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1.
Temperature and body size are widely agreed to be the primary factors influencing vital rates (e.g., growth, mortality) in marine fishes. We created a biophysical individual‐based model which included the effects of body size and temperature on development, growth and mortality rates of eggs, larvae and juveniles of Atlantic cod (Gadus morhua L.) in the North Sea. Temperature‐dependent mortality rates in our model were based on the consumption rate of predators of cod early‐life stages. The model predicted 35%, 53% and 12% of the total mortality to occur during the egg, larval and juvenile stages, respectively. A comparison of modeled and observed body size suggested that the growth of survivors through their first year of life is high and close to the growth rates in ad libitum feeding laboratory experiments. Furthermore, our model indicates that experiencing warmer temperatures during early life only benefits young cod (or theoretically any organism) if a high ratio exists between the temperature coefficients for the rate of growth and the rate of mortality. During the egg stage of cod, any benefit of developing more rapidly at warmer temperatures is largely counteracted by temperature‐dependent increases in predation pressure. In contrast, juvenile (age‐0) cod experiences a higher cumulative mortality at warmer temperatures in the North Sea. Thus, our study adds a new aspect to the ‘growth–survival’ hypothesis: faster growth is not always profitable for early‐life stages particularly if it is caused by warmer temperatures.  相似文献   

2.
How climatic variability and anthropogenic pressures interact to influence recruitment is a key factor in achieving sustainable resource management. However, the combined effects of these pressures can make it difficult to detect non‐stationary interactions or shifts in the relationships with recruitment. Here we examine the links between climate and Irish Sea cod recruitment during a period of declining spawning stock biomass (SSB). Specifically, we test for a shift in the relationship between recruitment, SSB and climate by comparing an additive (generalized additive model, GAM) and non‐additive threshold model (TGAM). The relationship between recruitment success, SSB and the climatic driver, sea surface temperature, was best described by the TGAM, with a threshold identified between recruitment and SSB at approximately 7900 t. The analysis suggests a threshold shift in the relationship between recruitment and SSB in Irish Sea cod, with cod recruitment being more sensitive to climatic variability during the recent low SSB regime.  相似文献   

3.
We estimated recent growth of Atlantic cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) larvae collected on the southern flank of Georges Bank in May 1992–94 from the ratio of RNA to DNA (R/D) and water temperature. Growth of both species increased with water temperature to about 7°C and then decreased. The highest growth rates were observed in May 1993 at water temperatures around 7°C. These data confirm an earlier observation of comparable temperature optima for growth of Atlantic cod and haddock larvae in the north‐west Atlantic. Comparisons of field growth rates and temperature optima with data for larvae cultured at high temperatures and prey densities in the laboratory suggest that growth may have been food‐limited at higher temperatures on Georges Bank. Given that 7°C is the long‐term mean water temperature on the southern flank in May and that climate models predict a possible 2–4°C rise in water temperatures for the western North Atlantic, our findings point to a possible adverse effect of global warming on Atlantic cod and haddock.  相似文献   

4.
Atlantic cod, Gadus morhua, harvested in US waters are currently managed as a Gulf of Maine stock and as a stock comprising Georges Bank and southern New England populations. Over the past two and a half decades, success of age‐1 recruitment to the Gulf of Maine stock has varied by more than an order of magnitude. To investigate the hypothesis that this variation is related to variation in the transport of larval cod to nursery areas, we carried out model simulations of the movement of planktonic eggs and larvae spawned within the western Gulf of Maine during spring spawning events of 1995–2005. Results indicate that the retention of spring‐spawned cod, and their transport to areas suitable for early stage juvenile development, is strongly dependent on local wind conditions. Larval cod retention is favored during times of downwelling‐favorable winds and is least likely during times of upwelling‐favorable winds, during which buoyant eggs and early stage larvae tend to be advected offshore to the Western Maine Coastal Current and subsequently carried out of the Gulf of Maine. Model results also indicate that diel vertical migration of later stage larvae enhances the likelihood of retention within the western Gulf of Maine. Consistent with model results is a strong correlation between age‐1 recruitment success to the Gulf of Maine cod stock and the mean northward wind velocity measured in Massachusetts Bay during May. Based on these findings, we propose a wind index for strong recruitment success of age‐1 cod to the Gulf of Maine stock.  相似文献   

