by freshwater runoff, provides a transport mechanism for pelagic eggs and larvae derived from the main spawning grounds off the southwest coast to the main nursery grounds off the north coast. In the present study, abundance and growth of larval and juvenile cod were recorded during a series of cruises conducted in June/July of 1998–2001 along the drift route southwest and west of Iceland. The cruises provided information on approximately 2–8-week-old individuals. Hatch dates and abundance varied greatly between years. Hatch dates ranged from Julian Day 92 to 167. Growth rate differed also between the years studied. Relative abundance was generally greatest in temperatures above 7.5 °C and in low-salinity waters, characteristic for the coastal current. The study demonstrates the link between the coastal current and larval/juvenile distribution, thus providing evidence for its importance in promoting successful recruitment of the Icelandic cod stock.     

Influence of water temperature and oxygenation on the aerobic metabolic scope of Atlantic cod (Gadus morhua)

Environmental influences (temperature and oxygenation) on cod metabolism and their impact on the ecology of this species were investigated. Limiting oxygen concentration curves (O₂ level ranging between 15 and 100% air saturation) were established at 2, 5 and 10°C. The standard metabolic rate (SMR), the maximum metabolic rate and the metabolic scope were then modelled as functions of temperature and/or oxygen saturation. The mean SMR at 2, 5 and 10°C were 19.8±4.9, 30.8±6.1 and 54.3±4.1 mg O₂ h⁻¹ kg⁻¹, respectively. Between 2 and 5°C, the active metabolic rate of cod almost doubled from 65 to 120 mg O₂ h⁻¹ kg⁻¹, to reach 177 mg O₂ h⁻¹ kg⁻¹ at 10°C. In terms of metabolic scope (MS), the temperature rise from 2 to 5°C resulted in a two-fold increase from 45 to 89 mg O₂ h⁻¹ kg⁻¹, with MS reaching 123 mg O₂ h⁻¹ kg⁻¹ at 10°C. Our proposed model describing the impact of temperature and oxygen level provides new insight into the energetic interactions which govern the relationship between Atlantic cod and its environment. We re-examined published experimental and field studies from the angle of the regulation of metabolic power. We suggest that, when faced with heterogeneous or unstable hydrological conditions, cod tend to behaviourally maximise their metabolic scope. Through this adaptive response, fish reduce energy budgeting conflicts and presumably increase the probability of routinely operating away from lethal boundaries.     

Cytochromes P-450 in cod (Gadus morhua): Effects of induces on regiospecificity of phenanthrene metabolism and immunochemical properties

The metabolism of some xenobiotics can lead to the formation of reactive intermediates with mutagenic/carcinogenic properties. With the carcinogenic PAH these have been identified as bay-region diol-epoxides.¹ Phenanthrene, a non-carcinogenic, bay-region containing model PAH, is metabolised in vivo by bony fish at the proximate bay-region position, whereas mammals and other marine organisms mainly form the K-region metabolite 9,10-dihydro-9,10-dihydroxy-phenanthrene.² We wanted to investigate this difference more closely by studying the regiospecificity of phenanthrene metabolism in vitro both with microsomes from differently pretreated cod and with isolated cytochrome P-450 isozymes from BNF-induced cod.³ Secondly, by preparing antibodies to the major isozyme isolated (called cod P-450c), we investigated the immunochemical properties of the variously treated cod liver microsomes.     

Modelling the influences of atmospheric forcing conditions on Baltic cod early life stages: distribution and drift

Retention or dispersion of larvae from the spawning ground has been identified as one of the key processes influencing recruitment success in fish stocks. An exercise combining 3-D hydrodynamic model simulations and field data on spatial distributions of juvenile Baltic cod was utilised to investigate the potential drift of larvae from the centre of main spawning effort in the Bornholm Basin, Baltic Sea. In the simulations cod larvae were represented as Lagrangian drifters. Habitats in which larvae and juvenile cod potentially dwell and where juveniles settle were identified to ascertain the importance of predicting transport. The transport of Baltic cod larvae was investigated by detailed drift model simulations for the years 1986 to 1999. The results yielded a clear dependency on wind-induced drift of larval cod, which is mainly controlled by the local atmospheric conditions over the Baltic Sea. Seasonally averaged distributions of drifters were compared with actual distributions of 0-group cod, as determined from bottom and pelagic trawl surveys conducted in autumn of the years 1993 to 2000 in and around the Bornholm Basin. The results suggest that juveniles caught in different areas can be assigned to different times of the spawning season. Because of seasonal differences in the circulation patterns, the southern coastal environment is on average most important for early and late spawners, whereas larvae hatching in mid-summer were on average transported towards the north or to a higher degree remained in the spawning ground.     

