Distribution of labile and residual particulate carbon in the Baltic Sea

During the September 1971 cruise of F. K. Alkor in the central Baltic Sea, the surface or summer water layer down to a depth of about 30 m was found to contain 187.4 g/l of particulate organic carbon, with a C:N (atoms) ratio of 8.97. This carbon was 44% (89.9 g/l) labile to bacterial decomposition, as determined by burning an aliquot of each sample in a CHN-analyser before and after maintenance in nutrientenriched sea water at 20°C for 3 months. The particulate material from the intermediate or winter water layer, with a depth ranging from 30 to 70 m, contained only 48% (43.1 g/l) of the labile carbon found in the summer surface layer, and had a significantly higher C:N ratio (11.25). These two facts indicate that a considerable breakdown of the organic material had taken place. The material, removed from the particulate state during this process, was 48% (89.7 g/l) of the original total particulate carbon, and was relatively nitrogen-rich, with a C:N ratio of 6.49. In this material, 52% (46.8 g/l) of the organic carbon was labile. The particulate material in the rest of the water column showed no significant changes until it reached the sediment. The slurry immediately above the sediment had a C:N ratio of 9.15, indicating the introduction of nitrogen from either dissolved or colloidal material. Labile carbon (44% of the total) was also present in sufficient quantities to support life and to make this an important diagenetic site.     

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).     

Observations of planktonic bioluminescence in the euphotic zone of the California Current

The distributions of bioluminescence, temperature, salinity, oxygen. pH, and chlorophyll a were measured at 10 m intervals, to a depth of 100 m at a station (33°46N; 119°36W) in the California Current from 17 to 20 July 1982. The distribution of bioluminescence showed a marked day-night change which was consistent over the sampling period. The nighttime maximum was at the surface, and the daytime maximum was between 30 and 40 m. The shapes of the day and night distributions were independent of the absolute intensity of bioluminescence and were also insensitive to advection, as inferred from changing temperature-salinity relationships. The nighttime depth distribution broadened during a period of high wind Day to night differences in the color spectrum at the depth of maximum bioluminescence suggest that the luminescent organisms differed from day to night.     

Nitrogenous nutrition in the euphotic zone of the Central North Pacific Gyre

The uptake rates of the three nitrogen compounds ammonium, nitrate, and urea were measured in the oligotrophic North Central Pacific Gyre in August–September 1985. The measurements were performed by using ¹⁵N-labelled substrates and incubating for short-time periods (3 to 4 h) under simulated in situ conditions. Ambient concentrations of the nitrogenous nutrients were generally below 0.10 mol l^-1. The average total daily nitrogen uptake rate, integrated over the euphotic zone, was 12.5 mmol N m^-2 d^-1. Diel studies in the upper water mass resulted in a calculated phytoplankton growth rate of 1.3 d^-1. Ammonium was the dominating nutrient, accounting for on the average 54% of the total nitrogen uptake, while urea uptake represented 32% and nitrate 14%. Ammonium uptake rates at a coastal station off the Hawaiian Islands were very close to the rates found at the oceanic station. Organisms <3 m dominated the nitrogen assimilation, being responsible for about 75% of the ammonium uptake. The nitrogen uptake rates in this study seem to be higher than those found by earlier investigations in the area, but correlated well with other productivity measurements performed during the same cruise.     

Strategies for carbon acquisition in the red marine macroalga Coccotylus truncatus from the Baltic Sea

Marine macroalgae need carbon-concentrating mechanisms because they have only limited access to CO₂ in their natural environment. Previous studies have shown that one important strategy common to many algae is the activity of periplasmic carbonic anhydrases that catalyse the dehydration of HCO₃^- into CO₂. The latter can then cross the plasma membrane by passive diffusion. We hypothesised that an active (energy-consuming) mechanism might also be involved in the membrane transport of CO₂, as is the case in a number of microalgae. Coccotylus truncatus was chosen as a model organism for this study because it belongs to a group of algae that usually lack direct HCO₃^- uptake: sublittoral red algae. The method used to study carbon uptake was pH drift of the seawater medium surrounding the algae in a closed vessel, with and without the addition of specific inhibitors or proton buffers. Measured parameters included pH, total inorganic carbon and alkalinity of the seawater medium. Our results suggest that, in C. truncatus, periplasmic carbonic anhydrase as well as H⁺ extrusion, probably driven by a vanadate-sensitive P-type H⁺-ATPase (proton pump), are involved in CO₂ uptake. No direct uptake of HCO₃^- was discovered. This paper also presents data on the buffer capacity of several proton buffers and the carbon-uptake inhibitors acetazolamide, 4,4'-diisothiocyano-stilbene-2,2'-disulfonate (DIDS) and orthovanadate in Baltic Sea water with a salinity of 6.5 psu.     

