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.     

Egg mortality and hatching rate of Baltic cod (Gadus morhua) in different salinities

Egg mortality of Baltic cod (Gadus morhua L.), collected off northern Gotland, Sweden, in 1990, was studied in four different salinities — 10 and 15 ppt (salinity of the principal spawning areas of Baltic cod) and 5 and 7 ppt (salinity above the halocline) — in laboratory experiments. Mortality was high during the first 4 d of development, but after gastrulation mortality was low in all salinities tested, except for 5 ppt, in which mortality increased slightly before hatching. Mortality during hatching varied considerably with salinity. No hatching occurred in 5 ppt salinity, and only a few larvae survived in 7 ppt salinity; in contrast, mortality during hatching was comparatively low in salinities of 10 and 15 ppt.     

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.     

Long-term trends in abundance of cladocerans in the Central Baltic Sea

Long-term dynamics (1960-1997) of the cladoceran species Bosmina coregoni maritima, Evadne nordmanni and Podon spp. are described for the Gdansk Deep and the Gotland Basin (Central Baltic Sea). By using correlation analyses on seasonal time-series, the influence of temperature and salinity on the abundance of cladoceran species was investigated. A clear affinity to higher temperature was found for B. coregoni maritima in summer as well as for E. nordmanni and Podon spp. in spring. In addition to temperature, association tests with salinity revealed besides species-specific preferences, regional and temporal differences. Contrary to B. coregoni maritima, both other species were positively associated to salinity in summer and autumn in the Gdansk Deep. In the Gotland Basin only E. nordmanni was positively correlated to salinity in autumn. Differences in the response to hydrographic variables are possibly stage specific, i.e. between resting eggs and adults, or due to a different adaptation to the abiotic environment.     

Physiologische Anpassungen eines marinen insekts. II. Die Eigenschaften von schwimmenden und absinkenden Eigelegen

The gelatinous egg-masses of Clunio marinus (Diptera; Chironomidae) consist of a tube of jelly containing up to 175 eggs. The swimming egg-mass of the Atlantic populations (Race A) and the sinking egg-mass of the Baltic populations (Race B) have a higher density than the sea water, on which the female spawns; salinity is not important to the swimming or sinking of the egg-masses. Density and sinking velocity of gelatinous egg-masses from Atlantic midges are significantly higher than those from Baltic midges if the eggs are laid on water of the same salinity. The density of the eggs and the quantitative relation of eggs to jelly are the same in both races. The jelly of the egg-masses from the Baltic Sea race swells more than that from the Atlantic Sea race (increase of volume by absorption of water = 3-fold and 2.4-fold, respectively). During swelling the jelly alters from a trough-like shape into a tube-like from which closes around the eggs. If the surface tension of the spawning medium is lowered by use of detergents, the A-spawn will sink. A- and B-spawns occupy different positions at an oil-water interface. Interference contrast microscopy revealed that the A-jelly forms a well-defined outer line in relationship to the surrounding aqueous medium; this line is missing in the B-jelly. The morphological fine surface structures, as seen by the scanning electron microscope, do not explain the different properties of the A- and B-spawn. Extracts and hydrolysates of jelly material were separated by thin-layer chromatography. The results indicate that the spawn jelly of C. marinus consists of a polysaccharide-protein-complex, similar to the jellies of other invertebrates. It is rather probable that the chemical structure or the components of the jellies from A- and B-populations are not exactly the same; this would explain the different properties described. The ecological significance of swimming and sinking gelatinous egg-masses and the taxonomical position of both midge populations are discussed.     

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.     

