Antimicrobial activity in serum of Atlantic cod Gadus morhua L.: comparison between cultured and wild fish

The present investigation determined the antibacterial activities in the sera of both cultured and wild Atlantic cod Gadus morhua . Serum samples from both groups of fish were quantified for total protein, and their effects against fish pathogens Listonella anguillarum , Aeromonas salmonicida and Yersinia ruckeri were determined using co-incubation assay. Total serum protein concentrations were not significantly different between farmed and wild Atlantic cod. Sera of cultured Atlantic cod significantly decreased growth of L. anguillarum by at least two-log₁₀ reductions in bacterial count, while those of the wild Atlantic cod were able to inhibit the growth of all three fish bacterial pathogens. The present study showed that sera from wild fish possess broader antibacterial activities than cultured Atlantic cod and that these could provide an explanation for the differences regarding their immunity to bacterial infections as well as their health status.     

Functional relationship between sealworm (Pseudoterranova decipiens, Nematoda, Ascaridoidea) burden and host size in Atlantic cod (Gadus morhua).

A model is proposed to describe the accumulation of sealworm parasites (Pseudoterranova decipiens, Nematoda, Ascaridoidae) in Atlantic cod (Gadus morhua). The model predicts the total amount of food eaten by a fish to reach its size. In young fish growth is linear in time and average parasite burdens are predicted to increase with the cube of the average length of the fish divided by its growth rate. An analysis of worm burdens in cod caught by the commercial fishery in the Firth of Clyde (Strathclyde, Scotland) between 1964 and 1970 illustrates how average parasite abundances at age, length, or fixed time of the year can be compared. It is concluded that exposure did not vary significantly over these years. Extensive apparent inter-annual variations in worm burdens of cod newly recruited to the fishery are well explained by differences in the age composition of cod catches.     

Effects of Body Size and Food Ration on Over-winter Survival and Growth of Age-0 Atlantic cod, Gadus morhua

This study examined the effects of body size and food ration on over-winter survival of age-0 Atlantic cod. Cod were divided into two groups based on standard length (large=76.95±1.10 mm SL; small=57.65±1.02 mm SL; mean±1 SE) and wet weight (large=4.02± 0.21 g; small=1.52±0.09 g). Replicate tanks (n=2) of 10 large and 10 small cod were exposed to one of two food rations (0.25% and 1.0% body weight day^-1) for the entire experiment (December to June). Tanks were examined daily for mortalities and feeding was adjusted accordingly. The experiment was run under ambient light and seawater conditions. All but large age-0 cod exposed to the low food ration grew over the course of the experiment. The specific growth rate (SGR) of small cod was significantly higher (0.2425% wet weight day^-1) than that of the large fish (0.0443% wet weight day^-1). Food ration had no significant influence on SGR or over-winter survival. Significantly more of the large age-0 cod survived the winter (58.5% of those originally introduced) compared to the smaller fish (14%). Our results are consistent with those from studies of several other fish species, and are discussed in relation to the ecology of age-0 cod, and their potential use in aquaculture in Newfoundland.     

Winter feeding ecology of cod (Gadus morhua) in a fjord of southern Norway

We investigated the winter feeding ecology of cod ( Gadus morhua ) in the Risørfjord and Flødevigen areas on the Skagerrak coast, southern Norway. Diets from the ice-covered Risørfjord were compared with diets from the more coastal Flødevigen area. In the Risørfjord area the diet featured numerically both decapods (56.0%) and fish (27.8%), but fish dominated by mass (75.1%). The most numerous dietary items from the Flødevigen area were decapods (48.1%) and isopods (30.6%), although fish still made up the bulk of the diet's mass (52.6%). Diets at Flødevigen shifted from winter to spring, as polychaetes became important numerically (67.8%) and also contributed substantially by weight (53.6%). Since cod feed opportunistically, differences between areas and seasons probably reflected differences in prey diversity and abundance. Prey size variability increased with increasing predator length, but maximum prey size relative to predator length was constant at about 9%. Prey fish increased in both length and numbers with the length of cod, particularly in the Risørfjord area, where fish were more important in the diet than at Flødevigen. However, the majority of fish consumed in both areas were small gobiids. Based on growth patterns observed in otoliths, with opaque zones (indicating faster growth) formed during winter, the quality and quantity of the winter diet may determine annual growth rates of cod on the Skagerrak coast. Fish and decapods constituted important, high energy food sources, and there was little indication that cod were food limited during winter in this area.     