5.
The demersal settlement of pelagic juvenile fish has been considered a critical period in which the final adjustment is made to the size of a year class. Distribution patterns of pelagic and recently settled juvenile cod (Gadus morhua) were examined from nine surveys on Georges Bank during the summer over 5 years, 1984–1989, to relate juvenile survival to the sedimentary environment. Pelagic juveniles were widespread across Georges Bank in June, and by mid‐July they occurred on all bottom types from sand to gravel on eastern Georges Bank. However, by late July‐early August they were mostly abundant on the northeastern edge gravel deposit, which with its complex relief, provides abundant prey and refuge from predators. A bank‐wide estimate of pelagic juvenile abundance in 1986 and 1987 was used to assess mortality of the recently settled juveniles and to evaluate the relative importance of survivors from the northeastern edge gravel area to recruitment of the Georges Bank population. Settlement mortality rates over 1–2 months on the northeastern gravel area ranged from 3 to 8% day?1, which compared reasonably with other studies. The seasonal abundance of the pelagic juveniles was almost an order of magnitude higher in 1987 than 1986; however, recruitment at age 1 was similar, indicating that a high mortality of the demersal juveniles occurred in 1987. The limited northeastern gravel area on Georges Bank may represent a survival bottleneck depending on the variability in the distribution and abundance of juvenile cod settlement in relation to that of their predators.  相似文献   

6.
Body movements of cod (Gadus morhua L.) embryos reared from fertilization to hatch at 5.4°C were observed at various stages of development and at six experimental temperatures ranging from 0–10°C. Frequency of cod embryo body movements increased from zero at 42 degree‐days post fertilization to maximal at 73–82 degree‐days (1 or 2 days prior to hatch). Embryos were most active at 2°C (mean of 5.5 movements per 10 min), with activity declining to less than 1/10 min at 8–10°C. Lengths of hatched cod larvae reared at a series of constant temperatures (from 4–10°C) from fertilization to hatch were greater at lower incubation temperatures. Incubation temperatures of 2–4°C were found to be optimal for incubation of cod eggs.  相似文献   

7.
The role of behavior, especially vertical migration, is recognized as a critical component of realistic models of larval fish dispersion. Unfortunately, our understanding of these behaviors lags well behind our ability to construct three-dimensional flow-field models. Previous field studies of vertical behavior of larval Pacific cod ( Gadus macrocephalus ) were limited to small, preflexion stages (≤11 mm SL) in a narrow range of thermal conditions. To develop a more complete picture of larval behavior, we examined the effects of ontogeny, temperature, and light on vertical responses of larval Pacific cod in experimental columns. While eggs were strictly demersal, yolk-sac larvae displayed a strong surface orientation as early as 1 day post hatch (∼ 5 mm SL). Consistent with field observations, small preflexion larvae (<10 mm SL) showed no response to varying light levels. However, there was a direct effect of temperature on larval behavior: Pacific cod larvae exhibited a stronger surface orientation at 4°C than at 8°C. The behavior of larger, postflexion larvae (>15 mm SL) in experimental columns was consistent with a diel vertical migration and independent of water temperature: fish were more widely distributed in the column, and median positions were consistently deeper at higher light levels. These laboratory observations are combined with observations from discrete-depth (MOCNESS) sampling in the Gulf of Alaska to characterize the vertical distribution of larval Pacific cod and contrast ontogenetic patterns with walleye pollock ( Theragra chalcogramma ). The vertical movements of larval Pacific cod described here will be applied in the development of dispersal projections from Gulf of Alaska spawning grounds.  相似文献   