Interactions between cod, herring and sprat in the changing environment of the Baltic Sea: A dynamic model analysis

The interactions between cod (Gadus morhua), herring (Clupea harengus) and sprat (Sprattus sprattus) in the Central Baltic Sea were examined with a simple dynamic model, an alternative to more complicated and data-demanding multispecies and ecosystem models. The main aims of the study were to compare the effect of alternative structures on the model output and examine the control relationships in the fish assemblage under different environmental conditions. The effect of environmental conditions was modelled using a stock-recruitment equation for cod incorporating an environmental index. The model output was especially sensitive to the functional response in predation by cod on herring and sprat. The type II functional response led to a collapse of the clupeid stocks when cod was abundant, while the type III response produced more realistic stock dynamics. According to the simulations, an abundant cod stock was able to keep the sprat stock at a low level, while the herring stock was less affected and benefited from the decreased density of sprat. Simulation of different fishing scenarios indicated that reducing fishing mortality to the level currently advised by ICES would allow the recovery of the cod stock even in unfavourable environmental conditions.     

Cycles and rates of reproduction of six Baltic Sea bivalves of different zoogeographical origin

All bivalves investigated (Astarte borealis, Astarte elliptica, Macoma calcarea, Cyprina islandica, Macoma baltica, Cardium lamarcki) are dioecious. The mean egg-diameters were determined by a series of measurements for 6 bivalves and compared with data from other authors. M. calcarea does not have a direct non-pelagic larval development as previously assumed, but probably produces pelagic planktotrophic larvae. The arctic species exhibit no modification of their mode of development in the boreal area. The boreal forms spawn over a relatively short period in spring and summer. The arctic forms revealed, over long periods, ripe eggs and sperms. Apparently they begin to spawn in winter or early spring. This extension and shift of the spawning season represents adaptation of the Aretic species to the specific conditions of the Baltic Sea. The number of eggs laid by the most important bivalves were determined and correlated with the larval ecology and length of generation time.     

Photosynthetic production and its regulating factors in the Baltic Sea

Measurements of primary production in the Baltic Sea over a period of 2 years (1969/1971) by the ¹⁴C method are reported. The rate of total annual production for the Baltic Sea proper and the Gulf of Finland including coastal areas appears to lie between 35 and 40 gC·m², a rate which can be assumed as characteristic for oligotrophic waters with a low production rate. An apparent increase in productivity between the 2 years could be noted. Calculations for the water mass below the halocline show that between 0.7 and 0.8 ml/l oxygen is used up during a period of 3 months required for the development of anoxic conditions. The major portion of the organic matter in the Baltic Sea undergoes decomposition in the sub-photic zone above the halocline. The ratio of organic material in deep water to the total available oxidizable matter appears to remain constant with time; therefore, the apparent increase in productivity in the photic layer does not affect apprecibly the development of anoxic conditions in deep water. Of the different areas investigated, the Gulf of Finland seems to be the most productive. In the southern Baltic Sea, the rate of production over 5 years between 1966 and 1971 has not changed appreciably. In fact, it shows a trend toward a slight decrease, whereas, in other areas of the Baltic Sea, the rate of production indicates an apparent increase. Comparison with oceanic coastal areas of the west coast of Sweden shows that the rate of production there is higher than in the Baltic Sea. Of the micronutrients controlling productivity in the photic layer of the Baltic Sea, nitrate has a stronger limiting effect on plant growth than phosphate, a fact which is in agreement with existing conceptions on the subject. Dissolved iron does not seem to exert any appreciable influence on the productivity.     

Calanoid resting eggs in the Baltic Sea: implications for the population dynamics of Acartia bifilosa (Copepoda)

Resting eggs of Acartia spp. (presumably A. bifilosa Giesbr.) at densities between 65 and 125 eggs cm^-2 were found in sediment off the southern coast of Finland and represent the first observations of dormant calanoid eggs in the Baltic Sea. The highest egg densities were found at sediment depths between 4 and 8 cm. The hatching percentage varied between 0 and 90% in the different sediment layers and at different experimental temperatures (13, 16 and 19°C). Some nauplii emerged from the eggs extracted from sediment layers 9 to 10 cm beneath the surface layer and other nauplii emerged from eggs after 82 d of dark storage at 3°C. Light was not needed to trigger hatching; the nauplii were able to hatch in darkness when the eggs were resuspended in filtered seawater. Indirect evidence suggests that the marked spring peak of Acartia spp. nauplii seen in a monitoring data from the years 1973 to 1984 was derived from benthic eggs and not from spawning. The possible mechanisms for induction of hatching and the adaptive benefits of resting egg production for A. bifilosa are discussed.     