Microbial clumps in seawater in the euphotic zone of Saanich Inlet (British Columbia)

Viable heterotrophic microorganisms were enumerated to be 3.7±7.3 bacterial cells per microbial clump during summer in the euphotic zone of Saanich Inlet, British Columbia, Canada. Large microbial aggregates were observed, especially after the phytoplankton bloom, when the phytoplankton biomass formed about 1/2 the total suspended organic matter in the sea. The cell number per microbial clump was minimal when the phytoplankton fraction in the total suspended organic matter was almost 0 (i.e., before the phytoplankton bloom), and again when the phytoplankton bloom occurred. The size of the microbial clumps is discussed, particularly in reference to the food chain in the sea.The work was carried out at the Fisheries Research Board of Canada, Biological Station, Nanaimo, during the tenure of a National Research Council Post-doctoral fellowship.     

Estimating carbon flux to pelagic grazers in the ice-edge zone of the eastern Bering Sea

Zooplankton ingestion of phytoplankton carbon in the iceedge zone of the Eastern Bering Sea was measured using a deck incubation approach in 1982. Using further samples collected in 1983, the plant cell carbon to cell volume ratio was estimated at 0.0604 pg m^–3 from an experimentally determined particulate carbon to seston volume relationship. The application of this conversion to the results of experimental incubations of natural plant stocks with net-caught zooplankton produced ingestion rates of 68.8 and 10.26 mg C g^–1 grazer d^–1 for copepods and euphausiids, respectively. Extrapolating these rates to in situ zooplankton biomass at the edge of the seasonal ice pack yielded carbon flux rates through the zooplankton community ranging between 6.5 and 32.8 mg C m^–2 d^–1. This consumption amounted to less than 2% of the daily phytoplankton production in the ice-edge zone.     

Bacterioplankton in the coastal euphotic zone: Distribution,activity and possible relationships with phytoplankton

Bacterioplankton were studied in the euphotic zone of the Southern California Bight, USA, with special attention to biological factors affecting bacterial distribution and activity. Measurements were made of bacterial abundance, thymidine incorporation into acid insoluble material, primary production (particulate and dissolved), chlorophyll, phaeopigments, total microbial ATP, particulate organic carbon and nitrogen, dissolved organic carbon, dissolved primary amines, and glucose and thymidine turnover rates. The data were analyzed by pairwise rank correlations with significance tested at the P<.005 level. Bacterial abundance and thymidine incorporation both declined progressively with increasing distance from shore (to 100 km); similar trends occurred for the phytoplankton, with several stations having subsurface maxima. Bacterial abundance, thymidine incorporation, and thymidine and glucose turnover rates were all significantly correlated to each other, suggesting they are comparable as relative measures of bacterial activity. Thymidine incorporation per cell, an indicator of specific growth rate, was not correlated to bacterial abundance, suggesting density independent specific growth rates. Bacterioplankton growth rate was evidently influenced more by the standing stock of phytoplankton than by the primary production of the phytoplankton. Thus, bacterial growth may possibly be stimulated by leakage of dissolved organic matter not so much from healthy photosynthesizing cells as from phytoplankton being disrupted and incompletely digested during predation by the zooplankton and nekton.     

Photosynthetic utilisation of carbon and light by two tropical seagrass species as measured in situ