Inter‐individual differences in rates of routine energy loss and growth in young‐of‐the‐year juvenile Atlantic cod

Inter‐individual differences in rates of routine (non‐feeding) metabolism and growth were evaluated in young‐of‐the‐year (YOY) juvenile Atlantic cod Gadus morhua . Rates of O₂ consumption, CO₂ production and ammonia (TAN) excretion were measured in 64, 25–43 mm standard length ( L _S) YOY growing at different rates (0·27–0·47 mm day⁻¹) in a common rearing tank. Parameter rates ( y ) increased allometrically ( y = a·M^b ) with increasing body mass ( M ) with b ‐values for O₂ production, CO₂ consumption and TAN excretion equal to 0·81, 0·89 and 0·56, respectively. In some cases, residuals from these regressions were significantly negatively correlated to fish growth rate. In no cases did residuals of parameter rates increase with increasing growth rate. These data suggest that, during unfed periods, relatively fast‐growing fish were more metabolically efficient than slower‐growing fish from the same cohort. The fish condition factor, derived from     , also significantly decreased with increasing growth rate. Results indicated differences in both the rates of routine energy loss and the patterns of growth allocation among YOY Atlantic cod. Since these physiological attributes were positively correlated with growth rate, they may be indicative of 'survivors' in field populations.     

The feeding, growth and behaviour of juvenile cod, Gadus morhua L., in cold environments

Most studies on feeding-related activities of fish in cold water (0° C) have not used eurythermal species or examined a broad suite of activities. In this study we report on the feeding, growth, behaviour and conversion-efficiency of 0 + Atlantic cod in response to water temperature and food availability. In one experiment, activity, opercular beat and growth decreased with decreasing temperature over the range 8·3–0·6° C. However, the conversion efficiency increased with decreasing temperature, a result partly explained by the relative change in activity and opercular rate. In another experiment, temperature had a linear effect on growth in cod fed different food rations. Both food availability and temperature were found to influence growth in group 0 + cod. The specific growth rate of cod in this experiment exceeded 2% day^-1. Growth data collected from cod sampled in the field during the cold-water period showed a progressive increase of mean length over the season. Overall, these results demonstrate that 0 + cod have adapted to life in cold-water environments, in contrast to adult cod which mostly inhabit warmer environments.     

Prevalence and influence of Loma branchialis (Microspora) on growth and mortality in Atlantic cod (Gadus morhua) in coastal Newfoundland

A study was conducted to determine the prevalence, geographical distribution, and effects of a microsporan parasite, Loma branchialis, in Atlantic cod in coastal Newfoundland. The parasite was identified microscopically as opaque, ovoid xenomas, primarily in the gills, heart, and spleen. Although widely distributed in young fish inhabiting 7 coastal localities, prevalence was 1-7% in general, <0.01% in older and large fish, but 78% in market-size cod from a commercial farm. Some of the latter fish were emaciated, and approximately 2 mo after the parasite was detected, 63% succumbed with a massive infection. Condition factor and blood values were significantly lower than those in reference fish. An experimental study conducted over 2 yr revealed that mass gained and condition factor were significantly lower in infected cod than in controls at the termination of the study. Feed consumption, mass gained, feed conversion efficiency, and condition factor also were significantly lower in another group of infected cod of marketable size studied over 12 wk. Xenomas also were observed in the heart and gills of year class II cod reared in a hatchery. Loma branchialis is considered to be a potential pathogen in both hatchery-reared and net pen-cultured cod in Newfoundland.     

Growth studies with fish—overcoming the problems of size variation

The relationship between growth rate and fish size is described by the equation log_e G _w= a —0·4 log_e W, where G_w is the specific growth rate and W is fish weight. Since the intercept (a) represents the log_e G_w of a fish unit size, the relationship presents a method allowing comparison of data from experiments involving fish of different sizes. The application of the method is demonstrated by examining the effects of environmental temperature on growth rates of cod, Gadus morhua , and it is suggested that the optimum temperature for growth of cod is 13–15° C.     

