Spectral sensitivity of the cod,Gadus morhua L.

The spectral sensitivity of the cod was determined under both dark adapted and light adapted conditions in the laboratory. Cod were trained by cardiac conditioning to detect a difference in radiance between an image of spots and the background radiance of a screen. Thresholds for this response were measured for a range of different wavelengths, and expressed as quantum adjusted values. Electroretino‐graphic studies were also performed on the eyes of cod, and spectral sensitivity curves prepared. Under dark adapted conditions both the behavioural and e.r.g. derived curves showed greatest sensitivity in the blue/green at 490 nm, matching the absorption curve for rhodopsin. A secondary peak in the behaviourally derived curve in the green/yellow at 550 nm indicated that a population of yellow cones may be implicated with the rods in scotopic vision. Under light adapted conditions the behavioural curves showed a shift to the blue, perhaps indicating an adaption to the high red content of the illuminating source. The e.r.g. curve showed greatest sensitivity to blue/green, as in the scotopic experiments but with an enhanced response at 550 nm, indicating greater cone activity. It is suggested that there is complex interaction between rods and cones in the cod retina, both types of receptor being active over a wide range of light intensities.     

The spawning activity of cod, Gadus morhua L.

Aspects of the reproduction of reared cod, Gadus morhua L., with special emphasis on the females, were studied under laboratory conditions. The fecundity and condition factor were 2–5 and 1–5 times, respectively, that of wild cod. A total of 18 spawning females were kept in separate tanks/ chambers, each with one or two males. Seven of the 18 females were classified as stressed, based upon behaviour, irregular spawning intervals and low fertilization rates of the eggs. The reared cod were found to spawn 17–19 batches. The number of eggs liberated in each batch normally followed a smooth, dome-shaped curve with time. The fertilization rate was normally 100%. Egg size decreased from first to last batch and the egg dry weight decreased by about 20–30%. The reared cod showed the same egg diameter to dry weight relation as wild cod. Egg diameter of first batch and maternal fish length were significantly positively correlated. The mean spawning interval for the female and the mean water temperature during its spawning were negatively correlated. The reared cod spawned in both the night and the day for about 50–60 days.     

Visual contrast thresholds in the cod Gadus morhua L.

The contrast discrimination of Gadus morhua L. was studied by means of a cardiac conditioning technique. Fish were trained to respond to a projected pattern of spots and the contrast of pattern and background then reduced until the minimum contrast required to elicit a response was established. This was repeated at seven different background light levels to cover the range of light conditions naturally encountered by the cod. The minimum detectable contrast decreased with increasing light level to a minimum of around 2.0%. The contrast threshold curve showed a discontinuity at a light level of approximately 8.0 × 10^-6 W sr^-1 m^-2. This was thought to be linked to the change from photopic to scotopic vision. Significant differences were found in cod taken from two different locations. Overall, cod contrast detection compared very favourably with figures available for other species including man. Using the obtained contrast discrimination figures some estimates were made of the cod sighting distance of hypothetical targets under natural conditions.     

Tracking cod Gadus morhua L. in a Scottish sea loch

Equipment has been developed and used to follow the movements of free-swimming individual fish in the sea. A pulsed ultrasonic transmitter inserted into the stomach is tracked by measuring differences in the time of arrival of the acoustic pulses at an array of omnidirectional hydrophones. The method was used to plot the movements of cod in an enclosed sea-loch. After an initial phase of active movement the cod became less mobile and adopted a nocturnal cycle of activity.     

Oxygen uptake in developing eggs and larvae of the cod, Gadus morhua L.

Unfertilised cod eggs showed a mean oxygen uptake rate at 5°C of 0.089 μl O₂, dry wt.⁻¹ h⁻¹; this gradually rose to 0.768 μl O₂ mg dry wt.⁻¹ h⁻¹ in eggs about to hatch. From hatching to complete yolk absorption larvae respired at 1.6 μl O₂, mg dry wt.⁻¹ h⁻¹. During starvation following yolk absorption, uptake fell significantly to 1.1 μl O₂, mg dry ⁻¹ h⁻¹. Much of this decrease in oxygen consumption was shown to be caused by reduction in activity. Loss of weight during the embryo and larval phases could not easily be reconciled with total oxygen consumption; it is suggested that cod embryos and larvae may not rely solely upon endogenous energy reserves during development.     

Light microscopical studies of chromosomes in embryos of cod, Gadus morhua L.

The chromosomes in the cells of 1–2-day-old cod embryos were studied using light microscopic techniques. The observations revealed slightly non-synchronous cell divisions, with at least three different mitotic phases present in a 1-day-old embryo. Surprisingly high percentages of abnormal metaphases and anaphases were found in normal-looking embryos. In the 2-day-old embryos, free-lying nuclei were observed in the periblast zone. Chromosome studies of these nuclei revealed a cleavage pattern of the chromosomes which differed from that of the other cells.     

The demography of a haemoglobin polymorphism in the Atlantic cod, Gadus morhua L.