In situ measurements of seagrass photosynthesis in relation to inorganic carbon (Ci) availability, increased pH and an inhibitor of extracellular carbonic anhydrase were made using an underwater pulse amplitude modulated (PAM) fluorometer. By combining the instrument with a specially designed Perspex chamber, we were able to alter the water surrounding a leaf without removing it from the growing plant. Responses to Ci within the chamber showed that subtidal plants of the seagrasses Cymodocea serrulata and Halophila ovalis had photosynthetic rates that were limited by the ambient Ci concentration depending on the irradiance that was available during short-term photosynthesis–irradiance trials. Relative electron transport rates (RETRs) at light saturation (up to 500 μ mol photons m⁻² s⁻¹) increased by 66–100% when the Ci concentration was increased from ca. 2.2 to 6.2 mM. On the other hand, intertidal plants of the same species exhibited a much lesser limitation of photosynthesis by Ci at any irradiance (up to 1500 μ mol photons m⁻² s⁻¹). Both species were able to use HCO⁻ ₃ efficiently, and there was stronger evidence for direct uptake of HCO⁻ ₃ rather than extracellular dehydration of HCO⁻ ₃ to CO₂ prior to Ci uptake. Subtidally, H. ovalis and C. serrulata grew to 10 and 12 m, respectively, where ambient irradiances were approximately 16 and 11% of those at the surface. Maximum RETRs (at light saturation) were lower for these deep-growing plants than for the intertidally growing ones. For both species, the onset of light saturation of photosynthesis (E _k) occurred at approximately 100 μ mol photons m⁻² s⁻¹ for the deep water populations, which was four and two times lower than for the shallow populations of C. serrulata and H. ovalis, respectively. This, and the differences in maximal photosynthetic rates (RETR _max), reflects an acclimation of the deep-growing populations to the lower light environment. The results presented here show that photosynthesis, as measured in situ, was limited by the availability of Ci for the deeper growing plants in Zanzibar, while the intertidally growing plants photosynthesised at close to Ci saturation. The latter result is contrary to previous conclusions regarding Ci limitations for these intertidal plants, and, in general, our findings highlight the need for performing similar experiments in situ rather than under laboratory conditions. Received: 4 April 2000 / Accepted: 31 August 2000     

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.     

Predator-inducible changes in blue mussels from the predator-free Baltic Sea

Blue mussels, Mytilus spp., have inhabited the brackish Baltic Sea, an environment lacking predatory crabs and starfish, for several thousand years. In this paper we examined whether Baltic Mytilus that were transplanted to the North Sea showed predator-inducible plasticity like their "marine" conspecifics. Our experiments showed that native North Sea Mytilus changed their morphology when exposed to waterborne scents from shore-crabs and starfish. These predators induced different kinds of changes, with emphasis on shell thickness and adductor muscle size, respectively. Baltic Mytilus responded in a similar way to crab scents, whereas starfish scents had a relatively weak effect on the morphology. Crab and starfish scent induced strengthening of the byssal attachment in North Sea Mytilus, with crabs providing more stimulation than starfish. Baltic mussels also improved the byssal attachment when exposed to either of the predators, but the attachment strength, as well as the response to crabs, were relatively weaker than that of North Sea mussels. We conclude that inducible plasticity still is present in Baltic Mytilus, despite their recent evolution in a predator-free environment. There is probably no strong selective pressure against inducible plasticity, but it could also be maintained in the population by gene flow from Mytilus in the adjoining North Sea. The question whether Baltic Mytilus are M. edulis or M. trossulus may also be relevant for the present results.     

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.     

活性有机碳含量在凋落物分解过程中的作用

土壤凋落物的分解不仅是生态系统养分循环的重要环节,也是生态系统碳释放源之一。将呼伦贝尔森林草原过渡带的草原凋落物、白桦林凋落物、落松林凋落物分别添加在棕色针叶林土里进行恒温培养,探讨了不同凋落物类型有机碳分解速率差异及其影响因子。结果表明:不同凋落物的有机碳矿化速率和矿化累积总量在分解初期不一致,但由高到低的次序均为:草原凋落物→白桦林凋落物→落叶松林凋落物,40d的有机碳矿化累积量分别为76.53、47.42、20.56mg/g。这主要与凋落物的化学性质有关,主要决定于凋落物中易被微生物分解的热水溶性有机碳含量和易分解有机物含量,而与凋落物的总有机碳含量、全氮含量、w(C)/w(N)比等关系不明显。     

Photoadaptation of zooxanthellae in the sponge Cliona vastifica from the Red Sea, as measured in situ

