Egg buoyancy of Baltic cod (Gadus morhua) and its implications for cod stock fluctuations in the Baltic

Successful spawning of Baltic cod (Gadus morhua L.) is restricted to the deep basins of the Baltic Sea and is dependent on abiotic conditions such as salinity and water oxygen content. Due to irregular inflows of saline water from the North Sea, the conditions for spawning fluctuate and consequently so does the cod stock. In May and June 1990 eggs were obtained from spawning cod caught off northern Gotland, Sweden. Our investigation revealed neutral egg buoyancy for Baltic cod at a salinity of 14.4±1.1 ppt, with a slight decrease some days before hatching. Today salinities of this magnitude occur only in the Bornholm basin, whereas in the Gdansk and Gotland basins the eggs sink toward the bottom and are exposed to lethal oxygen conditions.     

Egg buoyancy variability in local populations of Atlantic cod (Gadus morhua)

Previous studies have found strong evidences for Atlantic cod (Gadus morhua) egg retention in fjords, which are caused by the combination of vertical salinity structure, estuarine circulation, and egg specific gravity, supporting small-scaled geographical differentiations of local populations. Here, we assess the variability in egg specific gravity for selected local populations of this species, that is, two fjord-spawning populations and one coastal-spawning population from Northern Norway (66–71°N/10–25°E). Eggs were naturally spawned by raised broodstocks (March to April 2009), and egg specific gravity was measured by a density-gradient column. The phenotype of egg specific gravity was similar among the three local populations. However, the associated variability was greater at the individual level than at the population level. The noted gradual decrease in specific gravity from gastrulation to hatching with an increase just before hatching could be a generic pattern in pelagic marine fish eggs. This study provides needed input to adequately understand and model fish egg dispersal.     

Egg and early larval stages of Baltic cod, Gadus morhua, are robust to high levels of ocean acidification

The accumulation of carbon dioxide in the atmosphere will lower the pH in ocean waters, a process termed ocean acidification (OA). Despite its potentially detrimental effects on calcifying organisms, experimental studies on the possible impacts on fish remain scarce. While adults will most likely remain relatively unaffected by changes in seawater pH, early life-history stages are potentially more sensitive, due to the lack of gills with specialized ion-regulatory mechanisms. We tested the effects of OA on growth and development of embryos and larvae of eastern Baltic cod, the commercially most important fish stock in the Baltic Sea. Cod were reared from newly fertilized eggs to early non-feeding larvae in 5 different experiments looking at a range of response variables to OA, as well as the combined effect of CO₂ and temperature. No effect on hatching, survival, development, and otolith size was found at any stage in the development of Baltic cod. Field data show that in the Bornholm Basin, the main spawning site of eastern Baltic cod, in situ levels of pCO₂ are already at levels of 1,100 μatm with a pH of 7.2, mainly due to high eutrophication supporting microbial activity and permanent stratification with little water exchange. Our data show that the eggs and early larval stages of Baltic cod seem to be robust to even high levels of OA (3,200 μatm), indicating an adaptational response to CO₂.     

Effects of salinity on spermatozoa motility,percentage of fertilized eggs and egg development of Baltic cod (Gadus morhua), and implications for cod stock fluctuations in the Baltic

Variation in cod stock strength in the Baltic Sea is considered to be dependent on abiotic conditions such as salinity and water oxygen content in the spawning areas (the Baltic deep basins). Spawning cod were caught off northern Gotland, Sweden, from April to June in 1989 and 1990. Our investigation revealed a reduction in spermatozoan motility and a subsequent reduction in the percentage of fertilized eggs in salinities of 10 to 12. Normal egg development required a minimum salinity of 11. This coincides with the osmolality of the seminal plasma and egg yolk; i.e., the Baltic cod is adapted to hyperosmotic conditions for spawning and is thus totally dependent on periodical inflows of saline water from the North Sea.     

