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The objective was to investigate changes, throughout the spawning season, in body size attributes and quantitative semen characteristics of wild-caught and cultivated Atlantic cod, Gadus morhua L. Sperm velocity increased significantly throughout the spawning season of cod from both origins. Curvilinear velocity (VCL; 30 sec post-activation) increased from 78.9+/-6.5 to 128.2+/-6.5 microm/sec (mean+/-SEM) between the beginning and end of the spawning season, respectively, for wild-caught cod, whereas for cultivated fish, it increased from 26.6+/-2.4 to 48.9+/-3.1 microm/sec between January and March. Spermatocrit did not undergo a significant seasonal change in wild-caught cod but did thicken for cultivated cod (24.6+/-4.2% in January to 40.5+/-4.4% in April; P<0.01). Sperm head area, perimeter, length, and width declined significantly at the end of the spawning season of cod from both origins (all P values<0.01). Seminal plasma osmolality and Na(+) ion concentration followed a dome-shaped function through the spawning season for both wild-caught and cultivated cod (P<0.05). For cultivated cod, seminal plasma pH was significantly lower at the start of the spawning season (P<0.001), whereas Ca(2+) increased then decreased (P<0.05). Body size attributes, spermatocrit, and seminal plasma constituents had significant relationships with sperm activity variables. These relationships varied as a function of time post-activation, month, and fish origin. Our findings may be used to (i) assess spermiation stage without killing males; (ii) optimize semen collection for hatchery production; (iii) characterize the potential impact of farming on sperm quality; and (iv) improve success of sperm cryopreservation and short-term storage., (Copyright 2010 Elsevier Inc. All rights reserved.)

Fecundity data for four species (American plaice Hippoglossoides platessoides, yellowtail flounder Limanda ferruginea, witch flounder Glyptocephalus cynoglossus and Atlantic cod Gadus morhua) of north-west Atlantic demersal fishes, corresponding to nine populations, were examined in relation to fish size and condition in an attempt to explain the observed variability in potential fecundity. Both relative body (K(r)) and liver condition (H(r)) were poor single-factor predictors of fecundity, and in almost all cases fish body mass (M) was the best single-factor predictor. Annual variability in mean K(r) and H(r) existed for most populations. The inclusion of K(r) and H(r) in total length (L(T))-based predictive models improved model fit only slightly and not significantly in all cases. Multiple regression analyses to determine the best model for explaining the variability in fecundity often excluded K(r), H(r) and L(T) in favour of M. The amount of variability in fecundity that could be explained by the factors analysed here was species specific, with the highest proportion explained for H. platessoides and the lowest for L. ferruginea. The highly variable, and sometimes unpredictable, nature of north-west Atlantic groundfish fecundity suggests the need to continue collecting such reproductive data on an ongoing basis.

Low water oxygen content (hypoxia) is a common feature of many freshwater and marine environments. However, we have a poor understanding of the degree to which diminished cardiac function contributes to the reduction in fish swimming performance concomitant with acute exposure to hypoxia, or how fish cardiorespiratory physiology is altered by, or adapts to, chronic hypoxia. Thus, we acclimated adult Atlantic cod (Gadus morhua) to either approximately 8-9 kPa O(2) (40-45% air saturation) or approximately 21 kPa O(2) (100% air saturation; normoxia) for 6-12 weeks at 10 degrees C, and subsequently measured metabolic variables [routine oxygen consumption (M(O(2)), maximum (M(O(2)), metabolic scope] and cardiac function (cardiac output, Q; heart rate, f(H); and stroke volume, V(S)) in these fish during critical swimming speed (U(crit)) tests performed at both levels of water oxygenation. Although surgery (flow probe implantation) reduced the U(crit) of normoxia-acclimated cod by 14% (from 1.74 to 1.50 BL s(-1)) under normoxic conditions, exposure to acute hypoxia lowered the U(crit) of both groups (surgery and non-surgery) by approximately 30% (to 1.23 and 1.02 BL s(-1), respectively). This reduction in swimming performance was associated with large decreases in maximum M(O(2)) and metabolic scope (> or = 50%), and maximum f(H) and Q (by 16 and 22%), but not V(S). Long-term acclimation to hypoxia resulted in a significant elevation in normoxic metabolic rate as compared with normoxia-acclimated fish (by 27%), but did not influence normoxic or hypoxic values for U(crit), maximum M(O(2)) or metabolic scope. This was surprising given that resting and maximum values for Q were significantly lower in hypoxia-acclimated cod at both levels of oxygenation, because of lower values for V(S). However, hypoxia-acclimated cod were able to consume more oxygen for a given cardiac output. These results provide important insights into how fish cardiorespiratory physiology is impacted by short-term and prolonged exposure to hypoxia, and further highlight the tremendous capacity of the fish cardiorespiratory system to deal with environmental challenges.

Recent in vivo experiments on Atlantic cod (Gadus morhua) acclimated to chronic hypoxia (6-12 weeks at 10 degrees C; Pw(O(2)) approximately 8-9 kPa) revealed a considerable decrease in the pumping capacity of the heart. To examine whether this diminished cardiac performance was due to the direct effects of chronic moderate hypoxia on the myocardium (as opposed to alterations in neural and/or hormonal control), we measured the resting and maximum in situ function of hearts from normoxia- and hypoxia-acclimated cod: (1) when initially perfused with oxygenated saline; (2) at the end of a 15 min exposure to severe hypoxia (P(O(2)) approximately 0.6 kPa); and (3) 30 min after the hearts had been reperfused with oxygenated saline. Acclimation to hypoxia did not influence resting (basal) in situ cardiac performance during oxygenated or hypoxic conditions. However, it caused a decrease in maximum cardiac output (Q(max)) under oxygenated conditions (from 49.5 to 40.3 ml min(-1) kg(-1); by 19%), that was due to diminished values for maximum stroke volume (V(S)) and scope for V(S). Severe hypoxia reduced in both groups to approximately 20 ml min(-1) kg(-1), yet, the hearts of hypoxia-acclimated fish were better able to sustain this level of Q under hypoxia, and the recovery of Q(max) (as compared with initial values under oxygenated conditions) was significantly improved (94% vs 83%). These data show that acclimation to hypoxia has a direct effect on cod myocardial function and/or physiology, and suggest that the cod heart shows some adaptations to prolonged hypoxia.

Atlantic cod trypsin I is a highly active cold-adapted protease. This study aimed at further characterization of this enzyme with respect to kinetic parameters, sites of autolysis and stability. For that purpose, trypsin I was purified by anion exchange chromatography. Its purity and identity was verified by SDS-PAGE analysis and mass spectrometry. Concomitantly, another cod trypsin isozyme, trypsin X, previously only described from its cDNA sequence was detected in a separate peak from the ion exchange chromatogram. There was a stepwise increase in the catalytic efficiency (k(cat)/K(m)) of cod trypsin I obtained with substrates containing one to three amino acid residues. As expected, the activity of trypsin I was maintained for longer periods of time at 15 degrees C than at higher temperatures. The residues of the trypsin I molecule most sensitive to autolysis were identified using Edman degradation. Eleven autolytic cleavage sites were detected within the trypsin I molecule. Unfolding experiments demonstrated that autolysis is a contributing factor in the stability of trypsin I. In addition, the data shows that cod trypsin I is less stable towards thermal unfolding than its mesophilic bovine analogue.