Evidence that size-selective mortality affects growth of Atlantic cod (Gadus morhua L.) in the southern Gulf of St Lawrence

The effects of size-selective fishing mortality on the growth of Atlantic cod ( Gadus morhuu L.) in the southern Gulf of St Lawrence were investigated and compared between: (1) a period when fishing mortality was relatively high, growth was relatively rapid, and abundance low (1967–1972 year classes): and (2) a period when fishing mortality was lower, growth was slow, and density high (1977–1982 year classes). Cod first entered the fishery at age 3 during both periods. The 1967–1972 year classes (fast growing) were fully recruited to the fishery by age 5 or 6 and the fishery removed over twice as many fish from the lower than upper quartiles of length-at-age distributions for cod 4 to 10 years old (disproportionately high exploitation of slow-growing fish). In contrast. the 1977–1982 year classes (slow growing) did not fully recruit to the fishery until age 9 or 10 and the fishery removed four times as many fish from the upper than lower quartiles of the length-at-age distributions for 4- to 10-year-old cod (disproportionately high exploitation of fast-growing fish). The reduced mean lengths-at-age of the 1977–1982 year classes compared with the 1967–I972 year-classes is consistent with the different patterns of exploitation of fast- and slow-growing fish for the two periods.     

Is reduced body growth of cod exposed to the gill parasite Lernaeocera branchialis a cost of resistance?

Three hundred and sixty-nine cod Gadus morhua were individually marked and caged for 19 months. During this period, each cod was inspected several times for Lernaeocera branchialis . Growth in four groups of cod, identified by their infection history, were compared. During the caging, 79% of the cod remained uninfected, 8·5% were infected, but lost the parasite, 8% were infected with one parasite and 4·5% were infected with more than one parasite. The infected fish either harboured the parasite at caging or were infected during the study period. The highest rate of increase, both in body mass and in standard length ( L _S), was recorded in the group of male fish infected with one parasite throughout the experimental period. Conversely, those males free from infection showed significantly lower growth. The observed differences in growth could not be explained by changes in variables related to reproductive strategies. The alternative explanation for these results is that resistance to L. branchialis was associated with costs in terms of reduced growth of body mass and L _S.     

Specific growth rate and proximate body composition of Atlantic cod (Gadus morhua L.)

The effect of growth rate and maturation on the proximate composition and energy content of Atlantic cod, Gadus morhua L., was investigated over 10 months for each of two consecutive years, 1978–1980 at 5 and 8 °C. Relative energy and lipid content of whole cod increased with specific growth rate for all three sampling periods (November, January, March), each at 5 and 8 °C. Relative water content decreased with specific growth rate and temperature, and was lower in March than in January and November. Relative protein content was positively correlated with specific growth rate, but to a lesser degree than with temperature and age. Relative ash content was negatively correlated with specific growth rate. The effect of season and temperature on the proximate content of gonad, liver, muscle, and carcass was also determined. The major energy and lipid source in cod was the liver. Energy, lipid, and water were highly correlated to each other, and regressions are provided to allow for their prediction, given one of the components. Energy budgets for cod at 5 and 8 °C are calculated and the effect of increased ration size on the budget is estimated. The prediction of short-term specific growth rates of cod from the proximate composition is proposed. The proximate composition of cod is affected by growth rate and thus feeding level, and in turn directly affects behaviour. The relative proximate content of maturing and immature 3-yr-old cod was not found to be significantly different. Keywords: specific growth rate; proximate; bioenergetics; Atlantic cod; energy budget; temperature; season     

A century of fish growth in relation to climate change,population dynamics and exploitation