Foraging behaviour of larval cod (Gadus morhua) influenced by prey density and hunger

Fish larvae meet diverse environmental conditions at sea, and larval growth and chance of survival depend on a flexible response to environmental variability. The present study focuses on the flexibility of the foraging behaviour of larval cod in a series of laboratory experiments on larval search activity, prey selectivity, and hunger in a variable prey environment. Gadus morhua eggs were collected in March 1992 and 1993 from the Kattegat area, Denmark, fertilised and incubated in the laboratory. After hatching, the larvae were transferred to rearing tanks of 172 litres. The behaviour of larvae (6 to 7 mm long) was observed visually, and prey attacks, swimming activity and gut contents were registered across a range of 1 to 120 copepod nauplii l^-1. When prey density decreased, larvae increased their swimming activity, increased their responsiveness to prey (distance of reaction) and decreased their prey size selectivity. Behavioural response was to a large degree determined by the level of hunger, represented by the number of newly ingested prey in the gut. The findings show that cod larvae have a flexible response to changes in feeding conditions and imply that larvae can grow and survive even in the lower range of (mean) prey densities measured at sea.     

Movements of farmed and wild Atlantic cod (Gadus morhua) released in Trinity Bay,Newfoundland

Fishery scientists and managers are investigating the feasibility of enhancing annual recruitment to the northern Atlantic cod (Gadus morhua L.) stock complex off Labrador and northeastern Newfoundland through the release of farmed fish back into the sea. Release of newly matured fish and adults with farm-advanced fecundity would increase the spawning biomass. Enhancement efforts might be measurably successful in major bays that are year-round habitats for cod. To determine if farmed cod would remain and spawn in Trinity Bay, 14 fish with surgically implanted transmitters were released in November 1992. Sonic tracking confirmed that farmed cod released on the western side of Trinity Bay overwintered within the bay, and integrated with wild cod approaching spawning condition in April 1993. Blood plasma antifreeze levels confirmed that these wild cod had overwintered inshore in subzero waters. A spawning aggregation was found in July 1993, providing evidence that northern cod reproduce in Newfoundland bays. These findings suggest that it may be possible to increase the number of cod spawning inshore through the release of farmed fish.     

Experimental evidence of chemokinesis in newly hatched cod larvae (Gadus morhua L.)

The swimming behaviour of laboratory-reared newly hatched cod larvae (Gadus morhua L.) was observed in a control solution of artificial seawater and in seven solutions, each with a different concentration of arginine (10⁹ to 10^-3 M). The behaviour of 20 larvae was analysed in each of the eight solutions; the individual observation time was 1 min. Individual movements were recorded on video and analyzed using a computer-assisted program. The larvae swam in straight lines (a trajectory), rested, moved and started swimming again. For the parameters analyzed, i.e., number of movements, angle between successive trajectories and straightness index, there was no significant difference between the behaviour of the larvae in the different solutions. However, for the larvae in 10^-5, 10^-4 and 10^-3 M arginine solutions, the analyzed parameters, i.e., time active, frequency of trajectories (number of movements exceeding body length), distance swum min^-1, length of individual trajectories and trajectory velocity, were all significantly lower than for the larvae in the control solution of artificial seawater and for larvae in the solutions of 10^-9, 10^-8, 10^-7 and 10^-6 M arginine. The results show that the mean distance swum by cod larvae min^-1 was two to five times longer in artificial seawater without arginine and in the four lower concentrations of arginine than in the three higher concentrations. Scanning micrographs show that newly hatched (pre-feeding) cod larvae possess olfactory organs. It seems reasonalbe to assume that the observed changes in swimming behaviour are mediated by the olfactory sense and are important in the feeding strategy of cod larvae. We suggest that the observed behaviour increases the probability of the larvae localizing patches of prey organisms and remaining in the patch once they have found it. The results show that chemokinesis is a mechanism by which the spatial distribution of fish larvae will be correlated with their prey.     