Phagocytosis by fish cells has mostly been studied using adherent leucocytes, excluding suspended cells such as the majority of B-cells and neutrophils, but a recent study describes professional phagocytosis of latex beads and bacteria by B-cells from rainbow trout. In the present study, phagocytosis by B-cells and neutrophils from salmon and cod was studied. Leucocytes were isolated from peripheral blood (PBL) and head kidney (HKL). By flow cytometry analyses, proportions of MAb labelled cell populations with internalized fluorescent beads, as well as the number of beads within each cell, could be determined. Phagocytic capacity and ability were demonstrated in B-cells and neutrophils from salmon and cod. In salmon, B-cells had higher phagocytic ability than neutrophils in HKL, but not in PBL. For cod the phagocytic ability of B-cells were lower than for neutrophils in both HKL and PBL, but the phagocytic capacity of cod B-cells were higher than for neutrophils in both HKL and PBL. For salmon B-cells the phagocytic capacity was lower than or similar to neutrophils in HKL and PBL. The total phagocytic ability of leucocytes was different in the species studied. The highest phagocytic ability was observed in cod, showing similar values for PBL and HKL. Salmon PBL displayed about twice the phagocytic ability of cod PBL. There seemed to be some major differences between the two fish species concerning phagocytosis. In salmon, a rather large proportion of phagocytic leucocytes were phagocytic B-cells, indicating that B-cells may have an important function in particle clearance in this species. In cod, phagocytic leucocytes in HKL and PBL were mostly neutrophils, and only a small proportion of B-cells were phagocytic, supporting the more prominent role of innate immune functions in cod neutrophils., (Copyright 2009 Elsevier Ltd. All rights reserved.)

One of the main gastrointestinal hormones, cholecystokinin (CCK), was studied in order to advance understanding of the control of the digestive process in Atlantic cod Gadus morhua larvae after onset of first feeding. Larvae were fed either natural zooplankton or enriched rotifers in similar rearing systems and sampled from hatching to 22 days post-hatch (dph). CCK was visualized by immunohistochemistry and the first CCK-immunoreactive (IR) cells were detected at 8 dph corresponding to 6 days after first feeding. The CCK-IR cells were mostly found in the anterior midgut, and the number of CCK-IR cells was lower in the posterior midgut. They were also present in the hindgut of some of the larvae, but not in the foregut. No clear differences were found in the ontogenetic appearance and the distribution pattern of CCK-IR cells between the two dietary treatments. This indicates that the onset of CCK production in the gut as well as the spatial distribution of the CCK-IR cells is not differentially affected by these diets. To what extent the hormone production itself is influenced by dietary factors needs to be studied by more sensitive methods.

Hemoglobin (Hb) polymorphism in cod is associated with temperature-related differences in biogeographical distribution, and several authors have suggested that functional characteristics of the various hemoglobin isoforms (HbIs) directly influence phenotypic traits such as growth rate. However, no study has directly examined whether Hb genotype translates into physiological differences at the whole animal level. Thus, we generated a family of juvenile Atlantic cod consisting of all three main Hb genotypes (HbI-1/1, HbI-2/2, and HbI-1/2) by crossing a single pair of heterozygous parents, and we compared their metabolic and cortisol responses to an acute thermal challenge (10 degrees C to their critical thermal maximum [CTM] or 22 degrees C, respectively) and tolerance of graded hypoxia. There were no differences in routine metabolism (at 10 degrees C), maximum metabolic rate, metabolic scope, CTM (overall mean 22.9 degrees +/- 0.2 degrees), or resting and poststress plasma cortisol levels among Hb genotypes. Further, although the HbI-1/1 fish grew more (by 15%-30% during the first 9 mo) when reared at 10 degrees +/- 1 degrees C and had a slightly enhanced hypoxia tolerance at 10 degrees C (e.g., the critical O(2) levels for HbI-1/1, HbI-2/2, and HbI-1/2 cod were 35.56% +/- 1.24% , 40.56% +/- 1.99%, and 40.20% +/- 1.19% air saturation, respectively), these results are contradictory to expectations based on HbI functional properties. Thus, our findings (1) do not support previous assumptions that growth rate differences among cod Hb genotypes result from a more efficient use of the oxygen supply-that is, reduced standard metabolic rates and/or increased metabolic capacity-and (2) suggest that in juvenile cod, there is no selective advantage to having a particular Hb genotype with regards to the capacity to withstand ecologically relevant environmental challenges.

An experimental study was performed to disentangle parental and environmental effects on the growth of Atlantic cod Gadus morhua larvae and juveniles. Eggs were collected during the spawning season from spawning pairs (families) kept separately in specially designed spawning compartments. Newly hatched larvae were released simultaneously into two mesocosms of 2,500 and 4,400 m(3). Larval growth was monitored by sampling over a 10 week period, after which juveniles were transferred to on-growing tanks, where they were tagged and kept for up to 2 years. Maternal origin was determined by individual microsatellite genotyping of the larvae (n = 3949, 24 families) and juveniles (n = 600). The results showed significant positive correlations between egg size and larval size during the whole mesocosm period. Correlations, however, weakened with time and were no longer significant at the first tank-rearing sampling at an age of 9 months. Significant family-specific differences in growth were observed. The coefficient of variation (c.v.) was calculated in order to examine variation in standard length of larvae during the mesocosm period. Inter-family c.v. was on average 69% of intra-family c.v. Differences in zooplankton densities between the two mesocosms were reflected in larval growth, condition factor and c.v. Low food abundance appeared to reduce c.v. and favour growth of larvae that showed relatively slow growth at high food abundance. It is suggested that genetically determined variation in growth potential is maintained by environmental variability.

The presence and vasoactive effects of native calcitonin gene-related peptide (CGRP), substance P (SP), and neurokinin A (NKA) were studied on isolated small branches of the coeliac artery from Atlantic cod, Gadus morhua, using immunohistochemistry and myograph recordings, respectively. Immunohistochemistry revealed nerve fibers containing CGRP- and SP/NKA-like material running along the wall of the arteries. CGRP induced vasorelaxation of precontracted arteries with a pD(2) value of 8.54 +/- 0.17. Relaxation to CGRP (10(-8) M) was unaffected by L-NAME (3 x 10(-4) M) and indomethacin (10(-6) M) suggesting no involvement of nitric oxide or prostaglandins in the CGRP-induced relaxation. SP and NKA (from 10(-10) to 3 x 10(-7) M) contracted the unstimulated arteries at concentrations from 10(-8) M and above in 42% and 33%, respectively, of the vessels. It is concluded that the innervation of the cod celiac artery includes nerves expressing CGRP-like and tachykinin-like material, and that a vasodilatory response to CGRP is highly conserved amongst vertebrates while the response to tachykinins is more variable.

Bathymetric and regional variation in condition of Icelandic cod Gadus morhua in autumn is compared to that previously observed in spring. Once again, contradicting patterns in the hepato-somatic index and a morphometric index of fish condition were observed. The relevance of this persistent spatial variation in condition in relation to emerging evidence of behavioural types is discussed.