Marine ecosystems, particularly in high‐latitude regions such as the Arctic, have been significantly affected by human activities and contributions to climate change. Evaluating how fish populations responded to past changes in their environment is helpful for evaluating their future patterns, but is often hindered by the lack of long‐term biological data available. Using otolith increments of Northeast Arctic cod (Gadus morhua) as a proxy for individual growth, we developed a century‐scale biochronology (1924–2014) based on the measurements of 3,894 fish, which revealed significant variations in cod growth over the last 91 years. We combined mixed‐effect modeling and path analysis to relate these growth variations to selected climate, population and fishing‐related factors. Cod growth was negatively related to cod population size and positively related to capelin population size, one of the most important prey items. This suggests that density‐dependent effects are the main source of growth variability due to competition for resources and cannibalism. Growth was also positively correlated with warming sea temperatures but negatively correlated with the Atlantic Multidecadal Oscillation, suggesting contrasting effects of climate warming at different spatial scales. Fishing pressure had a significant but weak negative direct impact on growth. Additionally, path analysis revealed that the selected growth factors were interrelated. Capelin biomass was positively related to sea temperature and negatively influenced by herring biomass, while cod biomass was mainly driven by fishing mortality. Together, these results give a better understanding of how multiple interacting factors have shaped cod growth throughout a century, both directly and indirectly.     

The effects of body mass and feeding on metabolic rate in small juvenile Atlantic cod

Apparent specific dynamic action (SDA) amplitude in young juvenile Atlantic cod Gadus morhua (1 to 8 g wet mass), fed a standardized meal and then exercised in a circular swimming respirometer at a constant swimming speed of 0·5 ± 0·3 body lengths s^-1, occurred within l h after feeding in all juveniles. SDA amplitude were 1·4 to 1·8 times higher in fed fish compared to unfed fish, and rates of oxygen consumption decreased as body mass increased. SDA duration had a tendency to decrease with increasing body mass and was shortest, at 6 h, in the smallest fish (1–1·5 g), but increased to 10–11 h in the largest fish. Apparent SDA in fed fish ( R _r) scaled with a mass exponent of 0·89, while maximum metabolic rate ( R _max) determined by chasing fish to exhaustion and then measuring oxygen consumption for 12 h, and unfed routine metabolic rate (R_r) scaled with a mass exponent of 0·79 and 0·76 respectively. Relative aerobic scope ( R _max– unfed R _r) did not appear to vary over the 1 to 8 g increase in wet mass. These results show that as body mass increased in young juvenile Atlantic cod: (1) apparent SDA ( R _f) increased more rapidly than R _max, and (2) apparent SDA took up >98% of the relative aerobic scope and that young Atlantic cod allocated most of the energy to growth, and left little for other metabolic activities.     

Detection of TIMP-2-like protein in Atlantic cod (Gadus morhua) muscle using two-dimensional real-time reverse zymography

Matrix metalloproteinases (MMPs) have been proposed to participate in postmortem degradation of fish muscle connective tissues during storage. In the extracellular matrix (ECM) of mammals, a group of specific tissue inhibitors of metalloproteinases (TIMPs) contributes in regulating the MMPs present. However, little information exists on the presence of TIMPs in fish. In this paper, the presence of TIMPs in the muscle of Atlantic cod (Gadus morhua) was investigated using gelatin affinity chromatography, real-time reverse zymography (RTRZ) and mass spectrometry (MS). Using RTRZ inhibitory action against cod muscle, proteinases binding to gelatin were detected in the muscle. The inhibitor had similar molecular weight (21 kDa) as a human recombinant TIMP-2 used as a reference sample. Because isoforms of TIMP-2 homologues with similar molecular weight have been suggested in fish, a two-dimensional RTRZ (2D RTRZ) method was designed. The new method showed the existence of only one form with inhibitory action against cod muscle proteinases. Finally, de novo sequencing of two peptides derived from the cod muscle inhibitor showed high homology to TIMP-2s both from human and other teleosts.     

Restricted fish feeding reduces cod otolith opacity

The purpose of this work was to examine the effect of reduced feeding and constant temperature on cod otolith opacity. Three groups of juvenile cod were given restricted food rations at different times for 4 months, resulting in depressed somatic growth. Otolith opacity was measured on pictures of the otolith sections. The otolith carbonate deposited during the experimental period was generally opaque compared to the more translucent otolith material deposited prior to and after the experimental period, when the fish were kept in a pond and in sea‐cages at higher temperatures. Large variations in otolith opacity were found between individual fish both within groups and between groups. In two of the three groups significantly more translucent otolith material was deposited in response to reduced feeding. Our results show that variations in feeding and hence fish growth resulted in variation in otolith opacity, but the effect was minor compared to that of variations in ambient temperature. The combined influence of these effects, which both act on fish metabolism, are most likely controlling the seasonal opacity changes observed in wild fish. Our results help explain the variations seen in fish at constant temperatures.     

