First evidence of spawning of eastern Baltic cod (Gadus morhua callarias) in the Belt Sea,the main spawning area of western Baltic cod (Gadus morhua L.)

Two cod stocks (western Baltic cod, WBC, and eastern Baltic cod, EBC) are managed in the Baltic Sea which is characterized by different main spawning areas and different main spawning periods. In this study we analyse the spatial and temporal occurrence of spawning individuals of both cod stocks in the main spawning grounds of the Baltic Sea based on eight microsatellite loci. Our results suggest that EBC (Gadus morhua callarias) has formed currently temporally stable, substantially homogeneous population not only in the Bornholm Sea (ICES SD: 25) but also in the Arkona Sea (ICES SD: 24). The presented analyses proved that EBC (G. m. callarias) can temporarily also spawn in the Belt Sea.     

TaqMan DNA technology confirms likely overestimation of cod (Gadus morhua L.) egg abundance in the Irish Sea: implications for the assessment of the cod stock and mapping of spawning areas using egg-based methods

Recent substantial declines in northeastern Atlantic cod stocks necessitate improved biological knowledge and the development of techniques to complement standard stock assessment methods (which largely depend on accurate commercial catch data). In 2003, an ichthyoplankton survey was undertaken in the Irish Sea and subsamples of 'cod-like' eggs were analysed using a TaqMan multiplex, PCR (polymerase chain reaction) assay (with specific probes for cod, haddock and whiting). The TaqMan method was readily applied to the large number of samples (n = 2770) generated during the survey and when combined with a manual DNA extraction protocol had a low failure rate of 6%. Of the early stage 'cod-like' eggs (1.2-1.75 mm diameter) positively identified: 34% were cod, 8% haddock and 58% whiting. As previous stock estimates based on egg surveys for Irish Sea cod assumed that the majority of 'cod-like' eggs were from cod, the TaqMan results confirm that there was probably substantial contamination by eggs of whiting and haddock that would have inflated estimates of the stock biomass.     

Selectivity of fishing gears used in the Baltic Sea cod fishery

The Baltic cod (Gadus morhua) plays a very significant role in the Baltic Sea ecosystem being the major fish top predator and the most important commercial species for the Baltic Sea fishing industry. Consequently the management and understanding of the exploitation pattern of the stock is of major importance. Technical regulations, improving the selectivity of cod, have been a major management strategy and the Baltic Sea is likely to be the area where most fishing gear selectivity studies, focussing on size selectivity, have been conducted over time. The methodology for conducting and analysing selectivity data has been significantly improved in recent years. The subject is reviewed since the choice of methodology can have a significant effect on the interpretation of the outcome of selectivity experiments. Factors affecting the selectivity of trawl and gill nets are considered. Alternative ways to improve the size and species selectivity of trawls using selective devices are reviewed. Selectivity parameters from available literature are listed and the correlations of selectivity parameters to the mesh size for different gears are estimated. The historical legislation on selectivity is reviewed and the expected selectivity for trawls is estimated. Management considerations concerning the mortality of escaping and discarded fish and wider management impacts have to be considered if improving selectivity. The review is ended by conclusions including reflections on needed research in the future.     

Long-term variation in dates of mass spawning of cod Gadus morhua callarias (Gadidae) in the southeastern part of the Baltic Sea

Long-term data on the numbers of eggs of cod Gadus morhua callarias in ichthyoplankton and the seasonal variation of their distribution in the Gdansk Deep of the Baltic Sea are considered. The spring peak of spawning was observed for the years 1948–1954 and 1960–1964; the spring-summer peak for 1968–1977; and the summer peak for 1955–1959 and 1995–2001. Maximum egg numbers of cod were recorded from the late 1940s to the early 1950s; minimums were recorded in the late 1980s and the late 1990s. The decrease in the numbers of cod eggs in the ichthyoplankton, in general, coincided with a decrease in the proportion of spawners maturing at the beginning of the season of mass reproduction and a decrease in the oxygen content in the near-bottom layer of the Gdansk Deep. It is suggested that a shift of the spawning peak of cod from spring to summer is related to a decrease in the volume of near-bottom waters providing favorable conditions for the reproduction of this population.     