Thermal acclimation is frequently cited as a means by which ectothermic animals improve their Darwinian fitness, i.e. the beneficial acclimation hypothesis. As the critical swimming speed (U (crit)) test is often used as a proxy measure of fitness, we acclimated Atlantic cod (Gadus morhua) to 4 and 10 degrees C and then assessed their U (crit) swimming performance at their respective acclimation temperatures and during acute temperature reversal. Because phenotypic differences exist between different populations of cod, we undertook these experiments in two different populations, North Sea cod and North East Arctic cod. Acclimation to 4 or 10 degrees C had a minimal effect on swimming performance or U (crit), however test temperature did, with all groups having a 10-17% higher U (crit) at 10 degrees C. The swimming efficiency was significantly lower in all groups at 4 degrees C arguably due to the compression of the muscle fibre recruitment order. This also led to a reduction in the duration of "kick and glide" swimming at 4 degrees C. No significant differences were seen between the two populations in any of the measured parameters, due possibly to the extended acclimation period. Our data indicate that acclimation imparts little benefit on U (crit) swimming test in Atlantic cod. Further efforts need to identify the functional consequences of the long-term thermal acclimation process.

Individual behaviour of Atlantic cod Gadus morhua in the presence of hypoxic water was measured in situ in the vertically stratified Bornholm Basin of the Baltic Sea. Considering all recaptured individuals, the use of hypoxic habitat was comparable to data derived by traditional survey data, but some G. morhua had migrated towards the centre of the c.100 m deep basin and spent about a third of their time at oxygen saturation <50%, possibly to forage on zoobenthos. Maximal residence time per visit in such hypoxic water was limited to a few hours, allowing for the digestion of consumed prey items in waters with sufficient dissolved oxygen.

Key components of swimming metabolism: standard metabolism (R(s)), active metabolism (R(a)) and absolute aerobic scope for activity (R(a)-R(s)) were determined for small age 0 year Atlantic cod Gadus morhua. Gadus morhua juveniles grew from 0.50 to 2.89 g wet body mass (M(WB)) over the experimental period of 100 days, and growth rates (G) ranged from 1.4 to 2.9% day(-1), which decreased with increasing size. Metabolic rates were recorded by measuring changes in oxygen consumption over time at different activity levels using modified Brett-type respirometers designed to accommodate the small size and short swimming endurance of small fishes. Power performance relationships were established between oxygen consumption and swimming speed measurements were repeated for individual fish as each fish grew. Mass-specific standard metabolic rates (RsMWB-1) were calculated from the power performance relationships by extrapolating to zero swimming speed and decreased from 7.00 to 5.77 micromol O(2) g(-1) h(-1), mass-specific active metabolic rates (RaMWB-1) were calculated from extrapolation to maximum swimming speed (U(max)) and decreased from 26.18 to 14.35 micromol O(2) g(-1) h(-1) and mass-specific absolute scope for activity was calculated as the difference between active and standard metabolism (RaMWB-1-RsMWB-1) and decreased from 26.18 to 14.35 micromol O(2) g(-1) h(-1) as M(WB) increased. Small fish with low R(s) had bigger aerobic scopes but, as expected, R(s) was higher in smaller fish than larger fish. The measurements and results from this study are unique as R(s), R(a) and absolute aerobic scopes have not been previously determined for small age 0 year G. morhua.

Viral nervous necrosis (VNN) caused by beta-nodavirus affects many species of farmed marine fish, in particular juveniles. Apparently healthy, normally feeding, adult farmed Atlantic cod, Gadus morhua, were sampled in a farm 14 months after an outbreak of VNN with clinical signs. Following necropsy, brain and eye tissues were examined by histology, immunohistochemistry and polymerase chain reaction (PCR). Nodavirus-provoked cell death and inflammation was detected in eye and brain, particularly in the retina and cerebellum and differed from that previously described in Atlantic cod during clinical stages of VNN. Virus was detected both by PCR and immunohistochemistry. This is, to the best of our knowledge, the first description of pathological changes associated with chronic subclinical nodavirus infection in Atlantic cod. Our observations suggest that severe infection and pathological changes may go undetected if investigations are restricted to clinical examination and macroscopic evaluation at necropsy.

Anthropogenic CO2 emissions lead to chronically elevated seawater CO2 partial pressures (hypercapnia). The induced ocean acidification will very likely be a relevant factor shaping future marine environments. CO2 exposure concomitantly challenges the animal's capacity of acid-base and ionic regulation as well as the ability to maintain energy metabolism and calcification. Under conditions of acute hypercapnia, numerous studies have revealed a broad range of tolerance levels displayed by various marine taxa. Thus, it is well known that, in contrast to many marine invertebrates, most teleost fish are able to fully compensate acid-base disturbances in short-term experiments (hours to several days). In order to determine whether marine fish are able to preserve aerobic scope following long-term incubation to elevated CO2, we exposed two groups of Atlantic Cod for 4 and 12 months to 0.3 and 0.6 kPa P(CO2), respectively. Measurements of standard and active metabolic rates, critical swimming speeds and aerobic scope of long-term incubated cod showed no deviations from control values, indicating that locomotory performance is not compromised by the different levels of chronic hypercapnia. While the maintenance of high activity levels is supported by a 2-fold increased Na+/K+-ATPase protein expression and 2-fold elevated Na+/K+-ATPase activity in the 12 month incubated fish (0.6 kPa P(CO2)), no such elevation in Na+/K+-ATPase activity could be observed in the group treated with 0.3 kPa P(CO2). Owing to the relevance of Na+/K+-ATPase as a general indicator for ion regulatory capacity, these results point at an adjustment of enzymatic activity to cope with the CO2 induced acid-base load at 0.6 kPa P(CO2) while under milder hypercapnic conditions the 'standard' Na+/K+-ATPase capacity might still be sufficient to maintain acid-base status.

Precocious male puberty is a significant problem in Atlantic cod aquaculture. While photoperiod manipulation can inhibit testis growth, a detailed analysis of effects on spermatogenesis is missing. Starting July 1, 2004, prepubertal fish were exposed to different photoperiod regimens in indoor tanks for 17 mo. Testis histology, germ cell dynamics (proliferation and apoptosis), and plasma androgen levels were analyzed. In the natural light (NL) group, testis growth started in September 2004 and was completed in February 2005, when a 2-mo spawning period started. In the constant light (LL) group, none or very few spermatogenic cysts were recruited into spermatogenesis, and apoptotic germ cell loss was high. A change of photoperiod from NL to LL at winter solstice (December 21, 2004) resulted in premature (2 mo) completion of the reproductive cycle, while changing from LL to NL at winter solstice triggered faster than normal testis development. Plasma testosterone levels increased in the NL group from spermatogonial proliferation toward meiosis, while those of 11-ketotestosterone increased toward spermiogenesis and spermiation. Plasma androgen levels did not rise under LL conditions. Comparing fish with developing testes from all groups indicated that low androgen levels were associated with a high incidence of spermatogonial apoptosis; we also found that androgen receptor mRNA expression was most prominent in Sertoli cells in contact with growing spermatogonial clones. Our data show that an inhibitory photoperiod (LL) reduced or blocked differentiation of spermatogonia, increased apoptosis (particularly among proliferating spermatogonia), and was associated with reduced androgen levels, a situation possibly reflecting insufficient gonadotropic stimulation.

A major challenge in evolutionary biology is to identify the genes underlying adaptation. The oxygen-transporting haemoglobins directly link external conditions with metabolic needs and therefore represent a unique system for studying environmental effects on molecular evolution. We have discovered two haemoglobin polymorphisms in Atlantic cod populations inhabiting varying temperature and oxygen regimes in the North Atlantic. Three-dimensional modelling of the tetrameric haemoglobin structure demonstrated that the two amino acid replacements Met55beta1Val and Lys62beta1Ala are located at crucial positions of the alpha1beta1 subunit interface and haem pocket, respectively. The replacements are proposed to affect the oxygen-binding properties by modifying the haemoglobin quaternary structure and electrostatic feature. Intriguingly, the same molecular mechanism for facilitating oxygen binding is found in avian species adapted to high altitudes, illustrating convergent evolution in water- and air-breathing vertebrates to reduction in environmental oxygen availability. Cod populations inhabiting the cold Arctic waters and the low-oxygen Baltic Sea seem well adapted to these conditions by possessing the high oxygen affinity Val55-Ala62 haplotype, while the temperature-insensitive Met55-Lys62 haplotype predominates in the southern populations. The distinct distributions of the functionally different haemoglobin variants indicate that the present biogeography of this ecologically and economically important species might be seriously affected by global warming.