The influence of temperature on the development of Baltic Sea sprat (Sprattus sprattus) eggs and yolk sac larvae

In spring 2004 and 2005 we performed two sets of experiments with Baltic sprat (Sprattus sprattus balticus Schneider) eggs and larvae from the Bornholm Basin simulating ten different temperature scenarios. The goal of the present study was to analyse and parameterise temperature effects on the duration of developmental stages, on the timing of important ontogenetic transitions, growth during the yolk sac phase as well as on the survival success of eggs and early larval stages. Egg development and hatching showed exponential temperature dependence. No hatching was observed above 14.7°C and hatching success was significantly reduced below 3.4°C. Time to eye pigmentation, as a proxy for mouth gape opening, decreased with increasing temperatures from 17 days post hatch at 3.4°C to 7 days at 13°C whereas the larval yolk sac phase was shortened from 20 to 10 days at 3.8 and 10°C respectively. Maximum survival duration of non-fed larvae was 25 days at 6.8°C. Comparing the experimental results of Baltic sprat with existing information on sprat from the English Channel and North Sea differences were detected in egg development rate, thermal adaptation and in yolk sac depletion rate (YSDR). Sprat eggs from the English Channel showed significantly faster development and the potential to develop at temperatures higher than 14.7°C. North Sea sprat larvae were found to have a lower YSDR compared to larvae from the Baltic Sea. In light of the predictions for global warming, Baltic sprat stocks could experience improved conditions for egg development and survival.     

Biological effects of oil on early development of the Baltic herring Clupea harengus membras

The effects of petroleum hydrocarbons from two crude oils and one fuel oil (No. 1) were studied on the ontogenic development of the Baltic herring Clupea harengus membras L. Herring eggs exposed to water-soluble fractions of the oils at the time of fertilization showed no decrease in numbers of fertilized eggs compared to eggs exposed 6 or 72 h after fertilization. During embryongenesis, treatment with 3.1 to 8.9 ppm or 3.3 to 11.9 ppm total oil hydrocarbons from light fuel oil and the two crude oils respectively, gave rise to alterations in embryonic activity, decreased heart rate, and premature or delayed hatching. Although many larvae hatched from eggs exposed to contaminated water (3.1 to 11.9 ppm total oil hydrocarbons), the majority of the (70 to 100%) were malformed or dead 1 day after hatching. Exposure of eggs to 5.4–5.8 ppm total oil hydrocarbons resulted in significantly (P<0.001) decreased lengths of the larvae. Increased temperature (from 9° to 14°C) aggravated the effects of the oils. The results are discussed in relation to the potential effects of oil spills and chronic oil pollution on fish eggs and larvae in the Baltic Sea.     

Introgression and mitochondrial DNA heteroplasmy in the Baltic populations of mussels Mytilus trossulus and M. edulis

A strong clinal change in salinity occurs between the Baltic Sea and the North Sea, Atlantic Ocean, in the Danish Straits, where hybridization zone between mussels Mytilus edulis and M. trossulus has been reported. Eleven samples of mussels were studied from the Danish Straits and the inner Baltic Sea. Extensive introgression of M. edulis alleles from the North Sea into populations throughout the Baltic was ascertained for mitochondrial DNA (mtDNA) and two nuclear markers (ME15–16 and ITS). In the opposite direction, introgression of M. trossulus alleles into the M. edulis background was observed at the EFbis nuclear marker in populations from Kattegat (Danish Straits). While only M. edulis F (female) mtDNA was present in the Baltic, there were still strong differences in frequencies in the control region length variants between the Danish Straits and the inner Baltic samples, and weaker variation in coding region ND2–COIII haplotype frequencies. In the assays of the two mtDNA regions, various patterns of heteroplasmy were detected in 32% of all the studied individual mussels; this includes the presence of distinct, independently inherited M and F mitochondria in males, as well as the presence of two different distinguishable F genomes. The male-inherited M mtDNA genomes are quite common in the mussels from the Danish Straits, but very rare in males from the inner Baltic. Instead, a recombined control region variant (1r), which seems to have taken over the role of the M genome, was present in a number of specimens in the Baltic. Observations of heteroplasmy for two F genomes in some females and males confirm disruptions of the doubly uniparental inheritance mechanism in the hybrid Baltic Mytilus.     