Genetic stability was investigated at a polymorphic haemoglobin gene-locus in 13 124 Atlantic cod, Gadus morhua L. A total of 30 year-classes, sampled over 18 years, were variously sampled in 33 named localities. Regional populations showed distinctive allele frequencies, and each major cod fishery showed some degree of genetic imbalance, which was expressed as excessive numbers of homozygotes. This imbalance occurred seasonally in a proportion of the population samples in each of the major cod fisheries. The findings are attributable to migrations of distinctive genetic populations.     

Aerobic metabolic scope and swimming performance in juvenile cod, Gadus morhua L.

Juvenile cod (Gadus morhua) were made to swim in a tunnel respirometer to determine the oxygen consumption during swimming at different speeds. Results were compared with measurements of standard and active metabolic rates in static respirometers before and after intense exercise. The oxygen consumption at maximum sustainable swimming speed was considerably lower than the peak oxygen consumption following exhausting exercise. It is suggested that these fish have a poorly developed system of aerobic (red) locomotor muscles which do not normally make a major demand upon oxygen consumption. Apparent specific dynamic action following feeding and repayment of oxygen debt following anaerobic exercise can each give rise to greater rates of oxygen consumption. Following exhausting exercise there is a delay of about 1 h before oxygen consumption reaches a peak level some 40% higher than the peak level observed during sustained swimming.     

The detection of dissolved amino acids by the Atlantic cod, Gadus morhua L.

A classical conditioning technique was employed with cod, Gadus morhua L., to determine thresholds for the detection of the L-forms of some α amino acids which are thought to be attractants or feeding stimulants for fish. The amino acids investigated, in order of effectiveness, were tyrosine, cysteine, phenylalanine, glycine and methionine with mean threshold response levels ranging from 2·5 × 10^-8M to 7·4 × 10^-8M. Histidine and lysine resulted in similar thresholds with a mean value of approximately 3 ± 10^-7M while taurine and leucine were least effective with mean threshold levels of 2·1 × 10^-6M and 2·1 × 10^-5M respectively. Comparison is made with electrophysiological and behavioural response data from other species. The effect of raising the background level of glycine on the threshold to glycine for cod is described. The results are discussed with reference to data on levels of dissolved free amino acids in shallow sea waters which may have a bearing on determining chemosensory threshold levels. To detect a specific amino acid against a background level of the same substance the difference in level for detection is proportionally greater for higher background concentrations.     

Fertility of gynogenetic Atlantic cod (Gadus morhua L.)

In order to investigate whether meiotic gynogenetic Atlantic cod is fertile and able to produce viable offspring, meiotic gynogenetic females were produced in spring 2010 by activating cod eggs using irradiated sperm. The extrusion of the second polar body was prevented by the application of hydrostatic pressure (56.6 MPa) 36 min after fertilization. In February 2012, their mean round weight was 972 g, and 2580 g in March 2013. In 2012, when the fish were 2 years old, about 52% were mature, 33% were immature, and 13% had undifferentiated gonads. One year later, 77% were mature, 11% were immature, and 11% had undifferentiated gonads. Several of the mature females had malformed gonads, with only one developed ovary lobe or with the two lobes fused. The mean gonadosomic index (GSI) of the 2‐year‐old mature females was 5.2%, with an estimated relative fecundity of 581 000 eggs kg ovary‐free wet weight^?1. Females were stripped for eggs when 2 and 3 years old (2012 and 2013), and fertilized with sperm from normal males. Offspring were obtained from 12 of 17 and 12 of 15 egg batches incubated in 2012 and 2013, respectively, proving that the gynogenetic females are fertile. Furthermore, larvae in all but one of the hatched groups from 2013 had commenced feeding 2 h after being startfed using rotifers 4 days after hatch, indicating viable offspring.     

Immune parameters of immunised cod (Gadus morhua L.)

The immune response of cod (Gadus morhua L.) is unusual in that specific antibody response is limited or absent. In the present study cod was immunised with haptenated and non-haptenated protein antigen at two different temperatures and the antibody response monitored over a period of 18 months. Other humoral parameters of immunological importance were also analysed, namely total immunoglobulin concentration, anti-protease and spontaneous haemolytic activity. No specific antibody response was detected but increased activity of non-specific anti-TNP antibodies was observed 10-12 weeks after immunisation, irrespective of the antigen used. This antibody activity was attributed to the adjuvant used (FCA) and did not cross react with other antigens tested. Other parameters were probably not influenced by the immunisation but seasonal fluctuations were indicated. The immunoglobulin level appeared to peak in August-September and the anti-protease activity and the haemolytic activity in October-January.     

Reproductive potential of the Eastern Baltic cod Gadus morhua callarias L. population

Trends in interannual variation in maturation and spawning terms of various age cohorts in the Eastern Baltic cod population in 1997-2009 were studied. Specific features in the age structure of the mature population part that were established by the end of the first decade of the 21st century were clarified. The role of cod age cohorts in the current population reproduction was considered taking into account the data on cod recruitment and fecundity.     