Candidate biomarker discovery in plasma of juvenile cod (Gadus morhua) exposed to crude North Sea oil, alkyl phenols and polycyclic aromatic hydrocarbons (PAHs)

In this study we have investigated protein changes in plasma of juvenile Atlantic cod (Gadus morhua) induced by crude North Sea oil and North Sea oil spiked with alkyl phenols and polycyclic aromatic hydrocarbons, a surrogate produced water composition. Using a proteomic approach, we identified 137 differentially expressed proteins at different levels of crude oil exposure. Many of the induced protein changes occurred at low levels of exposure. The results obtained with protein expression profiles after exposure to oil and surrogate produced water indicate effects on fibrinolysis and the complement cascade, the immune system, fertility-linked proteins, bone resorption, fatty acid metabolism as well as increased oxidative stress, impaired cell mobility and increased levels of proteins associated with apoptosis. Although the number of individuals and samples in this study is limited within each treatment group, the protein changes observed in this study represent a first screening for potential biomarker candidates in cod plasma reflecting potential effects of crude oil and produced water exposure on fish.     

Toxicity of sea-surface microlayer: Effects of hexane extract on baltic herring (Clupea harengus) and atlantic cod (Gadus morhua) embryos

This study was designed to determine whether contaminated sea-surface microlayer was toxic to marine fish embryos in its ntaive form and as a hexane extract. Developing embryos of Atlantic cod (Gadus morhua) and Baltic herring (Clupea harengus) were exposed to hexane extracts of sea-surface microlayer collected from five locations in the North Sea and Baltic Sea. Extracts from two of these locations produced significant embryos mortality as well as severe deformities in live hatched larvae. A control sample of bulk water collected from 20 cm under the surface and extracted in the same way produced no significant mortality or deformities. Significaant changes in timing of hatching were also observed in those samples which produced embryo toxic effects. A comparison of these data with those obtained from code embryos exposed to unextracted microlayer showed a similar biological effect with both unextracted samples and hexane extracts.Chemical analyses revealed the greatest biological effect in samples with petroleum hydrocarbon concentrations between 180 and > 200 μg liter⁻¹. The bulk water control had 1 μg liter⁻¹ while the three samples that showed no biological activity had 3 to 8 μg liter⁻¹ petroleum hydrocarbons. Phthalic acid esters were detected in four samples and chlorinated hydrocarbons in one, but could not be positively correlated with any of the toxic responses. No other chemical contaminants were detected in the five samples.The data presented here show that some sites contain sea-surface microlayer which can be toxic to marine fish embryos: that Baltic herring and Atlantic cod embryos respond similarly to the toxic effects of contaminated microlayer: and that unextracted microlayer and hexane extract of microlayer produce essentially the same toxic effect(s) if only organic contaminants are considered.     

Bioavailability of PAH in effluent water from an aluminium works evaluated by transplant caging and biliary fluorescence measurements of Atlantic cod (Gadus morhua L.)

Fixed wavelength fluorescence (FF) of bile was used as a biomarker for PAH exposure in fish caged adjacent to an aluminium works. After 30 days of caging, a 20–50 fold increase of pyrene fluorescence was found in groups caged adjacent to the works as compared to two control groups. The PAH uptake occurred mainly from suspended particles rather than from PAH contaminated sediments at the caging sites. Our results demonstrates the combined use of fish caging and bile-fluorescence measurements as a powerful field model for the monitoring of coastal waters subjected to PAH pollution.     

Distributions of copepod nauplii and turbulence on the southern flank of Georges Bank: implications for feeding by larval cod (Gadus morhua)