Internally implanted data storage tags (DST) recording hydrostatic pressure (depth) and temperature in Atlantic cod Gadus morhua were used to track the behaviour of nine individuals during fishing operations. Gadus morhua showed identifiable and characteristic behavioural responses to different capture processes. In a unique observation, one fish survived being discarded and upon return to the seabed exhibited a cessation of activity comparable to observations on stressed fish under laboratory conditions.

The extent to which accumulations of polycyclic aromatic hydrocarbons (PAH) by semipermeable membrane devices (SPMD) may be correlated to the concentration of PAH metabolites in fish was examined. Atlantic cod (Gadus morhua) and SPMD were exposed over a 4-wk period to a mixture that was designed to simulate the composition of PAH in produced water discharges in the North Sea. Fish bile and whole SPMD were analyzed after exposure in a high, low, and a control treatment tank. PAH metabolites were detected and quantified by means of high-performance liquid chromatography (HPLC) or gas chromatography-time-of-flight mass spectroscopy (GC-ToF-MS) and PAH in SPMD by gas chromatography-mass spectroscopy (GC-MS). The level and duration of exposure were generally reflected in both matrices. Exposure water concentrations, calculated from SPMD accumulations, showed some considerable differences from nominal concentrations for volatile compounds. Concentration factors (CF) for fish based on SPMD-derived water concentrations ranged from 2.9 to 354.3 L/g. CF increased with hydrophobicity with more variation at higher log K(ow). Regression analysis of accumulations of individual compounds in fish and SPMDs showed reasonable but compound-specific correlation, r2 = .54-.85 (C1-phenanthrenes/anthracenes and pyrene, respectively). Comparison of the two exposures suggests that fish CF may not be entirely independent of water concentrations. Overall the results show that accumulation of up to four-ring PAH in SPMD may be related to internal exposure of fish to these substances when exposed through water, as evidenced by metabolite concentrations. The usefulness of using SPMD to measure exposure concentrations in a large-scale flow-through study is also examined.

During the course of the last century, populations of Atlantic cod Gadus morhua L. have undergone dramatic declines in abundance across their biogeographic range, leading to debate about the relative roles of climatic warming and overfishing in driving these changes. In this chapter, we describe the geographic distributions of this important predator of North Atlantic ecosystems and document extensive evidence for limitations of spatial movement and local adaptation from population genetic markers and electronic tagging. Taken together, this evidence demonstrates that knowledge of spatial population ecology is critical for evaluating the effects of climate change and commercial harvesting. To explore the possible effects of climate change on cod, we first describe thermal influences on individual physiology, growth, activity and maturation. We then evaluate evidence that temperature has influenced population-level processes including direct effects on recruitment through enhanced growth and activity, and indirect effects through changes to larval food resources. Although thermal regimes clearly define the biogeographic range of the species, and strongly influence many aspects of cod biology, the evidence that population declines across the North Atlantic are strongly linked to fishing activity is now overwhelming. Although there is considerable concern about low spawning stock biomasses, high levels of fishing activity continues in many areas. Even with reduced fishing effort, the potential for recovery from low abundance may be compromised by unfavourable climate and Allee effects. Current stock assessment and management approaches are reviewed, alongside newly advocated methods for monitoring stock status and recovery. However, it remains uncertain whether the rebuilding of cod to historic population sizes and demographic structures will be possible in a warmer North Atlantic.

It is generally assumed that there should be some relationship between tissue residues of contaminants and related biomarker responses in the same individual, with a possible additional modulation through physiological processes. The main basis for such assumptions originated from experimental studies, but it has been less easy to detect and quantify any such relationships in field-collected material. The aim of the current study was to clarify whether the responses of selected biomarkers (cytochrome P-4501A [CYP1A], delta-aminolevulinic acid dehydratase [ALA-D], metallothionein [MT]) could be correlated with physiological parameters and/or contaminant levels. Atlantic cod (Gadus morhua L.) were sampled over a decade at a number of locations along the Norwegian coast. The locations included both polluted and reference areas. In addition to contaminant-related responses (biomarkers), physiological characteristics were registered for each individual and the contaminant load determined. There were clear relationships between responses in all three biomarkers and tissue contaminants, but additional factors such as age or size and liver lipid or liver somatic index also contributed in multiple-regression models. The results indicated that responses in all three biomarkers reflected relevant, contaminant-related processes in the cod even though none of them provided a simple separation between polluted and pristine locations.

Available evidence suggests that androgens play critical roles in early oocyte growth and development in fish. However, the molecular mechanisms underlying this important aspect of reproductive endocrinology have not yet been established. In this study the effects of androgens (11-ketotestosterone [11-KT] and testosterone [T]) were determined on gene expression patterns and growth of cod previtellogenic oocytes, using an in vitro oocyte culture technique. Previtellogenic ovarian tissue was cultured for 5 and 10 d at different concentrations of 11-KT and T (0, 1, or 1000 microM) dissolved in ethanol (0.3%). The androgen concentrations were selected as they represent physiological and supra-physiological concentrations, respectively. Quantitative polymerase chain reaction (PCR) demonstrated increased mRNA expression for five genes recently identified as androgen responsive in our subtracted cDNA library in previtellogenic cod ovary exposed in vitro to androgens. Quantitative histological analyses showed a consistent stereological validation of oocyte growth and development after exposure to androgens. In general, both 11-KT and T induced previtellogenic oocyte growth and development, and these effects were more pronounced with 11-KT exposure. Taken together, our study reveals some novel roles of androgens on the development of previtellogenic oocytes, indicating control of early follicular and oocyte growth in cod ovary. The potent effects of 11-KT on oocyte growth support our earlier hypothesis that non-aromatizable androgens play significant roles in regulating early oocyte growth with potential consequences for the fecundity process. Therefore, these novel roles of androgens as promoters of ovarian growth and development presented in this study may be useful for the aquaculture industry and for breeding of new captive and endangered species. From a toxicological point of view, the cod is a marine species and exposure to complex chemical mixtures that may exert androgenic and/or anti-androgenic effects represents an environmental issue of reasonable concern in the marine environment. Therefore, the findings in the present study represent a novel basis that can be used to determine the effects of xenoandrogens on oocyte development and fecundity in this important marine species.

