Carbon, nitrogen, oxygen and sulfide budgets in the Black Sea: A biogeochemical model of the whole water column coupling the oxic and anoxic parts

Carbon, nitrogen, oxygen and sulfide budgets are derived for the Black Sea water column from a coupled physical-biogeochemical model. The model is applied in the deep part of the sea and simulates processes over the whole water column including the anoxic layer that extends from ?115 m to the bottom (?2000 m). The biogeochemical model involves a refined representation of the Black Sea foodweb from bacteria to gelatinous carnivores. It includes notably a series of biogeochemical processes typical for oxygen deficient conditions with, for instance, bacterial respiration using different types of oxidants (i.e denitrification, sulfate reduction), the lower efficiency of detritus degradation, the ANAMMOX (ANaerobic AMMonium OXidation) process and the occurrence of particular redox reactions. The model has been calibrated and validated against all available data gathered in the Black Sea TU Ocean Base and this exercise is described in Gregoire et al. (2008). In the present paper, we focus on the biogeochemical flows produced by the model and we compare model estimations with the measurements performed during the R.V. KNORR expedition conducted in the Black Sea from April to July 1988 (Murray and the Black Sea Knorr Expedition, 1991). Model estimations of hydrogen sulfide oxidation, metal sulfide precipitation, hydrogen sulfide formation in the sediments and water column, export flux to the anoxic layer and to the sediments, denitrification, primary and bacterial production are in the range of field observations.With a simulated Gross Primary Production (GPP) of 7.9 mol C m⁻² year⁻¹ and a Community Respiration (CR) of 6.3 mol C m⁻² year⁻¹, the system is net autotrophic with a Net Community Production (NCP) of 1.6 mol C m⁻² year⁻¹. This NCP corresponds to 20% of the GPP and is exported to the anoxic layer. In order to model Particulate Organic Matter (POM) fluxes to the bottom and hydrogen sulfide profiles in agreement with in situ observations, we have to consider that the degradation of POM in anoxic conditions is less efficient that in oxygenated waters as it has often been observed (see discussion in Hedges et al., 1999). The vertical POM profile produced by the model can be fitted to the classic power function describing the oceanic carbon rate (C_R=Z^−α) using an attenuation coefficient α of 0.36 which is the value proposed for another anoxic environment (i.e. the Mexico Margin) by Devol and Hartnett (2001). Due to the lower efficiency of detritus degradation in anoxic conditions and to the aggregation of particles that enhanced the sinking, an important part of the export to the anoxic layer (i.e. 33%, 0.52 mol C m⁻² year⁻¹) escapes remineralization in the water column and reaches the sediments. Therefore, sediments are active sites of sulfide production contributing to 26% of the total sulfide production.In the upper layer, the oxygen dynamics is mainly governed by photosynthesis and respiration processes as well as by air-sea exchanges. ?71% of the oxygen produced by phytoplankton (photosynthesis+nitrate reduction) is lost through respiration, ?21% by outgasing to the atmosphere, ?5% through nitrification and only ?2% in the oxidation of reduced components (e.g. Mn²⁺, Fe²⁺, H₂S).The model estimates the amount of nitrogen lost through denitrification at 307 mmol N m⁻² year⁻¹ that can be partitioned into a loss of ?55% through the use of nitrate for the oxidation of detritus in low oxygen conditions, ?40% in the ANAMMOX process and the remaining ?5% in the oxidation of reduced substances by nitrate.In agreement with data analysis performed on long time series collected since the 1960s (Konovalov and Murray, 2001), the sulfide and nitrogen budgets established for the anoxic layer are not balanced in response to the enhanced particle fluxes induced by eutrophication: the NH₄ and H₂S concentrations increase.     