Activity and metabolism of larval Atlantic cod (Gadus morhua) from Scotian Shelf and Newfoundland source populations

Patterns of activity and metabolism were investigated in larval Atlantic cod (Gadus morhua L.) between December 1991 and July 1992: (1) throughout larval development; (2) between two genetically discrete populations (Scotian Shelf and Newfoundland) and (3) as a function of two different culture temperatures. During the yolk-sac stage (0 to 5 d post-hatch), changes in swimming speed were not related to mass-specific metabolic rates; no portion of the mass-specific oxygen consumption could be explained by changes in activity. In the mixed feeding stage (6 to 14 d posthatch), there was a tendency for oxygen consumption to be related to changes in swimming speed. In the exogenous feeding stage (>14 d post-hatch), oxygen consumption significantly increased with swimming speed. These ontogenetic patterns of activity and metabolism were the same for larvae from the Scotian Shelf and Newfoundland populations. However, over the entire larval life and among ontogenetic stages, the metabolic cost of activity (mass-specific O₂ consumption/swimming speed) of Scotian Shelf larvae was significantly higher than that of Newfoundland larvae. When cod larvae, that had developed at 5°C, were acutely exposed to 10°C, Scotian Shelf larvae had a higher intrinsic cost of activity than Newfoundland larvae, over the entire larval life. During the exogenous feeding stage, the mean metabolic cost of activity for Newfoundland larvae raised at 10°C and tested at 10°C was significantly higher and more variable than that of larvae raised at lower temperatures. However, the metabolic cost of activity of larvae raised and tested at 10°C was not significantly different between source populations. Together these findings suggest that differences in swimming energetics reflect changing energy requirements for activity among ontogenetic stages, and reflect adaptation to regional environments among genetically discrete populations.     

Sub-lethal exposure to ultraviolet radiation reduces prey consumption by Atlantic cod larvae (Gadus morhua)

Levels of ultraviolet-B radiation (UV-B: 280–315 nm) reaching the earth’s surface have increased over the past few decades due to ozone depletion. It is well documented that exposure to UV-B radiation increases mortality in marine fish larvae. However, few studies have examined sub-lethal effects of UV-B radiation such as, for example, the possibility that it affects prey consumption by fish larvae. Atlantic cod larvae were exposed to a sub-lethal level of ultraviolet-B radiation (UV-B: 280–315 nm) for 15 h. After the exposure, rotifers (4/mL) were fed to cod larvae at three different post-exposure intervals (20 min, 3 and 6 h). Trials were replicated three times for each post-exposure interval. The number of rotifers in the gut and the percent of empty guts (number of fish with empty gut/number of fish examined × 100) were analyzed. Results were compared to those of unexposed fish (the control treatment). UV-B exposed cod larvae had consumed significantly fewer rotifers than control fish at all post-exposure intervals. There was no significant difference in the frequency of empty guts between fish in the UV-B treatment versus fish in the control treatment at any of the post-exposure time points (although the difference at 20 min post-exposure was borderline significant). These observations suggest that cod larvae exposed to sub-lethal levels of UV-B have lower net energy gain which may lead to reduced growth rate and possibly poorer survival.     

Wateriness of white muscle: A comparison between cod (Gadus morhua) and jelly cat (Lycichthys denticulatus)

Histological sections of starved cod (Gadus morhua Linnaeus, 1758) and jelly cats (Lycichthys denticulatus Kröyer, 1844) with similar water contents are compared. Starvation causes the white muscle fibres of cod to shrink into irregular shapes inside their connective tissue matrix, while those of jelly cats of similar water content are seen to be rounded. It appears to be less important for jelly cat muscle to be packed tightly with contractile fibres than for the muscle of cod, so the former may be adapted to a permanent state of unusually high hydration.     