The vertical distributions of copecod nauplii and water properties were sampled at well-mixed and stratified sites on Georges Bank using a pumping system, CTD and in vivo fluorometer during a four day period in late May 1992. At each stratified station at least one sample was taken within the thermocline and the fluorescence maximum, which usually co-occurred. Well-mixed sites had low average concentrations of nauplii, ca 41⁻¹, and showed little variation of abundance with depth. Stratified sites had from 4 to 16 times the integrated (0–50 m) abundance of nauplii compared to well-mixed sites and showed strong vertical patterns of distribution. Maximum concentrations of nauplii, up to 1601⁻¹, were associated with the thermocline at 7 of the 9 stratified stations. At the two remaining stratified sites the naupliar maximum was in the upper mixed layer, sampled at 5 m depth. The encounter rate between early feeding cod (Gadus morhua) larvae and their naupliar prey was calculated with and without turbulence. Turbulence was estimated from two sources: wind stress in the upper layer (calculated from wind observations during our cruise) and tidal shear in the lower layer (estimated initially from a tidal mixing equation). We ultimately replaced the lower layer estimates with turbulence data from a series of measurements made in 1995. The latter are more robust and had the advantage of providing dissipation rates for the pycnocline as well as the lower layer. Theory predicts an increase in encounters between a predator and its prey with the addition of turbulence parameters into standard models of encounter. We combined turbulence profiles with the vertical distribution of nauplii to examine the potential contribution of turbulent kinetic energy to predator-prey encounter rates at various depths in stratified and mixed water columns. Our calculations suggest the following increases due to turbulence at stratified sites on Georges Bank during our cruise: from 34 to 219% in the upper mixed layer, depending on wind speed and depth; approximately 8% in the pycnocline; and approximately 110% below the pycnocline. Mixed sites experience increases of at least 110% (tide only), but greater increases (118–192% in this study) occur when the wind blows because of the combined (spatially overlapped) effects of wind and tidal mixing at these sites. The absolute values for encounter rates and their modification by turbulence are sensitive to a number of assumptions in the models. We used a series of stated assumptions to generate estimates that range from 0.6 to 26.5 prey h⁻¹, depending on geographical location, physical forcing and depth.     

Influence of diet on growth, condition and reproductive capacity in Newfoundland and Labrador cod (Gadus morhua): Insights from stable carbon isotopes (δC)

Cod populations in Newfoundland and Labrador waters have shown differing growth, condition and recruitment since near-universal declines in these properties during the cold period of the late 1980s and early 1990s. To assess the influence of variable prey communities on these parameters, we compared cod energetics and diet in populations off Labrador and the northeast and south coasts of Newfoundland. Many properties were highest in the southern group(s) and lowest in the northern group(s), including growth, somatic condition, liver index and age-at-maturity. Most differences could be explained by variations in diet, as measured by stomach contents and stable carbon isotopes (δ¹³C). The diet of Labrador cod consisted almost entirely of northern shrimp (Pandalus borealis), and these cod displayed the most benthic δ¹³C signatures. Northeast cod had a more varied diet that included capelin and other fish, but still had mostly benthic δ¹³C signatures, suggesting the importance of benthic prey like shrimp in this population. South coast cod exhibited the most varied diet, including capelin (Mallotus villosus), zooplankton, crabs and other fish, and had the most pelagic δ¹³C signatures. Among and within populations, the benefits of a more pelagic diet in medium-sized (30–69 cm) cod included higher somatic condition, higher liver index (lipid stores) and greater spawning potential (decreased incidence of atresia). It is hypothesized that major rebuilding of Newfoundland and Labrador cod stocks will require a return to a system that supports mostly pelagic feeding (i.e. capelin) in cod.     

Trophic relations of capelin Mallotus villosus and polar cod Boreogadus saida in the Barents Sea as a factor of impact on the ecosystem

The purpose of the study is to assess the role of trophic relations of the dominant pelagic fishes capelin and polar cod in the Barents Sea with regard to distribution and accessibility as prey for the Atlantic cod in warm years (2004–2005). Unlike in the previous period, during these warm years a dramatic increase of the polar cod population resulted in a northwards expansion of the feeding grounds where overlapping of polar cod and capelin concentrations was observed. This caused an increased competition for copepods, which are the main food item for young fish. In the areas dominated by polar cod the shortage of copepods forced immature capelin to switch to the chaetognath Sagitta, which affected their fatness negatively.During the warm years the feeding grounds of Atlantic cod also expanded, to a large degree caused by the shortage of their main food, the capelin. In 2004–2005 the cod formed feeding concentrations in the north and northeast Barents Sea where they fed on the capelin. In this area the consumption of polar cod by cod increased, and in some local areas the polar cod practically replaced the capelin in the diet of cod. In general polar cod in the diet of Atlantic cod were more important in the northern than in the southern part of the Barents Sea. The fatness of cod was extremely low during the whole spring–summer period (until August), and after the feeding period the fatness index of the Atlantic cod became lower than the average long-term autumn value.     

Sublethal effects of the water-soluble fraction of ekofisk crude oil on the early larval stages of cod (Gadus morhua L.)