8.
Two approaches were used to qualify observed variability in Greenland cod (Gadus morhua) recruitment. In the first analysis, we used the linear trend of the Greenland cod recruitment time series and climatic variables, such as air temperatures from the Denmark Strait and wind conditions off East Greenland and Southwest Greenland, to explain the interannual variation in cod recruitment off Greenland. The model resulting from this ‘trend/environmental approach’, explained 79% of the interannual variation in cod recruitment off Greenland. In the second, analytical approach, the ‘regime approach’, multiple linear regression models were used, with the input data being the time series of cod recruitment and spawning stock biomass (SSB) from Iceland and Greenland, sea surface and air temperatures around Greenland, and zonal wind components between Iceland and Greenland. Model results indicated that, during the decades between 1950 and 1990, there were three different cause–effect regimes which significantly influenced the variability of cod recruitment. The three regimes included: (a) the 1950s and 1960s, a regime with favorable sea surface temperatures and a self‐sustaining cod stock off Greenland with high SSB that produced a series of above‐average, strong year classes; (b) the 1970s and 1980s, a regime of declining SSB and recruitment, with recruitment dependent on advection from Iceland; and (c) the 1990s, when the advective potential for recruitment from the Icelandic cod stock was the only available source for replenishment of the Greenland cod stocks, because cod recruitment in Greenland waters was negligible. The three models explained 76–77% of the observed interannual variation in cod recruitment off Greenland. Both approaches suggested that advective factors were the dominant influences for cod recruitment in the ‘Iceland–Greenland System’.  相似文献   

9.
We carried out an experiment to determine how rapidly the early incubation temperature of Atlantic cod eggs can be increased without affecting normal embryonic development and hatching. Atlantic cod eggs were incubated at a constant low temperature (4.5 ± 0.5°C; T5 – control) and four temperature increment treatments where the temperatures were increased stepwise from 4.5°C at zygote stage to 9.5 ± 05°C (T1‐8 h, T2‐32 h, T3‐64 h and T4‐96 h). Embryonic cell symmetry, embryonic mortality, hatching success and larval skeletal abnormalities, length and yolk sac volume were recorded. Larval samples were also taken at hatch for histological analysis. Except for higher egg mortality and lower hatching success in the T1, the differences among experimental groups were minor. Cell asymmetries and embryo mortalities were not significantly different between the control and T2–T4 treatment groups. Control larvae were significantly longer and had smaller yolk reserves at hatch than T1–T4 larvae and larvae from T2 had the largest yolk reserves. Tissue and organ histology of hatched larvae were similar. Considering embryonic cleavage pattern, hatching success and larval morphology and histology, a gradual increment of temperature in 32 h seems to be the better choice for future developmental programming studies in Atlantic cod.  相似文献   

10.
Commercial intensive fry production of Atlantic cod will be dependent on production of viable eggs independent of season. This can only be done by manipulation of maturation by photoperiod, but little is known about potential effects on egg characteristics and larval viability. In two cod broodstocks, maturation was successfully advanced or delayed 6 months compared with normal spawning season (March–April) by manipulation of photoperiod. The advanced broodstock spawned both in spring and autumn the same year. In two of the spawning tanks during autumn, ambient temperature was reduced after reaching 13.7°C during the first half of the spawning period. Egg quality and viability were monitored, and several egg batches were incubated, hatched and start‐fed for examination of growth and survival. Temperatures above 9.6°C resulted in significant reductions in fertilization and normal egg development. Concurrently, fractions of dead and unfertilized eggs increased with elevated temperature. Actual relative fecundity was not affected by temperature. Egg characteristics improved when temperature was controlled and lowered below 9.6°C. Occurrence of irregular spawners suggests that handling of broodstock fish should be avoided during maturation and spawning. Cod larvae originating from eggs of the advanced or delayed broodstocks were successfully reared beyond metamorphosis. Survival was 9.0–46.6% and 29.3% in green and clear water respectively. Survival correlated with both initial and average feeding conditions, but growth rate did not correlate with either of survival and feeding conditions. Specific growth rates (8.3–13.6% day−1) is comparable with other intensive rearing trials with cod, but were lower than reported from nature‐like systems.  相似文献   