Genetic Adaptation to Elevated Carbon Dioxide Atmospheres by Pseudomonas-Like Bacteria Isolated from Rock Cod (Sebastes spp.)

The microorganisms on rock cod fillets stored in a modified atmosphere (MA; 80% CO₂-20% air) at 4°C for 21 days were isolated. Only Lactobacillus sp. (71 to 87%) and tan-colored Pseudomonas sp.-like isolates (TAN isolates) were found. The TAN isolates grew more slowly in MA than in air at 8°C. When TAN isolates were grown in air at 8°C and then transferred to MA at 8°C, there was an initial decline in viable counts for 10 to 30 h followed by exponential growth. During this exponential growth phase in MA, the growth rates of the TAN isolates from MA-stored fish were significantly greater than those of the TAN isolates from fresh fish. When a TAN isolate from fresh fish was grown under MA for 21 days, it then grew as rapidly under MA as isolates from MA-stored fish. These results suggest that the TAN isolates genetically adapt to high levels of CO₂.     

Influence of Lernaeocera branchialis (Crustacea: Copepoda) on growth rate of Atlantic cod, Gadus morhua

Influence of Lernaeocera branchialis, a hematophagous copepod, on the growth rate of Atlantic cod, Gadus morhua, was investigated in fish of various sizes. Initially, it appeared that growth, over a 16-mo period, was reduced when single or multiple infections were present. We then examined growth rate by comparing food consumption, weight gain, food conversion efficiency, and condition (k) factor over a 2-mo period and subsequently over a 4 wk in control and experimentally infected fish. Total body weight and length of each fish, controls and infected were determined before and after feeding known weights of food. Adults infected with young parasites consumed more food and gained more weight than controls, especially in the first month, but food conversion efficiency and k-factor were greater than those of controls after 2 mo, but food efficiency was still lower than in controls. In 2 groups of young infected fish, all values were consistently lower than those of uninfected fish of comparable size during the 2-mo period. Three additional trials on immature cod confirmed that infected fish consumed less food and exhibited less growth than infected controls. Our results suggest that only adult cod infected with immature parasites compensate for the infections by increasing food consumption, thus resulting in a transitory weight gain. Additionally, parasites that have completed their reproductive activities still produce subtle effects in the form of low food conversion efficiency in their hosts.     

Diet of juvenile cod (age 0–2) in the Barents Sea in relation to food availability and cod growth

Diet investigations were carried out on 0-, 1- and 2-year-old Northeast Arctic cod (Gadus morhua) sampled in the Barents Sea during 1984–2002. Stomach-content analyses showed that the 0 and 1 group cod fed mainly on crustaceans, with krill and amphipods composing up to 70% of their diet. Krill (Thysanoessa spp. and Meganyctiphanes norvegica) and amphipods (Themisto spp.) were mainly found in cod stomachs sampled in the central and close to the Polar Front region in the Barents Sea where these prey organisms are reported to be abundant in summer. A shift in the main diet from crustaceans to fish was observed from age 1 to age 2. The diet of 2-year-old cod mainly comprised capelin (Mallotus villosus) and other fish, and to a lesser degree, krill and amphipods. Shrimp (mainly Pandalus spp.) was also an important prey in both age 1 and 2 cod. A statistically significant positive relationship was obtained between capelin stock size and the amount of capelin in the diet of 2-year-old cod. Results from this study also show that the larger age-2 cod preyed more on capelin in winter and that larger cod (>22 cm) prefer larger capelin (>12 cm). During periods of low capelin abundance, the 2-year-old cod shift their diet more to crustaceans, such as krill and amphipods. A positive significant relationship was also obtained between Total Fullness Index (TFI) and the amount of capelin in the diet and between TFI and the growth of 2-year-old cod, indicating that the growth of age-2 cod is to a large extent dependent on the amount of capelin consumed. Growth of age-1 cod was also positively correlated to TFI.