In the Red Sea, the zooxanthellate sponge Cliona vastifica (Hancock) is mainly present at >15 m depth or in shaded areas. To test whether its scarcity in unshaded areas of shallower waters is linked to the functional inefficiency of its photosymbionts at high irradiances, sponges were transferred from 30 m to a six times higher light regime at 12 m depth, and then returned to their original location. During this time, photosynthetic responses to irradiance were measured as rapid light curves (RLCs) in situ by pulse amplitude modulated (PAM) fluorometry using a portable underwater device, and samples were taken for microscopic determinations of zooxanthellar abundance. The zooxanthellae harboured by this sponge adapted to the higher irradiance at 12 m by increasing both their light saturation points and relative photosynthetic electron transport rates (ETRs). The ETRs at light saturation increased almost fourfold within 15–20 days of transfer to the shallower water, and decreased back to almost their original values after the sponges were returned to 30 m depth. This, as well as the fact that the photosynthetic light responses within an individual sponge were in accordance with the irradiance incident to specific surfaces, shows that these photosymbionts are highly adaptable to various irradiances. There was no significant change in the number of zooxanthellae per sponge area throughout these experiments, and the different photosynthetic responses were likely due to adaptations of the photosynthetic apparatus within each zooxanthella. In conclusion, it seems that parameters other than the hypothesised inability of the photosymbionts to adapt adequately to high light conditions are the cause of C. vastifica's rareness in unshaded shallow areas of the Red Sea. Received: 25 April 2000 / Accepted: 13 October 2000     

Occurrence of the semi-catadromous European eel Anguilla anguilla in the Baltic Sea

Strontium (Sr) and calcium (Ca) contents in the otoliths of yellow and silver European eels [Anguilla anguilla (L.)] collected from coastal waters of the Baltic Sea and a freshwater lake in Sweden were examined by wavelength dispersive X-ray spectrometry with an electron microprobe. The mean Sr/Ca ratios from elver check to otolith edge were significantly higher for the eels from coastal waters (5.39 ± 1.09‰) than for those from the lake (0.71 ± 0.89‰). The evidence indicates that European eels in the Baltic Sea do not necessarily migrate into freshwater streams during the growth phase. Received: 30 September 1999 / Accepted: 6 April 2000     

Metals in Zooplankton from the Baltic Sea, 1980-84

Between 1980 and 1984, plankton was collected for metal analysis during four expeditions in the Baltic Sea. for comparison, samples from adjacent areas of the northeastern Atlantic Ocean were also taken. the mixed net-plankton samples were analyzed by AAS for metals (Al, Cd, Co, Cr, Cu. Fe, Mn, Ni, Pb, and Zn). the results are discussed with regard to spatial and temporal trends and for comparison with data from other authors. Correlations among the contents of the different metals and between the metal contents and 'external factors' including salinity, season, percentage of co-collected phytoplankton, and concentration of dissolved and particulate metals in the water are considered.

There is a tendency for higher metal contents in plankton from the brackish Baltic Sea (Al, Mn and Hg), while other metals (e.g. Cd) show higher levels in samples taken from the marine environment. Except for mercury, no clear correlations could be found between the metal content in plankton and the dissolved concentration of the same metal in the ambient water. Otherwise, the contents of aluminium, iron, manganese and zinc in the suspended particulate matter and in the plankton seem to be partly related to each other.     

Stoichiometry of mesozooplankton in N- and P-limited areas of the Baltic Sea

The Baltic Sea is a very suitable site for stoichiometric studies, since its subbasins differ in their concentration of elemental components, and primary production can therefore be either nitrogen or phosphorus limited. To reveal if the nutrient limitation of mesozooplankton mirrors that of the primary producers, carbon, nitrogen and phosphorus content of both seston and grazers (Acartia sp., Centropages hamatus, Daphnia cristata, Eurytemora affinis, Limnocalanus macrurus, Temora longicornis) were measured in midsummer in the Baltic proper, the Gulf of Finland and the Gulf of Bothnia. The mineral ratios of the different taxa were equal, apart from L. macrurus with notably higher C:P and N:P ratios. Molar C:N ratios were relatively stable (5.1-6.3), whereas C:P and N:P ratios fluctuated more (41-144 and 6.6-24). However, zooplankton elemental composition and limitation did not depend on the limiting nutrient of the phytoplankton, the seston mineral ratio or the sea area. Both the seston-zooplankton elemental imbalance and the food threshold ratio indicated phosphorus limitation of most of the grazers. While L. macrurus may be C or N limited, the possible P deficiency of the other studied taxa suggests that the Baltic Sea zooplankton may act as a potential phosphorus sink, as the freshwater secondary producers do.     