Effects of temperature on morphological landmarks critical to growth and survival in larval Atlantic cod (Gadus morhua)

The morphology and function of structures important to energy acquisition were studied from spawning to the stage of transformation of larva to pelagic juvenile in Atlantic cod, Gadus morhua L., from December 1991 to July 1992. Fertilized eggs produced by adult fish from two genetically discrete populations (Newfoundland and Scotian Shelf) were raised under similar conditions in the laboratory at temperatures of 5 and 10°C. Subsamples of larvae were removed from cultures daily for 10 d, and then less frequently, and fixed for light microscopy and scanning electron microscopy. Nine functional morphological landmarks important to feeding, respiration and locomotion were chosen from observation of 280 ind. These landmarks defined 12 major developmental stages, from hatching to the pelagic juvenile stage. One of the feeding landmarks, intestinal stage, varied as a function of age and size and the variance in development was higher at 10°C than at 5°C; Newfoundland larvae developed more complex intestines than did Scotian Shelf larvae. In addition, Newfoundland larvae had significantly higher growth rates than those of Scotian Shelf larvae. Despite the higher growth rates and greater structural complexity of the intestine in Newfoundland larvae, the rate of yolk utilization was not significantly different between Newfoundland and Scotian Shelf larvae. Staging of respiratory landmarks showed that the gill arches were probably used preferentially in feeding while respiration was cutaneous. The gills, operculum and gill rakers developed late in larval life and accompanied the transition from cutaneous to branchial respiration. In the yolk-sac period, development of feeding and respiratory structures may be largely genetically controlled. During exogenous feeding, extrinsic factors also become important, as shown by the size and age-independent variation in intestinal development of larval cod raised at different temperatures.     

How does cod (Gadus morhua) cope with variability in feeding conditions during early larval stages?

Response of mesocosm-reared cod (Gadus morhua L.) larvae to different feeding conditions was investigated in 1988 in two mesocosms: a large basin and a smaller bag enclosure within the basin. The basin was filled with seawater, and a community of naturally occurring plankton developed. Plankton concentrations were monitored, and cod larvae stocked in the enclosures were sampled for determination of growth, survival, and gut content. In the bag, insufficient amounts of energetically favourable prey, as copepod nauplii, led to non-selective ingestion of plankton from a broad range of sizes, including considerable amounts of protozoans (tintinnid and oligotrich ciliates). Growth of larvae from the bag was low, with daily specific growth rates (SGR) less than 2.8% the first 3 wk post-hatch. This was followd by rapid increase of SGR to 21.7%, which coincided with a large increase in availability of copepod nauplii. In the basin, high nauplii concentrations led to SGR of 13.7 to 21.7% from onset of feeding to 16 d post-hatch, respectively. Under such conditions, the larvae were highly selective feeders. At 3 wk post-hatch, survival was 36.7 and 38.3% in the basin and bag enclosure, respectively. To cope with variations in the feeding conditions, the cod larvae were shown to be opportunists when nauplii were scarce, and included plankton from several trophic levels in their diet. When nauplii were abundant, cod larvae realized their high potential for growth. Both opportunism and realization of a high growth potential may enhance survival of the larvae.     

Residency and depth movements of a coastal group of Atlantic cod (Gadus morhua L.)

Atlantic cod stocks are subdivided into geographically and biologically discrete populations; some being locally resident, others migratory. The reasons for the variation in migration pattern and the consequences it may have for population structuring is poorly understood. We studied a group of cod from the coastal waters of the Shetland Isles in the northern North Sea. During the spawning season, electronic tags that record depth and temperature over time were implanted into 133 individuals and the fish released within a few kilometres of where they were captured. Thirty-nine cod have been recaptured up to 608 days later, throughout the year and, in all but two cases, within 15 km of their release site. Geolocation methods based on temperature and bathymetry also suggested that the cod had a limited home range, remaining resident year-round in coastal waters. The cod were deeper during the winter and moved to shallower water in late spring and summer. They showed diel, fortnightly and monthly cycles in depth movement that varied much within and between individuals over the season. Residency and the individual variability in vertical movement may reflect a combination of locally complex depth strata, variable seabed substrate and the wide range of seasonally available prey resources.     