Individuals migrate to exploit heterogeneities between spatially separated environments to modulate growth, survival, or reproduction. We devised a bioenergetics model to investigate the evolution of migration distance and its dependence on individual states. Atlantic cod Gadus morhua ranges from sedentary populations to stocks that migrate several thousand kilometers annually. We focused on the Northeast Arctic cod stock, which migrates south to spawn. A linear relationship between migration distance and the expected survival of offspring was assumed, here understood as the prospects for future survival and development that a fertilized egg faces at a particular spawning location. Reasons for why it may increase southward include warmer water that increases development rates, and thereby survival, along the pelagic drift trajectory. In the model, ingested energy can either be allocated to growth or stored for migration and reproduction. When migrating, individuals forgo foraging opportunities and expend energy. Optimal energy allocation and migration strategies were found using state-dependent optimization, with body length, age, condition, and current food availability as individual states. For both a historical and contemporary fishing regime we modeled two behaviors: (1) homing cod returning to the same spawning site each year and (2) roaming cod with no such constraints. The model predicted distinct regions of locally high spawning stock biomass. Large individuals in good condition migrated farthest, and these also tended to mature later in life. The roaming cod spread farther south as they grew larger and older. Homing cod did not have this freedom, and spawning was generally concentrated along a narrower stretch of the coastline. Under contemporary fishing, individuals matured earlier at a smaller size, had shorter migrations, spawned over a contracted geographical range, and tended to be in poorer condition. The effects were most pronounced for the homing behavior.

Serum-mediated control of Listonella anguillarum and transcriptional profiles of selected glucose transport and antioxidant defense genes, following short-term overcrowding in Atlantic cod, Gadus morhua were determined. Fish were subjected to overcrowding by reducing the water level in the tank for 1 h and this was repeated thrice over a 12 h period. Blood samples were collected before overcrowding (initial group) and at 2, 24 and 72 h post-crowding. The sera from fish obtained at 2 h post-crowding caused a significant reduction in L. anguillarum counts compared to the initial samples. There was a transcriptional upregulation of the glucose transport-4 and glyceraldehyde-3-phosphate dehydrogenase genes at 2 h after crowding. Gene transcripts of the antioxidant enzymes, Cu/Zn superoxide dismutase (Cu/Zn SOD), catalase and phospholipid hydroperoxide glutathione peroxidase also significantly increased at 2 h post-crowding, but thereafter they returned to their pre-crowding levels with the exception of Cu/Zn SOD that remained significantly higher than the initial group until 72 h. Thus, short-term overcrowding of Atlantic cod leads to a transient enhancement of in vitro serum antibacterial activity and enhanced transcriptional activity of glucose transport and antioxidant defense genes.

Warming of the global climate is now unequivocal and its impact on Earth' functional units has become more apparent. Here, we show that marine ecosystems are not equally sensitive to climate change and reveal a critical thermal boundary where a small increase in temperature triggers abrupt ecosystem shifts seen across multiple trophic levels. This large-scale boundary is located in regions where abrupt ecosystem shifts have been reported in the North Atlantic sector and thereby allows us to link these shifts by a global common phenomenon. We show that these changes alter the biodiversity and carrying capacity of ecosystems and may, combined with fishing, precipitate the reduction of some stocks of Atlantic cod already severely impacted by exploitation. These findings offer a way to anticipate major ecosystem changes and to propose adaptive strategies for marine exploited resources such as cod in order to minimize social and economic consequences.

Fluorescein isothiocyanate dextran (FITC-dextran) exchange between the primary (PCS) and secondary (SCS) circulatory systems in the Atlantic cod, Gadus morhua (Linnaeus, 1752), were studied using 20-kDa (n = 4) and 500-kDa (n = 4) FITC-dextran. In order to give a qualitative perspective of the general connection between the PCS and SCS, distribution of plasma-borne tracers (FITC-dextran) in the PCS and SCS were examined. In this study, a total of eight cod were cannulated in the ventral aorta (PCS) and dorsal cutaneous vessel (SCS), for investigation of FITC-dextran disappearance in the PCS and its subsequent appearance in the SCS. FITC-dextran of both sizes was found to be in equilibrium between the PCS and SCS in less than 20 min. This indicates a profound connection between the PCS and SCS in the Atlantic cod, and rapid mixing of tracers between the PCS and SCS. The destination of the injected 500-kDa FITC-dextran was also examined, and it was observed that of the 500-kDa FITC-dextran lost from the primary and secondary vascular systems, 63.0 +/- 9.2% could be recovered from the liver.

In this study the potential impact of food chain-based biotransformation and physico-chemical weathering of toxaphene on its tumour promoting potential was investigated in vitro and in vivo. Human exposure to toxaphene is mainly through consumption of contaminated fish, therefore fish-borne residues of toxaphene (cod liver extract, CLE) were prepared by exposing cod to technical toxaphene (TT) for 63 days. UV-irradiated toxaphene (uvT) was included to represent a physico-chemical weathered toxaphene mixture. In vitro, TT, uvT and CLE all showed a dose- and time-dependent inhibition of gap junctional intercellular communication (GJIC) with a relative potency of CLE>TT=uvT. Tumour promoting potency was further studied in vivo in a medium term two-stage initiation/promotion bioassay in female Sprague-Dawley rats, using an increase in altered hepatic foci positive for glutathione-S-transferase-P (AHF-GST-P) as read out. No increase in AHF-GST-P occurred following exposure to either TT, uvT, or CLE, except for the positive control group (2,3,7,8-TCDD). Based on this study the no observed adverse effect level (NOAEL) for tumour promoting potency is at least 12.5mg/kg/week, or higher for CLE. Considering current human exposure levels in Europe it is doubtful that consumption of fish at current levels of toxaphene contamination give rise to human health risk.

Despite recent evidence for sub-stock structuring, North Sea cod are assessed as a single unit. As a consequence, knowledge of sub-stock trends is poor. In particular, there are no recent evaluations of which spawning grounds are active. Here we report results from the first ichthyoplankton survey to cover the whole North Sea. Also, this survey, conducted in 2004, was the first to make extensive use of DNA-based molecular methods to unambiguously identify early developmental stage cod eggs. We compare the findings from the plankton survey with estimated egg production inferred from the distribution of mature cod in contemporaneous trawl surveys. Results from both approaches were in general agreement and showed hot spots of egg production around the southern and eastern edges of the Dogger Bank, in the German Bight, the Moray Firth and to the east of the Shetlands. These areas broadly coincide with known spawning locations from the period 1940 to 1970. We were, however, unable to directly detect significant numbers of cod eggs at the historic spawning ground off Flamborough (northeast coast of England). The results demonstrate that most of the major spawning grounds of cod in the North Sea are still active but that some localized populations may have been reduced to the point where it is now difficult to detect the presence of eggs in the plankton.

1. Relative body condition (the quantity of stored energy) is an important tool in understanding demographic variation and the ability of a population to respond to environmental stressors, varying food availability and competition. 2. A high-resolution database was used to examine causes of variation in the condition of north-east Arctic cod (Gadus morhua L.) for the period 1967-2004, over annual and monthly timescales. Community dynamics and climate variation were also tested as potential causes. 3. Temperature was shown to have a positive impact on condition at both inter- and intra-annual timescales. Inter-annually, temperature may affect stock distribution, in particular its overlap with the capelin stock. At shorter timescales it is likely that temperature directly affects the metabolism of the cod. 4. Intra-annually, the quantity of capelin in cod stomachs positively affected cod condition in the current and the preceding month for all lengths of cod. This indicated a time lag between a change in food consumption and a subsequent change in condition, or 'latency'. 5. Our study has shown that variation in temperature is a vital determinant of changes in condition, both at inter- and intra-annual timescales. Furthermore, the principle of latency has been demonstrated at the population level. Indirect effects of competition for energy-rich resources have been shown to have a negative effect on condition. This study supplements our knowledge of the implications for condition of changes in climate and in potential food resources.