Prehistoric versus modern Baltic Sea cod fisheries: selectivity across the millennia

Combining Stone Age and modern data provides unique insights for management, extending beyond contemporary problems and shifting baselines. Using fish chronometric parts, we compared demographic characteristics of exploited cod populations from the Neolithic Period (4500 BP) to the modern highly exploited fishery in the central Baltic Sea. We found that Neolithic cod were larger (mean 56.4 cm, 95% confidence interval (CI)+/-0.9) than modern fish (weighted mean length in catch =49.5+/-0.2 cm in 1995, 48.2+/-0.2 cm in 2003), and older (mean ages=4.7+/-0.11, 3.1+/-0.02 and 3.6+/-0.02 years for Neolithic, 1995, and 2003 fisheries, respectively). Fishery-independent surveys in 1995 and 2003 show that mean sizes in the stock are 16-17 cm smaller than reflected in the fishery, and mean ages approximately 1-1.5 years younger. Modelled von Bertalanffy growth and back-calculated lengths indicated that Neolithic cod grew to smaller asymptotic lengths, but were larger at younger ages, implying rapid early growth. Very small Neolithic cod were absent and large individuals were rare as in modern times. This could be owing to selective harvests, the absence of small and large fish in the area or a combination. Comparing modern and prehistoric times, fishery selection is evident, but apparently not as great as in the North Atlantic proper.     

Nitrogen Fixation During Blooms of Nodularia in Coastal Waters and Backwaters of the Arkona Sea (Baltic Sea) in 1974

Nitrogen fixation was investigated by means of the acetylene reduction method during the development of a water bloom of Nodularia in coastal waters of the Baltic Sea west of the island of Hiddensee and in backwaters showing different degrees of eutrophication. Depending on plankton density, the values found varied greatly. The maximum of nitrogen fixation values found in extremely dense water blooms under special conditions (Baltic Sea, 2250 μg N₂/l · h; Kleiner Jasmunder Bodden, 374 μg N₂/l · h) are up to 10³ times higher than from other parts of the Baltic Sea or from inland waters. The average nitrogenase activity determined for coastal water populations of the Baltic Sea is 2.15 pg N₂/heterocyst · h and that of the inmost backwaters 0.77 pg N₂/heterocyst · h. The relationship between N₂-fixation and nutrient content in water is discussed.     

The potential fecundity of cod in the Baltic Sea from 1993 to 1999

Investigations on the reproduction biology of cod in the Baltic Sea (Gadus morhua L. and G. morhua callarias L.) were carried out beginning in 1993, with yearly sampling of gonads to estimate the potential absolute and relative individual fecundity in the Belt Sea (subdivision, SD: 22) and in the Arkona Sea (SD: 24). Additional samples were taken in the Bornholm Sea (SD: 25). The potential absolute fecundity of the total area of investigation is described by the regression models: F_a = 6.89 × TL^3.09 and F_a = ?40 586 + 977.7 × W. As a further result the mean relative fecundity was 978 oocytes per gram body weight. Furthermore, it was shown that there was a difference in development of the potential individual fecundity between the Belt Sea cod and the original eastern Baltic cod stock. In contrast to the stable fecundity of cod in the Belt Sea, the fecundity of the large females in the Bornholm Sea significantly increased from 1993 to 1999. This change is related to a decrease in the dry weight of oocytes in the ovaries. In the same period this cod stock dramatically decreased.     

Impact of hypoxia on consumption of Baltic cod in a multispecies stock assessment context

The Baltic Sea is characterised by a heterogeneous oceanographic environment. The deep water layers forming the habitat of Baltic cod (Gadus morhua callarias L.) are subjected to frequently occurring pronounced anoxic conditions. Adverse oxygen conditions result in physiological stress for organisms living under these conditions. For cod e.g. a direct relationship between oxygen availability and food intake with a decreasing ingestion rate at hypoxia could be revealed. In the present study, the effects of oxygen deficiency on consumption rates were investigated and how these translate to stock size estimates in multi‐species models. Based on results from laboratory experiments, a model was fitted to evacuation rates at different oxygen levels and integrated into the existing consumption model for Baltic cod. Individual mean oxygen corrected consumption rates were 0.1–10.9% lower than the uncorrected ones. At the currently low predator stock size, however, the effect of oxygen‐reduced consumption on the total amount of eaten prey biomass and thus predation mortalities was only marginal. But should successful management lead to higher cod stock sizes in the future, then total predation mortalities will greatly increase and thus improved precision of these estimates would be valuable for the assessment of prey stocks.     

The ability of Baltic cod eggs to maintain neutral buoyancy and the opportunity for survival in fluctuating conditions in the Baltic Sea

In the brackish water of the Baltic Sea successful spawning of Baltic cod Gadus morhua is restricted to the Bornholm, Gdansk and Gotland basins below the halocline, occurring at 50–80 m depth. Due to irregular mixing of the deep water, stagnant conditions occur regularly accompanied with unfavourable oxygen conditions. In avoiding stressful oxygen conditions maintenance of egg buoyancy is considered a major limiting factor for successful spawning of Baltic cod. Batches of eggs were incubated experimentally in a density gradient column. Egg specific gravity changed during development, decreasing from the time of gastrulation, then increasing prior to hatching. The changes in specific gravity varied among egg batches from different females and were related to egg quality, egg size and ambient salinity. Eggs achieve different specific gravity depending on incubation salinity. Initial egg specific gravity together with the ability of eggs to gain and maintain buoyancy up to hatching, determine larval specific gravity and the depth where hatching will occur, and thus opportunities for larval survival, avoiding stressful oxygen conditions and developing at favourable feeding conditions.     