11.
This study investigated the effects of photoperiod and temperature on plasma melatonin secretion in Atlantic cod (Gadus morhua L.). Initial work confirmed the presence of a diel profile of melatonin synthesis, with elevated levels during the dark phase. Unusually for fish, the peak in plasma melatonin occurred towards the end of the dark phase, which is indicative of a type `A' melatonin profile. When exposed to 60 hours of continuous darkness a clear endogenous rhythm of melatonin synthesis was observed, which continued for 4 cycles with a periodicity which, approximated to 24 h. When acclimated to varying temperatures (4, 8, 12 or 16 °C) no variation in melatonin production was seen, however, body size appeared to be an important influence, with the smallest fish exhibiting significantly higher levels of dark phase melatonin. Finally, the application of additional night-time illumination to cod maintained in sea cages i.e. without blackout, did not significantly reduce dark phase plasma melatonin, suggesting that cod are less sensitive to photoperiod manipulation in cages than salmonids.  相似文献   

12.
In vitro cell culture methods are crucial for the isolation, purification and mass propagation of intracellular pathogens of aquatic organisms. Cell culture infection models can yield insights into infection mechanisms, aid in developing methods for disease mitigation and prevention, and inform commercial‐scale cultivation approaches. This study details the establishment of a larval cell line (GML‐5) from the Atlantic cod (Gadus morhua) and its use in the study of microsporidia. GML‐5 has survived over 100 passages in 8 years of culture. The line remains active and viable between 8 and 21°C in Leibovitz‐15 (L‐15) media with 10% foetal bovine serum and exhibits a myofibroblast phenotype as indicated by immuno‐positive results for vimentin, α‐smooth muscle actin, collagen I and S‐100 proteins, while being desmin‐negative. GML‐5 supports the infection and development of two microsporidian parasites, an opportunistic generalist (Anncaliia algerae) and cod‐specific Loma morhua. Using GML‐5, spore germination and proliferation of L. morhua was found to require exposure to basic pH and cool incubation temperatures (8°C), in contrast to A. algerae, which required no cultural modifications. Loma morhua‐associated xenoma‐like structures were observed 2 weeks postexposure. This in vitro infection model may serve as a valuable tool for cod parasitology and aquaculture research.  相似文献   

13.
We use a simple model of temperature-dependent egg development and mortality to develop several hypotheses concerning the effect of temperature on the occurrence of eggs of Atlantic cod, Gadus morhua , in Placentia Bay, Newfoundland. Predictions from this exploratory model were tested through a series of synoptic ichthyoplankton surveys throughout spawning and postspawning periods during 1997 and 1998. Although several egg mortality relationships were explored through the simulation, a constant mortality rate best represented the pattern observed in the two years of data. Peaks in late stage egg densities occurred in August of both 1997 and 1998 and were apparently decoupled from egg production peaks in April. We observed a decrease in mortality and the distance dispersed during egg development with increases in water temperature. We suggest that the effects of predation are small relative to the advective effects within this system, and that the interaction between advection and temperature-dependent vital rates of eggs may have dramatic consequences for coastal retention of propagules produced by inshore spawning events.  相似文献   

14.
The robustness of a previously described environmentally mediated stock–recruitment relationship for Pacific cod in Hecate Strait, BC, Canada was tested with 10 yr of additional data. The original analysis tested several alternative hypotheses and concluded that water transport through Hecate Strait, as indicated by sea level height, coupled with cod spawning biomass formed the best model. The present analysis indicates the relationship held through the 1990s. The implications of variation in sea level on stock production were investigated with a delay‐difference stock production model that included an environmentally mediated stock–recruitment relationship. The model predicted that the maximum fishery yield would vary between 1750 and 3670 t yr?1 over the observed range of sea level height, and the estimated unsustainable fishing mortality during periods of low productivity would be only 0.5 times that in periods of high productivity.  相似文献   