A previous study has reported that Atlantic cod can be conditioned to detect ultrasonic sound pulses of high intensity. This capability has been proposed as a mean for detection and avoidance of echolocating toothed whales that emit intense ultrasonic clicks. In this study, we use acoustic playbacks to test the hypotheses that unconditioned cod can detect and respond to intense ultrasound from toothed whales and from echosounders. Intense ultrasound exposure of 210 dB re. 1microPa (pp) did not cause a short-term stress response in the form of bradycardia in unconditioned cod. Free-swimming cod exposed to ultrasonic clicks and echosounder pulses with received levels of more than 204 dB re. 1 microPa (pp) did not elicit flight responses as seen in ultrasound detecting Alosinae. Furthermore, we tested the debilitating effects of high intensity ultrasound on swimming cod with no detected changes in swimming ability when exposed to more than 213 dB re. 1 microPa (pp). It is concluded that intense ultrasound exposure induces neither an antipredator nor a stress response in Atlantic cod, and that echosounder pulses and biosonar clicks therefore most probably play no ecophysiological role in wild cod populations.

Sea-caged cod are limited in their movements in the water column, and thus can be exposed to large seasonal ( approximately 0-20 degrees C) temperature fluctuations. To investigate the physiological response of Atlantic cod to summer-like increases in temperature, we exposed 10 degrees C acclimated juvenile cod to a graded thermal challenge (1 degrees C increase every 5 days) and measured: (1) plasma cortisol and glucose levels; (2) the respiratory burst activity of blood leukocytes; and (3) the expression of specific immune-related genes [MHC Class I, Interleukin-1beta (IL-1beta), beta2-microglobulin (beta2-M), Immunoglobulin M (IgM)-light (L) and -heavy (H) chains] in the blood using quantitative reverse transcription-polymerase chain reaction (QRT-PCR). The experiment was stopped at 19.1 degrees C, with 26.7% of the fish surviving to this point. Plasma glucose levels increased slightly at 16 and 18 degrees C (by 1.39- and 1.74-fold, respectively), in contrast, cortisol levels were elevated significantly (by 2.9-fold) at 16 degrees C but returned to control levels thereafter. The effect of increasing temperature on the expression of immune related genes in blood cells (leukocytes) was variable and depended on the gene of interest. The expression of IgM-H remained stable for the duration of the experiment. In contrast, IL-1beta expression was increased significantly (by approximately 25-fold) at 19 degrees C as compared to time-matched control fish, and changes in the expression of beta2-M, MHC Class I and IgM-L followed a pattern similar to that seen for cortisol: increasing at 16 degrees C (by 4.2-, 5.3- and 17-fold, respectively), but returning to pre-stress levels by 19 degrees C. Interestingly, increasing temperatures had no effect on respiratory burst activity. This study is the first to examine the effects of a chronic regimen of increasing temperature on the stress physiology and immunology of a marine teleost, and suggests that immune function is influenced by complex interactions between thermal effects and temperature-induced stress (elevated circulating cortisol levels).

Power produced by red myotomal muscles of fish during cruise swimming appears seldom maximized, so we sought to investigate whether economy may impact or dominate muscle function. We measured cost of transport (COT) using oxygen consumption and the strain trajectories and electromyographic activity of red muscle measured at anterior (ANT) and posterior (POST) locations while Atlantic cod (Gadus morhua) swam steadily at speeds between 0.3 and 1.0 body lengths (BL) s(-1). We then measured the power produced by isolated segments of red muscle when activated either as in the swimming cod or such that maximal net power was produced. Patterns of activation during swimming were not optimal for power output and were highly variable between tail beats, particularly at the ANT location and at slow swim speeds. Muscle strain amplitude did not increase until swimming speed reached 0.9 (ANT) versus 0.5 (POST) BL s(-1). These limited power to only 53% (ANT) and 71% (POST) of maximum at slower swim speeds and to 70%-80% of maximum at high swim speeds. COT (resting metabolism subtracted) was minimal at the slowest swim speed, surprisingly, where power was most impaired by activation and strain. Thus, production of powered forces for maneuverability/stability appeared to greatly impact red muscle function during cruise swimming in cod, particularly at slow speeds and in ANT muscle.

Basic knowledge about learning capacities and awareness in fish is lacking. In this study we investigated which temporal gaps Atlantic cod could tolerate between two associated events, using an appetitive trace-conditioning paradigm with blinking light as conditioned stimulus (CS) and dry fish food as unconditioned stimulus (US). CS-US presentations were either temporally overlapping (delay conditioning, CS duration 24 s, interstimulus interval 12 s) or separated by 20, 60, or 120 s (trace conditioning, CS duration 12 s) or 2 h (control, CS duration 12 s). The percentage of fish in the feeding area increased strongly during CS presentation in all delay, 20 s, and 60 s trace groups and in one out of two 120 s trace groups, but not in the control groups. In the 20 and 60 s trace procedures, the fish crowded together in the small feeding area during the trace interval, showing strong anticipatory behaviour. In all the conditioned groups, the fish responded to the CS within eight trials, demonstrating rapid learning. At 88 and 70 days after the end of the conditioning experiments, the delay and 20 s trace groups, respectively, were presented the CS six times at 2-h intervals without reward. All groups responded to the light signal, demonstrating memory retention after at least 3 months. This study demonstrates that Atlantic cod has an impressively good ability to associate two time-separated events and long time retention of learnt associations.

Population growth, and hence the population's persistence, is affected by several factors such as climate, species interaction, and harvesting pressure. Proper resource management requires an understanding of these factors. We apply techniques based upon age-structured population matrices to analyze estimated stock sizes derived from annual bottom trawl sampling in the winter feeding area of northeast Arctic cod (Gadus morhua L.) from 1981 to 2003. We run generalized additive models to explain population growth rate by different explanatory variables. Cod population growth was found to be positively related to the abundance of capelin (Mallotus villosus Miller), negatively related to the number of cannibalistic cod with a two-year lag, and marginally positively related to the winter North Atlantic Oscillation index (NAO). This model remains true independently from the population status (i.e., fished or non-fished). Capelin abundance is the main variable that to some degree can be adjusted in order to maintain the population size at a given level of cod harvesting. Our results point to the importance of managing conjointly cod and capelin stocks.

The present study describes the possibilities for using computer vision-based methods for the detection and monitoring of transient 2D and 3D changes in the geometry of a given product. The rigor contractions of unstressed and stressed fillets of Atlantic salmon (Salmo salar) and Atlantic cod (Gadus morhua) were used as a model system. Gradual changes in fillet shape and size (area, length, width, and roundness) were recorded for 7 and 3 d, respectively. Also, changes in fillet area and height (cross-section profiles) were tracked using a laser beam and a 3D digital camera. Another goal was to compare rigor developments of the 2 species of farmed fish, and whether perimortem stress affected the appearance of the fillets. Some significant changes in fillet size and shape were found (length, width, area, roundness, height) between unstressed and stressed fish during the course of rigor mortis as well as after ice storage (postrigor). However, the observed irreversible stress-related changes were small and would hardly mean anything for postrigor fish processors or consumers. The cod were less stressed (as defined by muscle biochemistry) than the salmon after the 2 species had been subjected to similar stress bouts. Consequently, the difference between the rigor courses of unstressed and stressed fish was more extreme in the case of salmon. However, the maximal whole fish rigor strength was judged to be about the same for both species. Moreover, the reductions in fillet area and length, as well as the increases in width, were basically of similar magnitude for both species. In fact, the increases in fillet roundness and cross-section height were larger for the cod. We conclude that the computer vision method can be used effectively for automated monitoring of changes in 2D and 3D shape and size of fish fillets during rigor mortis and ice storage. In addition, it can be used for grading of fillets according to uniformity in size and shape, as well as measurement of fillet yield measured in thickness. The methods are accurate, rapid, nondestructive, and contact-free and can therefore be regarded as suitable for industrial purposes.