Seasonal changes in energy and the energy cost of spawning in Gulf of Alaska Pacific cod

In the Gulf of Alaska, adult Pacific cod exhibited an annual cycle of condition, gonad index and liver index in which maximum values occurred in ripe fish in March and minima in July. About 30–31 % of prespawning stored energy was expended during the spawning effort. The energy associated with spawning derived from liver (24% and 18%), somatic tissue (22% and 33%) and gonad (53% and 48%) for females and males, respectively. Liver index and gonad index at the time of sampling were directly related in females, but in males gonad index was best related to liver index 1–3 months earlier.
The Pacific cod is very similar to the Atlantic cod in terms of energy cycling, maximum gonad sizes, energy expended during spawning and gonadal contribution to energy expenditure. However, in Pacific cod, somatic tissue contributes markedly to energy expended during reproduction. The Pacific cod cod differs from the walleye pollock with respect to gonad index (13% and 20%ν. 20% and 8% for females and males, respectively), spawning weight loss (25%ν. 38%), liver energy loss during spawning (71%ν. 55%) and energy cost of spawning.     

Physiological-biochemical aspects of omission of the spawning period in the Barents Sea cod Gadus morhua

Using histological and biochemical methods, ovaries of the Barents Sea cod Gadus morhua were studied in different physiological states (immature fish, normal maturation, omission of spawning). Among the fish missing spawning, two categories are identified: without signs of occyte maturation and with massive resorption of maturing oocytes. The maximal activity of lysosomal and cytoplasmic proteolytic enzymes was revealed in the fish ovaries with histological signs of massive oocyte resorption. In the ovaries of maturing fish the highest concentrations of water-soluble proteins was found. The molecular-mass protein composition and the fraction responsible for the cytosol proteinase activity were determined by electrophoresis and high-performance liquid chromatography (HPLC). The obtained results allow some suggestions to be made about mechanisms of degradation of cellular proteins and functions of proteinases in various physiological states of the cod gonads.     

The role of spawning stock in formation of recruitment of northeast atlantic cod Gadus morhua

On the basis of statistical treatment of the data for 1946–2003 (correlation analysis and analysis of variance) on the biomass of the spawning stock (SSB) and of the resulting recruitment (R), expressed as abundance of three-year-old fish (N₃), of the Northeast Atlantic cod Gadus morhua, the relationship SSB → N₃ is revised and the extent of the influence of SSB on the formation of R is assessed. The relationship of SSB → N₃ turned out to be significant only at SSB ≤ 250 000 t. The influence of SSB is only 4.3–14.7%, in comparison to the other background factors controlling R. This casts doubt on the adequacy of the so-called precautionary approach in fishery management, popular in international fishery organizations, which is based on efforts to support the spawning stock at a certain level (B_pa) taken as the optimum one.     

Description of growth by simple versus complex models for Baltic Sea spring spawning herring

The objective was to find a length–growth model to help differentiate between herring stocks (Clupea harengus l.) when their length–growth shows systematically different patterns. The most essential model restriction was that it should react robustly against variations in the underlying age range which varies not only over time but also between the different herring stocks. Because of the limited age range, significance tests as well as confidence intervals of the model parameters should allow a small sample restriction. Thus, parameter estimation should be of an analytical rather than asymptotic nature and the model should contain a minimum set of parameters. The article studies the comparative characteristics of a simple non‐asymptotic two‐parameter growth model (allometric length–growth function, abbreviated as ALG model) in contrast to higher parametric and more complex growth models (logistic and von‐Bertalanffy growth functions, abbreviated as LGF and VBG models). An advantage of the ALG model is that it can be easily linearized and the growth coefficients can be directly derived as regression parameters. The intrinsic ALG model linearity makes it easy to test restrictions (normality, homoscedasticity and serial uncorrelation of the error term) and to formulate analytic confidence intervals. The ALG model features were exemplified and validated by a 1995 Baltic spring spawning herring (BSSH) data set that included a 12‐year age range. The model performance was compared with that of the logistic and the von‐Bertalanffy length–growth curves for different age ranges and by means of various parameter estimation techniques. In all cases the ALG model performed better and all ALG model restrictions (no autocorrelation, homoscedasticity, and normality of the error term) were fulfilled. Furthermore, all findings seemed to indicate a pseudo‐asymptotic growth for BSSH. The proposed model was explicitly derived for of herring length‐growth; the results thus should not be generalized interspecifically without additional proof.     