15.
Coupled three-dimensional (3-D) physical oceanographic modelling and field sampling programmes were carried out in May 1988 and August 1991 to investigate the potential drift of larval cod ( Gadus morhua L.) in the Bornholm Basin of the Baltic Sea. The goals were to predict the transport of cod larvae, thus aiding the identification of physical processes influencing larval retention/dispersal. Numerical simulations were performed using a 3-D eddy-resolving baroclinic model based on the Bryan–Cox–Semtner code adapted for the Baltic Sea. Within the Bornholm Basin, the model was initialized with ground truth data of physical parameters obtained on the research cruises, and all simulations were forced with actual wind data. Outside the basin, generalized hydrographic features of the Baltic Sea were utilized by incorporation of simulated hydrographic fields from previous model runs typical for the time periods considered. Larval drift was simulated either by incorporation of passive drifters, or as the initial horizontal distribution of larvae implemented into the model. Drift model simulations of larval transport agreed relatively well with field observations. The influence of variations in the vertical distribution on a smaller scale, i.e. vertical deviations of ± 6 m from the observed mean centre of mass, on the drift was examined, revealing no significant differences in the drift of larvae depending on their vertical distribution. The different wind forcing during the investigated time periods was linked to a retention situation in May 1988 and to a dispersal situation in August 1991. Finally, observed spatial distribution patterns of 1-group cod based on Baltic Young Fish Surveys (BYFS) were compared with their predicted transport in the larval phase and examined with respect to recruitment.  相似文献   

16.
Atlantic cod (Gadus morhua) is one of the commercially most important fish species in the North Atlantic and plays a central role in several ecosystems. Fishing pressure has been heavy over a prolonged period and the recent decades have shown dramatic decline in abundance of many stocks. The Arcto‐Norwegian (or North‐east Arctic) cod stock in the Barents Sea is now the largest stock of Atlantic cod. Recruitment to this stock has varied extensively during the last 60 yr. There is evidence for fluctuations in climate, particularly sea temperature, being a main cause for this variability, higher temperatures being favourable for survival throughout the critical early life stages. Our studies of time series present compelling evidence for a strengthening of the climate–cod recruitment link during the last decades. We suggest this is an effect of the age and length composition of the spawning stock having changed distinctly. The age of the average spawner has decreased by more than 3 yr from between 10 and 11 in the late 1940s to 7–8 in the 1990s, average length from just above 90 cm to around 80 cm. The number of age classes contributing to the spawning stock has also decreased, while the number of length groups present increased slightly. Significant decrease in age of spawners has frequently been described for other heavily fished stocks worldwide. We therefore find it likely that the proposed mechanism of increased influence of climate on recruitment through changes in the spawning stock age and size composition is of a general nature and might be found in other systems.  相似文献   

17.
Sea temperature has earlier been shown to have a large influence on the recruitment of Arcto-Norwegian cod, Gadus morhua. We here hypothesize that this linkage is partly due to the direct effect of temperature on larval and juvenile growth. Secondly, temperature acts as a proxy for both biotic and abiotic factors influencing recruitment. Indices of abundance of early juvenile cod (2–3 months old), 0-group cod (4–5 months old) and 3-year-old cod are analysed in more detail against the environmental temperature, wind stress components, wind-induced turbulent energy and the spawning stock biomass. To deal with autocorrelation, non-stationar-time and nun-normality, which complicate a statistical time series analysis, randomization and Box-Jenkins methods are applied. In addition to the important effect of high sea temperature during the early life stage in forming strong year classes, the results show that the spawning stock biomass is nearly as important. Also, alongshore southerly wind stress anomalies during the period of pelagic drift (from April through summer) and offshore wind stress anomalies during egg and early larval stages (in April) act favourably on recruitment. The beneficial effect of southerly wind anomalies could he linked partly to high temperature, but the flux of zooplankton-rich water from the Norwegian Sea into the feeding areas of the Barents Sea may also be increased. The favourable influence of offshore winds in April is less predominant and causal links are also less clear; possible explanations for this might be increased offshore spreading of eggs and early larvae, resulting in reduced risk of predation, and increased compensation inflow of intermediate Norwegian Sea water which, in this restricted period of time, has a high concentration of spawning copepods suitable as prey for the developing cod larvae.  相似文献   