Wind induced upwelling as a possible explanation for mass occurrences of epiphyticEctocarpus siliculosus (Phaeophyta) in the northern Baltic Proper

The quantity of epiphytic filamentous algae varies considerably in macroalgal vegetation along rocky shores of the northern Baltic Proper. The main species responsible for irregular mass occurrences is the summer annual brown algaEctocarpus siliculosus (Dillw.) Lyngb. In this study, data collected over a 3-yr monitoring period are related to hydrographical parameters. The tideless and brackish Baltic Sea is salinity stratified, and the salinity difference between surface and bottom waters can be used to indicate upwelling events. Mass occurrences developed when the salinity difference was low. These salinity differences explained 87% of the observed increase inE. siliculosus cover. We propose that the slight salinity changes themselves do not affect the growth ofE. siliculosus, but rather they can be used as indicators of short nutrient pulses. These nutrient pulses are not detected in standard water sampling, as they are utilised by both pelagic and benthic organisms within a few hours or days. If the proposed mechanism is true, changes in wind speed and direction have an important effect on the nutrient dynamics of littoral algal communities. A higher frequency of SW winds may cause mass occurrences of filamentous algae, which are often believed to indicate eutrophication of an anthropogenic origin.     

Radioactivity in the Baltic Sea

The Baltic Sea is, like the Mediterranean, a marginal sea, which with the Black Sea, were marine environments contaminated from the Chernobyl accident.

Radiocaesium and plutonium isotopes were studied in water, sediment and macroalgae in the Baltic Sea since 1982. the inventory of ¹³⁷Cs in the Baltic increased from 0.65 PBq to 5.85 PBq following the Chernobyl accident. the corresponding increase for ^{239 + 240}Pu was less significant and yielded 1.5 TBq to a total value of 16.5 TBq.

For plutonium, 98% is trapped in the sediment and the net-exchange of this element through the Baltic straits is very small (1 GBq/year), while for radiocaesium, 45% is in the water phase and there is a net-loss of 60 000 GBq annually into the adjacent water (Kattegatt).     

In-situ observations of Baltic herring (Clupea harengus membras) spawning behaviour in the Askö-Landsort area,northern Baltic proper

In a Baltic herring (Clupea harengus membras L.) spawning ground study, carried out in summer 1982 in the northern Baltic proper, spawning herring were encountered on two line transects on June 14. The spawning behaviour of minor groups as well as of a large school was studied in situ by four divers for approximately three hours. No behavioural differences or systematic coordinated behaviour between the sexes could be detected. Most of the spawning took place in a limited depth interval, 0.5 to 4 m, and Ceramium tenuicorne was the substrate most used.     

Icy heritage: ecological evolution of the postglacial Baltic Sea reflected in the allozymes of a living fossil,the priapulid Halicryptus spinulosus

Genetic variation of 16 allozyme loci in 397 Halicryptus spinulosus (Priapulida) revealed overall polymorphism of P=0.438 and Hardy-Weinberg expectations for heterozygosity of H _e=0.060 for Baltic Sea stocks, H _e=0.143 for the White Sea and H _e=0.121 for Iceland. Maximal unbiased standard distances of D=0.0693 separated Baltic and White Sea populations. Nordic and Baltic populations could be distinguished by allozymes, but Baltic subsamples proved cohesive. Gene flow amounted to effective exchange values per generation of N _m=2.94 over 650 km of continuous habitat, N _m=10.65 over 175 km, and N _m=13.85 over 20 km. Gene flow started to decrease with geographic distance beyond a dispersal threshold of 20 km, but hierarchical G _ST-statistics indicated light isolation by distance beyond a minimum of 8 km. Gene flow is high for a benthic worm assumed to lack dispersal by pelagic larva, a paradox which cannot be resolved now. Baltic populations are characterized by lower heterozygosity than Nordic stocks. In the Baltic Basin, temporally continuous brackish-water conditions have only existed for the past 7000 years. The two possible colonization routes of H. spinulosus to the geologically young Baltic Sea imply genetic drift, whether by founder effect (sweepstake colonization from Iceland) or by refugial bottlenecking during the Ancylus phase of the Baltic Basin after a direct connection to the White Sea had been sequestered. Continued genetic drift is emphasized by lower heterozygosity in the ecologically unstable Belt Sea compared to the central Baltic. Allozymes falsify the reduced-mutability hypothesis to explain bradytelic evolution of Priapulida. Regional genetic homogeneity, ample polymorphism, and preference for anoxic black mud qualify H. spinulosus populations as indicators of microevolutionary responses to water circulation regimes or pollution in the Baltic Sea.     