Contrasting population structures in two sympatric fishes in the Baltic Sea basin

Detailed multispecies studies on the patterns of genetic variability and differentiation in marine environments are still rare. Using mitochondrial and nuclear genetic markers, we compared genetic variability and population structuring of threespine (Gasterosteus aculeatus) and ninespine (Pungitius pungitius) sticklebacks from the same eleven marine and six freshwater locations within the Baltic Sea basin. Analyses of both marker types revealed a significantly lower degree of genetic structuring in both marine and freshwater populations of threespine than those ninespine sticklebacks. Isolation-by-distance (IBD) was detected across the marine populations in both species, suggesting spatially limited gene flow. However, the levels of genetic diversity and differentiation across the localities were uncorrelated between the two species in both marine and freshwater environments. Accordingly, estimates of effective population sizes were larger and migration rates were higher for three- than for ninespine sticklebacks. Hence, ninespine stickleback populations from the Baltic Sea basin appear to be subject to stronger genetic drift than sympatric threespine sticklebacks, and the proximate reason for this difference is likely to be found from autecological differences between the two species. In accordance with the earlier studies, genetic variability was higher and the degree of genetic differentiation was lower in marine than in freshwater populations in both species.     

Contribution of salps to carbon flux of marginal ice zone of the Lazarev Sea, southern ocean

In order to estimate the in situ grazing rates of Salpa thompsoni and their implications for the development of phytoplankton blooms and for the sequestration of biogenic carbon in the high Antarctic, a repeat-grid survey and drogue study were carried out in the Lazarev Sea during austral summer of 1994/1995 (December/January). Exceptionally high grazing rates were measured for S. thompsoni at the onset of a phytoplankton bloom (0.2 to 0.8 μg chlorophyll a l⁻¹) in December 1994, with up to ≃160 μg of plant pigments consumed by an individual salp of 7 to 10 cm length per day. Dense salp swarms extended throughout the marginal ice zone, consuming up to 108% of daily phytoplankton production and 21% of the total chlorophyll a stock. Due to the much faster sinking rates and higher carbon content of salp faecal pellets, the efficiency of downward carbon flux through salps is much higher than through the other major grazers, krill and copepods. S. thompsoni can thus export large amounts of biogenic carbon from the euphotic zone to the deep ocean. With the observed ingestion rates during December 1994, this flux could have attained levels of up to 88 mg C m⁻² d⁻¹, accounting for the bulk of the vertical transport of carbon in the Lazarev Sea. However, in January 1995, when phytoplankton concentrations exceeded a threshold level of 1.0 to 1.5 μg chlorophyll a l⁻¹, salps experienced a drastic reduction in their feeding efficiency, possibly as a result of clogging of their filtering apparatus. This triggered a dramatic reversal in the relationship, during which a dense phytoplankton bloom developed in conjunction with the collapse of the salp population. Increases in the biomass and geographic range of the tunicate S. thompsoni have occurred in several areas of the southern ocean, often in parallel with a rise in sea-surface temperature during sub-decadal periods of warming anomalies. Received: 10 August 1997 / Accepted: 21 October 1997     

Floristic and physiological differences between the shallow and the deep nanophytoplankton community in the euphotic zone of the open tropical Atlantic revealed by HPLC analysis of pigments

Suspended matter sampled in 1982 in the North Equatorial Current, in the open Atlantic to the west of West Africa, was analyzed by high performance liquid chromatography. The pigment fingerprint of samples taken in the surface mixed layer was dominated by zeaxanthin and chlorophyll a, in agreement with observed dominance of coccoid cyanobacteria. Near the bottom of the euphotic zone the fingerprint was more complicated, with a sharp transition at the depth of the deep chlorophyll maximum layer to dominance of chlorophyll b, 19-hexanoyloxyfucoxanthin and an unknown fucoxanthin derivative in the lower part of this layer; this fingerprint suggests dominance of eukaryotes (green algae, Prymnesiophyceae and Chrysophyceae) at depth. Up to 90% of the chl a was contained in particles smaller than 8 m, and in the surface mixed layer even more than 50% in particles smaller than 1 m. The high concentration of zeaxanthin relative to chl a near the surface suggests adaptation of the cyanobacteria to exposure to high irradiance. Evidence of this adaptation was the very high specific phytoplankton growth rate between sunrise and sunset (=0.16 h^-1), measured by recording ¹⁴C incorporation into organic carbon and into chl a carbon after isolation of the latter by HPLC. The high concentration of chl b relative to chl a at depth was possibly caused by shade-adapted green algae containing more chl b than chl a. The specific growth rate of the deep shade community was low (<0.04 h^-1), yet net primary production, calculated on the basis of chl a increase during incubation, was greatest at depth.