Low levels of dissolved oil hydrocarbons are demonstrated to produce sublethal effects on the early cod larval stages. These effects are discussed in relation to larval survival through the first critical stages.Continuous exposure of cod eggs and larvae to 0·05 and 0·25 ppm of the water-soluble fraction (WSF) of Ekofisk crude oil for 14 days caused significant reduction in growth and change in neutral buoyancy. The larvae exposed to 0·25 ppm showed malformation in the foremost part of the head and jaw, which reduced their ability to capture prey organisms at first feeding.     

Comparison of the response of Atlantic cod (Gadus morhua) in the high-latitude regions of the North Atlantic during the warm periods of the 1920s–1960s and the 1990s–2000s

Concern about future anthropogenic warming has lead to demands for information on what might happen to fish and fisheries under various climate-change scenarios. One suggestion has been to use past events as a proxy for what will happen in the future. In this paper a comparison between the responses of Atlantic cod (Gadus morhua) to two major warm periods in the North Atlantic during the 20th century is carried out to determine how reliable the past might be as a predictor of the future. The first warm period began during the 1920s, remained relatively warm through the 1960s, and was limited primarily to the northern regions (>60°N). The second warm period, which again covered the northern regions but also extended farther south (30°N), began in the 1990s and has continued into the present century. During the earlier warm period, the most northern of the cod stocks (West Greenland, Icelandic, and Northeast Arctic cod in the Barents Sea) increased in abundance, individual growth was high, recruitment was strong, and their distribution spread northward. Available plankton data suggest that these cod responses were driven by bottom-up processes. Fishing pressure increased during this period of high cod abundance and the northern cod stocks began to decline, as early as the 1950s in the Barents Sea but during the 1960s elsewhere. Individual growth declined as temperatures cooled and the cod distributions retracted southward. During the warming in the 1990s, the spawning stock biomass of cod in the Barents Sea again increased, recruitment rose, and the stock spread northward, but the individual growth did not improve significantly. Cod off West Greenland also have shown signs of improving recruitment and increasing biomass, albeit they are still very low in comparison to the earlier warming period. The abundance of Icelandic cod, on the other hand, has remained low through the recent warm period and spawning stock biomass and total biomass are at levels near the lowest on record. The different responses of cod to the two warm events, in particular the reduced cod production during the recent warm period, are attributed to the effects of intense fishing pressure and possibly related ecosystem changes. The implications of the results of the comparisons on the development of cod scenarios under future climate change are addressed.     

Metabolism of the S-triazine herbizide atrazine in early life stages of the zebrafish, Danio rerio

Increasing anthropogenic pollution of the environment can have adverse consequences for organisms which are more subtle than direct toxicity. Detectable levels of persistent pollutants remain even if the substance is no longer used. An example is atrazine, a herbicide in common use throughout the world but one which has been banned in Germany since 1992. As a lipophilic substance, atrazine is bioconcentrated which may lead to chronic intoxication or physiological stress. In order to withstand chemical stressors, many organisms possess detoxication enzymes, for example certain P-450 monoxygenases and glutathione S-transferases. But detoxication demands cellular energy, and in developing organisms, such as fish embryos which have particularly high energy needs for sustaining growth and organogenesis, the additional energy needs of detoxication may present additional stress. In this study, uptake of atrazine, cytochrome P-450 binding spectra, effects on microsomal and soluble glutathione S-transferase activities, and the initial detoxication steps of atrazine via microsomal and soluble glutathione S-transferases were studied using early life stages of zebrafish.     

Causes of different impact of Major Baltic Inflows on cod reproduction in the Gotland Basin of the Baltic Sea

The impact of the Major Baltic Inflows (MBIs) of 1976, 1993, and 2003 on cod reproduction in the Gotland Basin of the Baltic Sea is considered based on calculations of the eastern Baltic cod egg abundance and offspring survival indices. The reproductive success (1976–1977) was defined by the elevation of the 11 psu isohaline and deepening of the 2 mL/L isooxygene surface, which provided the formation of a water layer suitable for cod spawning at a depth of 90–130 m. Prerequisites of the appearance of the reproductive layer in 1976–1977 were a previously high salinity level and penetrating two successive MBIs in one year. The cause of the weak MBI impact in 1993 was the previous salinity minimum; in 2003 it was rapid oxygen depletion due to an increase in temperature. As a result of their weaker influence, small and fragmented cod spawning biotopes appeared in the Gotland Basin.