High natural mortality of Baltic herring (Clupea harengus) eggs caused by algal exudates?

Results from field studies in 1978 and 1982 together with results from a laboratory study carried out in 1984 indicate that the unusually high natural mortality of Baltic herring (Clupea harengus L.) eggs observed in situ is presumably caused by toxic exudates released by filamentous brown algae during a limited period which happens to coincide with the peak of spawning. Oxygen deficiency is no longer believed to be the major cause of the observed unnaturally high mortality. It cannot, however, be entirely ruled out.     

Do cod form spawning leks? Evidence from a Newfoundland spawning ground

Catch and acoustic telemetry data were used to test three predictions of lekking behaviour in Atlantic cod (Gadus morhua L.) at a small-scale spawning ground (∼25 km²) in Placentia Bay, Newfoundland: (1) that cod form male-skewed aggregations during spawning; (2) spawning arenas occur at consistent sites; and (3) on the arenas, there will be higher proportions of spawning fish. Forty-two fishing sets from 1998 to 2003 indicated male-skewed sets early in the spawning season (April) in 5 of 6 survey years. Male-skewed sets were consistently distributed at depths < 50 m and mostly in one part of the ground. The proportions of spawning females and spent males were significantly higher in male-skewed sets, whereas sets with equal sex ratios had significantly higher proportions of immature males and spent females. In addition, cod of both sexes were significantly larger in male-skewed sets. Telemetric tracking of 25 cod in 2002 and 2003 (12 males, 13 females) indicated that both males and females were highly mobile while present on the spawning ground. Overall, our results add support to the hypothesis that cod employ a lekking mating system.     

Mesozooplankton resting eggs in the Baltic Sea: identification and vertical distribution in laminated and mixed sediments

Benthic resting eggs of rotifers, cladocerans and calanoid copepods were studied at two sites with different hydrographical and sediment properties off the SW coast of Finland, the northern Baltic Sea between 1991 and 1993. Vertical distribution of the resting eggs in the sediment was studied down to 10 cm depth, and hatching of the eggs extracted from different 1-cm thick sediment layers was experimentally tested. The larvae of eight mesozooplankton taxa emerged from the eggs: Calanoida (mostlyAcartia bifilosa), cladoceransBosmina spp.,Daphnia spp.,Evadne nordmanni, andPodon spp. (mostlyP. polyphemoides), and rotifersSynchaeta spp.,Keratella spp. andAsplanchna priodonta. Some calanoid and cladoceran eggs hatched after 7 mo storage of the sediment at 3°C. A few calanoid and cladoceran eggs hatched from the 8 to 10 cm deep sediment layer. Their age, estimated by means of¹³⁷Cs analysis, was ca. 8 to 10 yr. Vertical distribution of the benthic eggs reflected the different sediment structures of the study sites. The eggs of all zooplankton taxa were more stratified in the laminated sediment than in the disturbed sediment. We suggest that the intensity of deep water mixing is the key factor for regulating the vertical distribution and hatching probability of mesozooplankton resting eggs, since it largely determines the stratification of the sedimenting material, distribution of oxygen within the sediment, and the living conditions of benthic animals causing bioturbation.