Energetic costs at different levels of feeding in juvenile cod, Gadus morhua L.

The energetic costs associated with feeding by juvenile cod were determined by means of an open-circuit respirometer. Fish acclimated to several temperatures (7, 10, 15 and 18°C) were kept at natural lighting levels, and fed inside their individual respirometers. They consumed a diet compounded from natural foods, at five different ration levels, their oxygen consumption being monitored continually over an 11–16 day period.
After each meal the rate of oxygen consumption increased to above the pre-feeding level, reaching a peak 8–10 h later. With each successive meal the oxygen consumption showed a cumulative increase, reaching a maximum usually after the last meal.
The elevation in metabolic rate associated with feeding was dependent upon ration size, increasing linearly as the food intake increased. The effect was also dependent upon temperature; for fish fed to satiation the total energy cost was equivalent to 11.9, 10.9, 16.4 and 17.1% of the ingested energy at 7, 10, 15 and 18°C respectively. For resting satiated fish the rate of oxygen consumption was close to the maximum rate for active fish.     

Induction of meiotic gynogenesis in Atlantic cod,Gadus morhua (L.)

Gynogenesis is one of several chromosome‐manipulating techniques used in fish. In gynogenesis the male does not contribute to the genetic material of the offspring, and the sperm cells act only as stimulators in order for the egg to start development. This technique has several applications, both in aquaculture and in biological research: gynogenetic fish may be used as a step in the production of all‐female populations, the production of isogenetic‐ and inbred lines, revealing of the sex determination mechanism, construction of genetic maps, and testing of environmental vs genetic control of different traits. The aim of this study was to develop a simple protocol for production of gynogenetic cod (Gadus morhua L.) for further use in aquaculture research. Various milt dilutions and UV‐irradiation doses were tested, in order to inactivate the sperm without destroying its ability to induce egg development. This was followed by pressure treatment of the eggs shortly after ‘fertilization’ to suppress the completion of meiosis II, and thereby restoring diploidy. A dose of 9000 erg mm^?2, followed by a 5‐min pressure treatment (58.6 MPa) 180 min‐degrees after fertilization gave 100% gynogenetic larvae. Histologically, sexual differentiated fish were all females, possibly confirming female homogamety in Atlantic cod. No particular signs of reduced growth, survival or enhanced deformity rates were observed after the fish had reached the juvenile phase. Mortality was, however, high during the egg and larval stages. This protocol has made capable the production of gynogenetic cod juveniles in significant amounts using relatively simple means; the next step will be to elaborate on the technique in order to produce mitotic gynogenetic (double haploid) individuals, which are 100% homozygous.     

Neutrophils and B-cells in Atlantic cod (Gadus morhua L.)

Proportions of leucocytes from head kidney, blood and spleen were identified as B-cells and neutrophils using a polyclonal antibody to cod IgM and a monoclonal antibody which previously has been shown to bind specifically to salmon and trout neutrophils. The cell specific binding of the antibodies was supported by double immunostaining. The morphology of isolated leucocytes was examined on Diff Quick stained slide preparations, and myeloperoxidase positive neutrophils were identified by diaminobenzidine staining. The antibodies clearly identified distinct cell populations. Using flow cytometry, high proportions of neutrophils were observed in peripheral blood leucocytes and high proportions of B-cells were found in head kidney leucocytes when compared to proportions of these cells in Atlantic salmon (Salmo salar L.). The spleen contained the highest proportion of B-cells. Cytoplasmic staining of immunoglobulin positive cells in slide preparations indicated that plasma cells were present, but not strikingly abundant, in head kidney, spleen and peripheral blood. Staining for myeloperoxidase identified, in accordance with the flow cytometry results, a large number of neutrophils, especially in peripheral blood leucocytes. The neutrophil nucleus was not clearly segmented, but appeared more irregular than rounded. The findings of high proportions of neutrophils in peripheral blood suggest that these cells of the innate immune system might have a central role in defence and protection against infections in cod.     

Chitinase activity in the digestive tract of the cod, Gadus morhua (L.)

The study was made to determine if enzymatic degradation of chitin occurs in the digestive tract of the cod, Gadus morhua . The method employed corresponds to the end product measurement of Jeuniaux (1966), using 'native' chitin as the substrate. The following results were obtained.

• (1) 

  Chitinolytic enzyme of high activity is present in enzyme solutions from the stomach contents, gastric mucosa and intestinal contents.
• (2) 

  Lower chitinase activities are found in samples of the intestinal mucosa and the pyloric caeca.
• (3) 

  The optimum pH ranges for the action of the enzymes in the stomach and the intestine differ: 4.5–5.1 and 5.1–6.5, respectively.
• (4) 

  The role of chitin-decomposing bacteria is discussed, based on bacterial numbers and pH conditions in the digestive tract. The existence of two different enzyme systems is indicated.