Genetic variation and population structure of cod, Gadus morhua L., in some fjords in northern Norway

Biochemical genetic polymorphism in cod from several fjords in Troms, northern Norway was analysed. Gene frequencies at several polymorphic loci ( Hb, Pgi, Ldh, Pgm, Gpd and Idh ) are given. Significant variation was found both within and between fjords, and even between samples from the same locality sampled in different years, indicating a mosaic structure of the cod population in the area studied. The results of the haemoglobin variation are compared to similar results obtained more than 20 years ago in the same and adjacent areas. The biochemical genetic variation is discussed in relation to stability of gene frequencies, isolation mechanisms and migration patterns.     

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.     

The parasite fauna of cod (Gadus morhua L.) in Balsfjord,North Norway

Summary Samples of cod from Balsfjord, a subarctic fjord in North Norway, were collected and the fauna of macro-parasites was classified zoogeographically. The results revealed an impoverished and to a large extent boreal parasite fauna compared with that of cod from the Barents Sea. Because of the last glaciation Balsfjord is a relatively young ecosystem, and the silled entrance surrounded by an archipelago also makes the fjord a rather isolated system. This together with its small size may be important factors in explaining the character of its parasite fauna. The colonization by boreal parasites of both the Barents Sea and Balsfjord probably reflects the direction of the main currents.     

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.     

Oogenesis in cod, Gadus morhua L., studied by light and electron microscopy

The development of the oocytes in the cod, Gadus morhua L., is described by light and transmission electron microscopy. The oocyte volume increases about 700 times during the stages preceding hydration. The size of the nucleoli increases 35-fold, the cortical alveoli increase 6-fold and the yolk granules increase 70-fold as the oocyte grows. The maximal number of yolk granules is nearly 100000 per oocyte; this number is reduced to less than the half prior to hydration. A significant reduction occurs also in the number of nucleoli at that time. The number of cortical alveoli increases steadily towards hydration. Yolk is deposited in the oocyte as crystalline granules. The lattice is broken down at hydration, leaving the egg transparent. Follicle cells go through a primordial stage and later change to a squamous and to a cuboids shape. The presence of lipid droplets in their interior and the virtual lack of interfollicular spaces are characteristic. The chorion grows to a tripartite structure: an outer thin porous layer, an intermediate homogenous layer and an inner thick helicoidally layer. A mucous substance covers the porous layer. The occurrence of the so-called lamellae in the helicoidal layer is considered a function of the orientation of its micro fibrils and the plane of sectioning, i.e., not caused by alternating chemical arrangements.     

Gene flow and lack of population differentiation in Atlantic cod, Gadus morhua L., from Iceland, and comparison of cod from Norway and Newfoundland

Restriction analysis of mitochondrial DNA was used to study genetic variation and geographic population structure of Atlantic cod from localities around Iceland. Gene phylogenies were constructed and geographic locations superimposed on these. The variation was not localized. Estimated gene flow was large. Thus, Atlantic cod in Iceland belong to a single genetic population. Analyses of published sequence variation of cod from Norway and Newfoundland showed that the extensive continuity of intraspecific phylogeny of cod in Iceland, which is very similar to other marine organisms with a similar life history, extends from Norway to Newfoundland and possibly to larger areas of the Atlantic.     

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.     

Seasonal variations in biochemical composition and energy content of liver, gonad and muscle of mature and immature cod, Gadus morhua (L.) from Balsfjorden, northern Norway

Seasonal energy changes in mature and immature cod of the same size were studied. For most of the year, the variations in energy content in the tissues of both mature and immature cod are similar. In the period from October to January, the fish accumulate energy. From January until spawning in early April there is a fall in the total energy content. The growth of gonads from January until spawning can explain about one-third of the energy decrease in liver and muscle.     