18.
Field-derived growth rates (RNA-DNA based) of cod (Gadus morhua) larvae collected on the southern flank of Georges Bank were higher on average in May 1993 than May 1994, despite the apparent higher abundance of potential prey in 1994. A biophysical modeling study is presented here in which factors are examined that may have led to the difference in population mean growth. A one-dimensional physical model, forced by winds and tides, was used to simulate the vertical structure (of currents, temperature field, and turbulent kinetic energy dissipation rate) following a column of water in a Lagrangian sense at a site on the southern flank of Georges Bank over 5-day periods in late May of 1993 and 1994. The biophysical model and observed zooplankton abundance allowed us to explore the vertical structure and temporal (hourly) evolution of feeding and growth for cod larvae in relation to environmental conditions. Our trophodynamic model is improved over previous versions and now includes the effect of light on larval feeding response, as well as the effect of temperature on larval metabolic costs, ingestion, and digestion. Larval prey profiles, comprising four copepod species, were used from a time series of 1/4-m2 MOCNESS tows to define the prey field. Data from a collateral time-series of larval gut contents (1-m2 MOCNESS tows) was used to define maximum ingestion (satiation level) and prey selection. Model outputs provide depth-dependent estimates of growth, prey biomass ingested, larval length, and larval weight. Water-column growth-rate profiles were made for four size classes of larvae (5, 6, 7 and 9 mm) under the environmental conditions observed in May 1993 and 1994. A weighted-mean growth rate based on the mean vertical distribution of larvae was estimated for each size class. In all cases, when using all available potential prey, the model-derived 1994 growth rates were higher (by 3–6% day−1) than those for 1993. However, simulations in which 7-mm larvae followed the field-derived weighted mean depth over the sampling period, and were limited to their preferred Pseudocalanus prey, resulted in average growth of 12.2% day−1 for 1993 and 9.7% day−1 for 1994. These compared closely to the field growth means of 11.3% day−1 in 1993 and 9.8% day−1 in 1994. Thus, the lower observed growth in May 1994 may have resulted from depth-dependent food limitation and prey-selectivity coupled with the greater metabolic costs induced by the higher temperature that year.  相似文献   

19.
The objective of this paper was to compare the growth and gut morphology of juvenile diploid and triploid Atlantic cod (Gadus morhua) reared under similar conditions. Individually tagged 36‐week‐old diploid (mean weight 49.3 ± 13.8 g) and triploid (mean weight 43.6 ± 11.2) juvenile cod were measured at intervals during a 29‐week growth trial. Data for weight, length, condition factor (K), hepato‐somatic index (HSI), gonado‐somatic index (GSI), Relative Gut Length (RGL) and pyloric caeca number were collected and results were analysed in relation to ploidy status, gender and family contribution. At the end of the experiment, only one family (M2xF3) had many representatives with a relatively even distribution of sexes and ploidies. Diploid females were significantly heavier and had higher K than triploid females in the M2xF3 family (body weight 371.2 ± 120.2 vs. 298.4 ± 100.7 g; K 1.1 ± 0.1 vs. 0.93 ± 0.1), but no differences were found between diploid and triploid males. In the other families (pooled data), no differences in body weight were found between the ploidy groups. In general, triploids had a shorter intestine (RGL) and fewer pyloric caeca than their diploid siblings regardless of gender suggesting possible impairments in nutrient utilization and growth.  相似文献   

20.
Escapes of cod (Gadus morhua) from sea cages represent an economic problem for farmers and a potential environmental problem. We estimate that 0–6% of cod held in sea‐cage farms in Norway were reported to have escaped each year from 2000 to 2005, which is a high proportion compared with salmon. We interviewed employees at 19 coastal sea‐cage cod farms in Norway to investigate both how and why cultured cod escape and to document cage handling and management strategies that were effective in minimizing escapes. Based on the interviews, we describe five working hypotheses that may explain why a greater proportion of cod than salmon escape: (1) cod are more willing to escape than salmon; (2) cod bite the net cage and create wear and tear; (3) net cages have insufficient technical standards for cod culture; (4) cod are placed in sea cages at considerably smaller sizes than salmon; and (5) cod are more popular feed for predators. Preliminary testing of the hypothesis that cod bite netting and create holes was done by placing pre‐damaged net panels with cut twines and control panels inside sea cages. Holes in the pre‐damaged net panels increased in size over a period of 3 months. The type of damage indicated that biting of netting twines was the likely cause.  相似文献   

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