Habitat use by juvenile Atlantic cod (Gadus morhua) in the presence of an actively foraging and non-foraging predator

Experiments were conducted in the autumn and winter of 1992/93 to examine habitat use by juvenile (age 0+) Atlantic cod, Gadus morhua L., before, during and following exposure to a passive or actively foraging predator (age 3+ cod). Experiments presented groups of juvenile cod (n=5 fish/group) with one of two combinations of three substrates; (1) gravel, sand, and a patch of artificial kelp (kelp), or (2) cobble, sand, and kelp. Cobble is known to provide juvenile cod with a refuge from predation. Kelp was used to test the hypothesis that juvenile cod associate with fleshy macroalgae in nature because of the safety it provides from predators. There was little difference in habitat use by juvenile cod before, during or following exposure to a passive predator. Under these conditions, juvenile cod appeared to prefer finer grained mineral substrates and avoided the kelp. The extent of the juvenile response to a passive predator was to avoid the predator's location in the experimental tank. In contrast, juvenile cod showed a significant shift in habitat use when exposed to an actively foraging predator, hiding in cobble or, when cobble was not available, in kelp. Use of both these habitats resulted in a significant reduction in predation risk to the juvenile cod. Our results suggest that: (1) an association with kelp provides safety from predation to juvenile cod, and (2) juvenile cod are capable for assessing the risk a predator represents and adjust their response accordingly.     

Condition of larval cod (Gadus morhua) off southwest Nova Scotia in 1983 in relation to plankton abundance and temperature

In our study of the condition of larval cod (Gadus morhua) collected off southwest Nova Scotia in winter-spring, 1983, we (1) examined relationships between larval condition and ambient environmental conditions, and (2) compared the use of simple morphometric indices of larval condition and of multivariate statistics to obtain information relevant to larval condition. Twelve indices of relative condition were obtained, based upon seven measurements made on each larva, and a principal component (PC) analysis was performed on these condition indices. Most condition indices and the first PC were significantly correlated with numbers of nauplii and of zooplankters per m³ in the water column, which are direct measures of food available to the larvae, but not to less direct measures of environmental quality for the larvae, such as plankton displacement volume or chlorophyll concentration. Conventional indices based upon relations of dry weight or body height at the anus to length were most sensitive to environmental conditions. There was no apparent advantage in performing a multivariate analysis based upon a larger series of measurements.     

Feeding opportunities of larval and juvenile cod (Gadus morhua) in a Greenlandic fjord: temporal and spatial linkages between cod and their preferred prey

Feeding of fish depends on a spatial and temporal match with prey, and since larval and juvenile feeding can be highly selective, their preferences for given prey sizes and taxa should be considered when quantifying the actual availability of potential prey. We investigated the diet and prey preferences of the early-life stages of Atlantic cod (Gadus morhua) to quantify the availability of prey during a spring-summer season in a West Greenlandic fjord. We hypothesized that abundances of larval and juvenile cod at size were synchronized to optimal availability of preferred prey in space and time. The present analysis is based on nine cruises each covering 5 stations visited between 24 May and 5 August 2010 comparing zooplankton abundance, cod gut content and distribution patterns. Cod 4–25 mm in length preferred prey of about 5 % of their own length. During ontogeny, their preferences changed from calanoid nauplii towards Pseudocalanus spp. and Calanus spp. copepodites. The larvae/juvenile had an exceptionally high dietary contribution from cladocerans, which were highly preferred by cod larger than 9 mm, while the abundant Metridia longa and the non-calanoid copepods contributed less. These findings stress the importance of focusing on abundance of preferred prey when assessing the actual prey availability to young fish. We found a spatio-temporal overlap between cod and their preferred prey, and observations suggest that advection of both zooplankton and cod contributed to this overlap. Hence, the larval feeding opportunities might be sensitive to climate-related changes affecting the circulation patterns in this fjord.     