On differentiation of cod (Gadus morhua L.) groups in Baltic Sea

Russian Journal of Genetics - Using the AGP*, PGI-1*, PGI-2*, LDH*, IDH*, and PGM* allozyme markers, the differentiation of cod groups during the spawning period in Baltic Sea was evaluated. It was...     

The dangers of ignoring stock complexity in fishery management: the case of the North Sea cod

The plight of the marine fisheries is attracting increasing attention as unsustainably high exploitation levels, exacerbated by more extreme climatic conditions, are driving stocks to the point of collapse. The North Atlantic cod (Gadus morhua), a species which until recently formed a major component of the demersal fisheries, has undergone significant declines across its range. The North Sea stock is typical of many, with a spawning stock biomass that has remained below the safe biological limit since 2000 and recruitment levels near the lowest on record. Cod within the North Sea are currently managed as a single stock, and yet mounting empirical evidence supports the existence of a metapopulation of regionally variable, genetically distinct, sub-stocks. Applying the same management strategies to multiple stocks that differ in their resilience to exploitation inevitably results in the overfishing and likely collapse of the weaker components. Indeed, recent studies have identified two North Sea spawning stocks that have undergone disproportionally large collapses with very substantial reductions in egg production. Similarly affected cod stocks in the northwest Atlantic have shown little evidence of recovery, despite fishery closures. The possible implications of ignoring sub-structuring within management units for biocomplexity, local adaptation and ecosystem stability are considered.     

Histology of pseudobranchial tumours in Atlantic cod(Gadus morhua) from the North Sea and the Baltic Sea

Atlantic cod bearing pseudobranchial tumours were investigated from both North Sea and Baltic Sea areas. Light and electron microscopic studies revealed a tumorous structure previously described in cod from Atlantic and Pacific waters. Additionally, cysts, multinucleated cells and early stages of mitosis are described. These findings support the hypothesis that pseudobranchial tumours are instigated by a protozoan infection causing xenotumours associated with the pseudobranch organ. The cells contained in these lesions showed similarities to amoebae.     

Assessing seasonal changes in stock composition of Atlantic salmon catches in the Baltic Sea with genetic stock identification

The feasibility of using genetic stock identification to analyse seasonal changes in stock compositions of Atlantic salmon catches in the Baltic Sea was examined. The analysis employed seven variable allozyme loci from most of the potentially contributing stocks (16) from Finland and Sweden. Catch samples were collected from Finnish salmon fisheries in the eastern Bothnian Sea during the 1992 fishing season. Simulation studies were used to evaluate the feasibility of identifying Baltic salmon stocks with allozyme data. Special attention was paid to analysing the wild production of salmon stocks. Clear seasonal differences in stock composition were found. The estimates were compared with smolt production and Carlin-tag data. The proportions of the Neva and Oulujoki river stocks could be estimated as individual stocks, whereas the contributions of the remaining stocks were estimated as four composite stock groups. One of the groups consisted of wild stocks from the rivers Kalixälven and Simojoki. Identification of this group, which could be used as an index of wild production in the catches, requires catch sample sizes >300 salmon if <15% error is required.     

Epidemiology of pseudobranchial tumours in Atlantic cod(Gadus morhua) from the North Sea and the Baltic Sea

Investigations during several cruises (1979–1981) showed that Atlantic cod with pseudobranchial tumours occurred in all areas investigated. Statistical analysis of data revealed that there was a marked prevalence of the condition in the centre of the German Bight, whereas a lower incidence of the condition was observed on the Dogger Bank, off the Dutch coast and in the western Baltic Sea. The higher prevalence of pseudobranchial tumours in the German Bight coincides with a dense population of cod and impacts of pollution.     

Seasonal prevalence and intensity of follicular atresia in Baltic cod Gadus morhua callarias L.

In the present study, 307 ovaries of eastern Baltic cod Gadus morhua callarias sampled during the prespawning and spawning season 2000 were analysed histologically to estimate the seasonal prevalence and intensity of atresia. The number of atretic oocytes per ovary was estimated using a combination of the physical disector method and volume fraction (Delesse principle). Atretic oocytes were observed in 32% of the ovaries. Prevalence of atresia was independent of female size, but increased significantly with declining female condition from prespawning and through the spawning stages. The relative intensity of atresia, i.e. number of atretic oocytes in relation to normally developed vitellogenic oocytes, was low amounting to 1·4% on average. Similar to prevalence, relative intensity of atresia differed significantly between maturity stages and increased with decreasing female condition. The population egg loss due to atresia amounted to 4·6% indicating that Baltic cod was performing close to maximum productivity, i.e. potential egg production.