Precocious male puberty significantly compromises sustainability aspects of aquaculture in a number of finfish species. As part of a program aiming to understand and eventually control testis maturation in farmed Atlantic cod, we studied the first reproductive cycle. The gonadosomatic index shows a 41-fold increase from immature (August) to mature (March) stages, reaching almost 10% of the total body weight. The paired cod testes are composed of several lobes arranged around a central collecting duct. In each individual lobe, spermatogenesis occurs in a marked gradient of development, with undifferentiated spermatogonia in the periphery of the lobe and the most advanced germ cells in the vicinity of the collecting duct, suggesting a tight spatiotemporal organization of spermatogenesis in the testis lobes of this species. Spermatogonial proliferation starts in August and continues for about 6 mo. Meiosis and spermiogenesis are first observed in October and are completed in all cysts by February, when a 2-mo-long spawning season starts. Spermatogonia go through 11 mitotic divisions before differentiating to primary spermatocytes. Apoptosis is rare, but when observed it occurs mainly during the last spermatogonial generations. Our observations suggest a model in which a maturational wave progresses through each growing lobe that is first driven by appositional growth from the lobe's periphery, reflecting spermatogonial proliferation and cyst formation which, when ceasing, is terminated by completing spermiogenesis and spermiation that progress toward the lobe's periphery.

Increasing turbidity in coastal waters in the North Atlantic and adjacent seas has raised concerns about impacts on Atlantic cod (Gadus morhua) using these areas as nurseries. A previous experiment (Meager et al. 2005 Can. J. Fish. Aquat. Sci. 62, 1978-1984) has shown that turbidity (up to 28 beam attenuation m-1) had little effect on the foraging rate of juvenile cod. Although this was attributed to cod using chemoreception in conjunction with vision to locate prey, foraging rates may also be maintained by increased activity. Higher activity, however, is energetically costly and may offset benefits from increased foraging return. We examined the effects of turbidity on prey searching and spontaneous activity of juvenile cod in the laboratory, by measuring activity with and without prey cues. Activity of juvenile cod was nonlinearly affected by turbidity and was lower at intermediate turbidity, regardless of the presence of prey odour. Activity increased over time when prey odour was present and decreased when absent, but the effects of prey odour were similar across all turbidity levels. Position in the tank was unaffected by turbidity or prey odour. Reduced activity at intermediate turbidities is likely to offset longer prey-search times. At high turbidity (greater than 17m-1), both longer prey-search times and higher activity indicate that increased energetic costs are probable.

Traditionally, critical swimming speed has been defined as the speed when a fish can no longer propel itself forward, and is exhausted. To gain a better understanding of the metabolic processes at work during a U(crit) swim test, and that lead to fatigue, we developed a method using in vivo (31)P-NMR spectroscopy in combination with a Brett-type swim tunnel. Our data showed that a metabolic transition point is reached when the fish change from using steady state aerobic metabolism to non-steady state anaerobic metabolism, as indicated by a significant increase in inorganic phosphate levels from 0.3+/-0.3 to 9.5+/-3.4 mol g(-1), and a drop in intracellular pH from 7.48+/-0.03 to 6.81+/-0.05 in muscle. This coincides with the point when the fish change gait from subcarangiform swimming to kick-and-glide bursts. As the number of kicks increased, so too did the Pi concentration, and the pH(i) dropped. Both changes were maximal at U(crit). A significant drop in Gibbs free energy change of ATP hydrolysis from -55.6+/-1.4 to -49.8+/-0.7 kJ mol(-1) is argued to have been involved in fatigue. This confirms earlier findings that the traditional definition of U(crit), unlike other critical points that are typically marked by a transition from aerobic to anaerobic metabolism, is the point of complete exhaustion of both aerobic and anaerobic resources.

Much previous research has demonstrated the plasticity of myoglobin concentrations in both cardiac and skeletal myocytes in response to hypoxia and training. No study has yet looked at the effect of thermal acclimation on myoglobin in fish. Atlantic cod (Gadus morhua) from two different populations, i.e. the North Sea and the North East Arctic, were acclimated to 10 and 4 degrees C. Both the myoglobin mRNA and myoglobin protein in cod hearts increased significantly by up to 3.7 and 2.3 fold respectively as a result of acclimation to 4 degrees C. These increments were largest in the Arctic population, which in earlier studies have been shown to possess cold compensated metabolic demands at low temperatures. These metabolic demands associated with higher mitochondrial capacities may have driven the increase in cardiac myoglobin concentrations, in order to support diffusive oxygen supply. At the same time the increase in myoglobin levels may serve further functions during cold acclimation, for example, protection of the cell against reactive oxygen species, and scavenging nitric oxide, thereby contributing to the regulation of mitochondrial volume density.

World-wide many cetaceans drown incidentally in fishing nets. To reduce the unwanted bycatch in gillnets, pingers (acoustic alarms) have been developed that are attached to the nets. In the European Union, pingers will be made compulsory in some areas in 2005 and in others in 2007. However, pingers may effect non-target marine fauna such as fish. Therefore in this study, the effects of seven commercially-available pingers on the behaviour of five North Sea fish species in a large tank were quantified. The species tested were: sea bass (Dicentrarchus labrax), pout (Trisopterus luscus), thicklip mullet (Chelon labrosus), herring (Clupea harengus), and cod (Gadus morhua). The fish were housed as single-species schools of 9-13 individuals in a tank. The behaviour of fish in quiet periods was compared with their behaviour during periods with active pingers. The results varied both between pingers and between fish species. Sea bass decreased their speed in response to one pinger and swam closer to the surface in response to another. Thicklip mullet swam closer to the bottom in response to two pingers and increased their swimming speed in response to one pinger. Herring swam faster in response to one pinger, and pout and cod (close relatives) showed no behavioural responses to any of the pingers. Of the seven pingers tested, four elicited responses in at least one fish species, and three elicited no responses. Whether similar responses would be elicited in these fish species in the wild, and if so, whether such responses would influence the catch rate of fisheries, cannot be derived from the results of this study. However, the results indicate the need for field studies with pingers and fish. Based on the small number of fish species tested, the present study suggests that the higher the frequency of a pinger, the less likely it is to affect the behaviour of marine fish.

A model is developed to describe the interaction between a predator and two prey types located in different regions. Conditions for stability and persistence are analysed. The effects of harvesting the predators are investigated by making the predator mortality rate habitat dependent. Results demonstrate that for any given set of parameter values there is a value of the intrinsic preference of the predator for each prey type at which the system undergoes a Hopf bifurcation. Above this critical value the system evolves towards a stable equilibrium, whereas below it, stable limit cycles arise by Hopf bifurcations. Simulations demonstrate that the presence of demographic stochasticity may destabilise oscillatory populations, thereby causing population extinctions. An application of the model to the foraging behaviour of North Sea cod is described. It is shown that if the preferred prey is more productive, it is likely that the equilibrium will be stable, whereas if the less preferred prey is more productive, populations are likely to display cycles and in the stochastic case become extinct. As cod fishing mortality is increased, the point of bifurcation and region of parameter space for which the system is unstable decreases. An increased understanding of how cod behave may enable fish stocks to be managed more successfully, for example by indicating where marine reserves should be placed.