Exposure to variable spatial information in the early rearing environment generates asymmetries in social interactions in cod (Gadus morhua)

Many re-introduction programs used for conservation of populations and species threatened with extinction advocate the use of enriched rearing environments to train animals how to behave appropriately in the wild. Curiously, most of the current fish re-stocking programs have paid little attention to lessons previously learned in bird and mammal re-introductions. Many rehabilitation programs that use releases of hatchery fish observe higher mortality in released fish compared to wild, with most mortality arising shortly after release. One explanation for this mortality is based purely on selection processes; many hatchery fish normally selected out of the population thrive in the predator free, food-rich hatcheries. Alternatively, mortalities may be high because hatchery nursery environments fail to shape fish behaviour appropriately. Here, we empirically address the effect of enrichment in the early rearing environment in coastal cod (Gadus morhua). We find asymmetries in aggressive behaviour when fish reared in plain or enriched environments are allowed to interact. Furthermore, cod reared in standard, impoverished, hatchery environments spend less time in shelter, are more active, and show weaker anti-predator responses than fish reared with access to heterogeneous spatial cues. These results suggest that the constant, plain environments of fish farms may generate behavioural deficits that could reasonably be expected to be associated with lower survival in fish released into the wild.     

Maternal allocation of lipid classes and fatty acids with seasonal egg production in Atlantic cod (Gadus morhua) of wild origin

A suite of characteristics is often used to assess egg quality as these properties potentially play important roles in progeny survival and growth. Our objective was to assess egg characteristics including lipid biocomposition of an iteroparous, batch-spawning teleost of wild origin. Maternal allocation to egg number was generally dome-shaped (5 of 8 females) and egg size declined over the breeding season for eight breeding pairs of wild Atlantic cod (Gadus morhua) (n = 43 batches). Egg lipid composition ranged considerably among females and between egg batches within females (e.g., phospholipids 40–86 %; polar 47–87 % and neutral lipids 15–52 % of total lipids; polyunsaturated fatty acids 16–50 % of total fatty acids). Principal component analyses revealed significant inter-relationships among maternal traits, batch sequence and fecundity, and egg size and composition. Seasonal trends with regard to lipid deposition were variable; three females showed consistent declines in lipid parameters (μg egg⁻¹) with both batch number and egg diameter, one female showed consistent increase and the four remaining females showed no trend. The three females that exhibited seasonal declines in egg lipid content were characterized as having high fertilization success (>75 %). Our findings highlight the variability in lipid allocation to eggs of batch spawners of wild origin and characterize the composition of endogenous reserves available during embryogenesis and yolk sac larval stages.     

Free amino acids as energy substrate in developing eggs and larvae of the cod Gadus morhua

The content of free amino acids (FAA) in the cod (Gadus morhua L.) egg is about 200 nmol at spawning, decreasing by about 100 nmol/egg during the egg stage and about 75 nmol/larva during the yolksac larval stage. Together, alanine, leucine, serine, isoleucine, lysine, and valine account for about 75% of the decrease. Ammonium accumulates gradually during the egg stage and is quickly excreted after hatching. The body protein content is maintained during the egg and yolksac larval stages. The measured oxygen uptake of the cod embryo during the egg and yolksac larval stages accounts for about 85% of the oxygen necessary to catabolize the FAA disappearing during this period. Ammonia excretion of the cod embryo, as taken from literature data, is similar to the expected ammonia production from catabolism of the FAA. Our data suggest that FAA are a major substrate for aerobic energy production in cod eggs and yolksac larvae. The implication of this finding for the production of a favourable first-feed for cod and other cultivated marine fish larvae, and for the selection of high quality eggs of marine fishes, is stressed.