Factors affecting gastric evacuation in cod, Gadus morhua L., fed single-meals of natural prey

Functional relationships were derived in order to describe the effects of fish weight (262–2066 g), temperature (1–14° C), prey size and ration size (0.4–8.4% body weight) on gastric evacuation in cod fed four natural prey types. Interindividual (unexplained) differences among voluntary fed cod were relatively large. Power functions were appropriate for describing the effects of fish weight and ration size on the half-time of evacuation. Effects of meal and/or prey size appeared to counterbalance the effects of cod size. It is suggested that when cod of different sizes are fed fixed proportions of their body weight evacuation time is constant and independent of body size. There was found to be an exponential relationship between temperature and gastric evacuation time but, observations made at similar temperatures at different times of the year suggested that seasonal effects were of minor importance.     

Robertsonian polymorphism in plaice, Pleuronectes platessa L., and cod, Gadus morhua L., (Pisces Pleuronectiformes and Gadiformes)

Karyotypic analysis of plaice, Pleuronectes platessa L., and cod, Gadus morhua L., revealed that the chromosome numbers in both species vary but that chromosome arm numbers (NF) stay constant. The C-, Q- and R-banding patterns also confirmed that the population of plaice studied shows Robertsonian polymorphism. The tendency of reduction in chromosome number in fishes suggests that Robertsonian fusions play a role in karyotype evolution in fishes.     

Early development of the digestive tract of cod larvae, Gadus morhua L., during start-feeding and starvation

Cod larvae, Gadus morhua L., were reared in the laboratory and released to a large marine enclosure 4 to 5 days after hatching (6–8° C). The development of the digestive system was studied until day 24 after hatching. Morphological investigations of the jaw apparatus and the digestive tract showed that the larvae are able to absorb ingested food well before exhaustion of the yolk sac. The foregut, and especially the midgut, were particularly active in lipid absorption, and the hindgut was characterized by pinocytotic activity. Duhng the first days of feeding, no distinct prey organisms were observed in the gut, and signs of food absorption in the epithelial cells of the gut were sparse.A distinct red fluorescence, restricted to the hindgut, was observed from day 11 to day 19. On the basis of changes in absorptive pattern in the gut we suggest that changes in digestive and absorptive abilities, as well as in nutritional needs, take place around days 15–17 after hatching.
In starved larvae, signs of degeneration of the gut tissue were first visible in the foregut. By day 9 after hatching, microvilli was degenerated to such an extent that the ability to absorb food must have been severely restricted. If larvae are starved longer than this, they will probably not survive.     

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.     

Mitochondrial cytochrome b DNA sequence variation of Atlantic cod Gadus morhua, from Norway

We studied sequence variation of a fragment of the mitochondrial cytochrome b gene using polymerase chain reaction (PCR) and direct sequencing among 85 Norwegian Atlantic cod Gadus morhua, from nine sampling localities representing three overall areas: arctic, coastal and middle. In the analysis we include an additional 15 cod studied by Carr & Marshall (1991a). Nine base changes were found among the 100 sequences defining 11 haplotypes which differ from each other by one to five mutations. Two sites have been hit twice. All but one mutation are at third position silent sites. The variation passes several neutral-theory tests and is thus suitable as marker for studying population differentiation. Coefficients of coancestry or intraclass correlation coefficient are negative both at the level of individuals within localities and localities within areas. This implies greater differences among individuals within populations than between populations. Intralocality nucleotide diversity is high and masks interlocality nucleotide divergence such that the net interlocality nucleotide divergence is nil. Similarly the net interarea nucleotide divergence is nil.     

The hardening process of the egg chorion of the cod, Gadus morhua L., and lumpsucker, Cyclopterus lumpus L.

When transferred to sea water the chorion of cod and lumpsucker eggs harden, reaching a resistance of about 150 and 2000 g respectively. In sea water this hardening process is independent of fertilization. Studies of eggs kept in artificial sea water with various ionic compositions indicated, in the lumpsucker eggs, that a cortical reaction seems to be a prerequisite for the hardening process. Calcium is necessary for the reaction both in unfertilized and inseminated eggs, whereas hardening takes place in the absence of potassium, magnesium or sulphate. Addition of barium or strontium as a substitute for calcium only caused hardening in the presence of activating spermatozoa. Activating spermatozoa were also necessary for hardening of lumpsucker eggs kept in ovarian fluid, but such hardening only occurred if calcium and/or magnesium were added. The hardening of lumpsucker eggs was associated with profound changes in the thickness and surface appearance of the chorion.     

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.