Atlantic cod populations live in a wide thermal range and can differ genetically and physiologically. Thermal sensitivity of metabolic capacity and swimming performance may vary along a latitudinal gradient, to facilitate performance in distinct thermal environments. To evaluate this hypothesis, we compared the thermal sensitivity of performance in two cod stocks from the Northwest Atlantic that differ in their thermal experience: Gulf of St Lawrence (GSL) and Bay of Fundy (BF). We first compared the metabolic, physiological and swimming performance after short-term thermal change to that at the acclimation temperature (7 degrees C) for one stock (GSL), before comparing the performance of the two stocks after short-term thermal change. For cod from GSL, standard metabolism (SMR) increased with temperature, while active metabolism (AMR, measured in the critical swimming tests), EMR (metabolic rate after an exhaustive chase protocol), aerobic scope (AS) and critical swimming speeds (U (crit) and U (b-c)) were lower at 3 degrees C than 7 or 11 degrees C. In contrast, anaerobic swimming (sprint and burst-coasts in U (crit) test) was lower at 11 than 7 or 3 degrees C. Factorial AS (AMR SMR(-1)) decreased as temperature rose. Time to exhaustion (chase protocol) was not influenced by temperature. The two stocks differed little in the thermal sensitivities of metabolism or swimming. GSL cod had a higher SMR than BF cod despite similar AMR and AS. This led factorial AS to be significantly higher for the southern stock. Despite these metabolic differences, cod from the two stocks did not differ in their U (crit) speeds. BF cod were better sprinters at both temperatures. Cod from GSL had a lower aerobic cost of swimming at intermediate speeds than those from BF, particularly at low temperature. Only the activity of cytochrome C oxidase (CCO) in white muscle differed between stocks. No enzymatic correlates were found for swimming capacities, but oxygen consumption was best correlated with CCO activity in the ventricle for both stocks. Overall, the stocks differed in their cost of maintenance, cost of transport and sprint capacity, while maintaining comparable thermal sensitivities.

The North Sea has warmed in recent years and there is an ongoing debate into how this is affecting the distribution of fishes and other marine organisms. Of particular interest is the commercially important Atlantic cod (Gadus morhua L.), which has declined sharply in abundance in the North Sea over the past 20 years. Observations of the temperature experienced by 129 individual cod throughout the North Sea were made during a large-scale electronic tagging programme conducted between 1999 and 2005. We asked whether individual cod fully occupied the thermal habitat available to them. To this end, we compared the temperature experience of cod with independently measured contemporaneous sea-bottom temperature data. The majority of cod experienced a warmer fraction of the sea than was potentially available to them. By summer, most of the individuals in the south experienced temperatures considered superoptimal for growth. Cooler waters were within the reach of the cod and a small number of individuals migrated to areas that allowed them to experience lower temperatures, indicating that the cod had the capacity to find cooler water. Most did not, however, suggesting that the changing thermal regime of the North Sea is not yet causing adult cod to move to cooler waters.

The majority of survival analyses focus on temporal scales. Consequently, there is a limited understanding of how species survival varies over space and, ultimately, how spatial variability in the environment affects the temporal dynamics of species abundance. Using data from the Barents Sea, we study the spatiotemporal variability of the juvenile Atlantic cod (Gadus morhua) survival. We develop an index of spatial survival based on changes of juvenile cod distribution through their first winter of life (from age-0 to age-1) and study its variability in relation to biotic and abiotic factors. Over the 25 years analyzed (1980-2004), we found that, once the effect of passive drift due to dominant currents is accounted for, the area where age-0 cod survival was lowest coincided with the area of highest abundance of older cod. Within this critical region, the survival of age-0 cod was negatively affected by its own abundance, by that of older cod, and by bottom depth. Furthermore, during cold years, age-0 cod survival increased in the eastern and coldest portion of the examined area, which was typically avoided by older conspecifics. Based on these results we propose that within the examined area top-down mechanisms and predation-driven density dependence can strongly affect the spatial pattern of age-0 cod survival. Climate-related variables can also influence the spatial survival of age-0 cod by affecting their distribution and that of their predators. Results from these and similar studies, focusing on the spatial variability of survival rates, can be used to characterize species habitat quality of marine renewable resources.

We present the first data on the effect of hypoxia on the specific dynamic action (SDA) in a teleost fish. Juvenile cod (Gadus morhua) were fed meals of 2.5% and 5% of their wet body mass (BM) in normoxia (19.8 kPa Po(2)) and 5% BM in hypoxia (6.3 kPa Po(2)). Reduced O(2) availability depressed the postprandial peaks of oxygen consumption, and to compensate for this, the total SDA duration lasted 212.0+/-20 h in hypoxia, compared with 95.1+/-25 h in normoxia. The percentage of energy associated with the meal digestion and assimilation (SDA coefficient) was equivalent between the different feeding rations but higher for fish exposed to hypoxia. Comparing peak oxygen consumption during the SDA course with maximum metabolic rates showed that food rations of 2.5% and 5% BM reduced the scope for activity by 40% and 55%, while ingestion of 5% BM in hypoxia occupied 69% of the aerobic scope, leaving little energy for other activities.

Neuropeptide Y (NPY) and cocaine and amphetamine regulated transcript (CART) are two neuropeptides involved in the regulation of feeding in both mammals and fish. NPY stimulates feeding whereas CART inhibits feeding. In this study, we have cloned the full-length cDNA and complete genomic DNA sequences for NPY and CART in Atlantic cod. Atlantic cod preproNPY share a 45-85% identity with preproNPY from other fish whereas preproCART shows a 70% identity to CART peptides from zebrafish and goldfish. RT-PCR revealed that NPY mRNA is expressed in brain, in particular the forebrain, and in peripheral tissues, including intestine and kidney. CART mRNA is expressed throughout the brain and in ovaries. In order to assess the role of these peptides in the regulation of feeding, we examined changes in mRNA expression in the forebrain before, during and after a meal. NPY and CART mRNA both undergo peri-prandial changes in expression, with NPY levels being elevated around meal time and CART showing a decline 2 h after a meal. Food deprivation for 7 days induced a decrease in CART mRNA expression in the brain but did not affect NPY mRNA expression. Overall, our results suggest that NPY and CART are conserved peptides that might be involved in the regulation of feeding and other physiological functions in Atlantic cod.

Produced water, a by-product of offshore oil production, contains significant amounts of alkylphenols (APs). Many studies have shown that APs cause endocrine disruption in marine organisms, but relatively little is currently known about their long-term effects on the biology of pelagic fish. Here, we describe in detail the effects of APs on the reproductive potential of first-time spawning Atlantic cod (Gadus morhua). Cod were fed with feed paste containing four APs (4-tert-butylphenol, 4-n-pentylphenol, 4-n-hexylphenol and 4-n-heptylphenol), at a range of concentrations, for either 1 or 5 weeks. AP-exposed fish were compared to unexposed fish and to fish fed paste containing natural estrogen (17beta-estradiol). Our results showed that in female fish AP exposure impaired oocyte development, reduced estrogen levels, and delayed the estimated time of spawning by 17-28 days. Similarly, in male fish, we observed that AP-exposure reduced 11-keto-testosterone concentrations, and caused a small induction of VTG levels. We also observed impaired testicular development, with an increase in the amount of spermatogonia and a reduction in the amount of spermatozoa present. Taken together these results suggest that APs released into the sea via produced water may have a negative influence on the overall reproductive fitness of cod populations.