Sulfide-oxidizing bacteria in the burrowing echinoid,Echinocardium cordatum (Echinodermata)

Symbiotic filamentous bacteria thrive in the intestinal caecum of the deposit-feeding echinoid Echinocardium cordatum. Specimens of E. cordatum were collected at Wimereux (Nord Pas-de-Calais, France) in 1991. Their symbiotic bacteria build nodules by forming multilayered mats around detrital particles that enter the caecum. The morphological features of the bacteria are those of Thiothrix, a sulfide-oxidizing genus. The filaments, which may form rosettes, are sheathed and made by a succession of hundreds of rod-shaped bacteria which store elemental sulfur in the presence of external sulfide. Live bacteria are restricted to the outer layers of the nodules. Their sulfide-oxidizing activity was investigated, using a Biological Oxygen Monitor, by measuring the O₂-consumption when reduced sulfur compounds are provided. They oxidize thiosulfate and sulfide. Optimal sulfide oxidation occurs at intermediary pO₂ (100 to 160 M O₂l^-1). Spectrophotometry has confirmed that the sulfur content of the filamentous symbiotic sulfideoxidizing bacteria depends on the presence of external sulfide. This is the first report of symbiotic intradigestive Thiothrixspp.-like bacteria; it lengthens the list of symbioses between sulfide-oxidizing bacteria and invertebrates from sulfide-rich habitats.     

Oxygen dependent sulfide detoxification in the lugwormArenicola marina

The lugwormArenicola marina L. oxidizes entering sulfide to thiosulfate. After 8 h of normoxic incubations with sulfide concentrations of 0.2 to 1.0 mmoll^-1 thiosulfate in the coelomic fluid amounted up to about 4 mmoll^-1 whereas sulfite concentrations were 100-fold lower and no accumulation of sulfate in the coelomic fluid was found. The sulfide oxidation was highly oxygen dependent. An increase of oxygen partial pressure ( ) in the medium was followed by enhanced thiosulfate production and by a decrease of sulfide concentration in the coelomic fluid. Under normoxia, the sulfide oxidation rate was sufficient to compensate the influx of sulfide into the coelomic fluid when the sulfide concentration in the medium was below 0.33 mmoll^-1. When external sulfide was raised beyond this level, sulfide up to 5 moll^-1 in the coelomic fluid appeared. Succinate in the body wall tissue was low as long as no sulfide appeared in the coelomic fluid, indicating the maintenance of an aerobic metabolism. The oxidation of sulfide to thiosulfate was localized in the mitochondria of the body wall tissue. The oxygen consumption of mitochondria was stimulated by the addition of sulfide. The mitochondrial sulfide oxidation rate depended on the amount of mitochondrial protein and followed a Michaelis-Menten kinetic. An apparentK _m of 0.68±0.29 moll^-1 and aV _max of 41.9±22.3 nmol min^-1 mg^-1 protein was calculated. Sulfide was stoichiometrically oxidized to thiosulfate with 1 mol sulfide consuming 1 mol oxygen. Sulfide oxidation was not inhibited by sulfide concentrations as high as 100 moll^-1. At low concentrations of cyanide or azide, when respiration without sulfide was already inhibited, sulfide oxidation could still be stimulated, tentatively indicating the existence of an alternative terminal oxidase. Specimens examined in the present study were collected near St. Pol de Leon, France, from 1989 to 1992.     

The bacterial symbiont from the hydrothermal vent tubewormRiftia pachyptila is a sulfide specialist

We have investigated the metabolic adaptations of the chemolithotrophic bacterial symbionts ofRiftia pachyptila. Specimens of the tubeworm were collected by submersible from depths of 2600 m at 13°N on the East Pacific Rise in 1987, and 2450 m at the Galápagos Rift in 1988. Isolated bacteria utilize sulfide, but not thiosulfate or sulfite, as their sole reduced-sulfur energy source. The bacteria rapidly oxidize a wide range of sulfide concentrations (5µM to 2 mM), with maximal respiration rates at concentrations >1 mM, and unlike many sulfur-oxidizing bacteria, show no inhibition in oxygen consumption at sulfide concentrations up to 2 mM. Incubations of freshly homogenized trophosome tissue or isolated bacteria with sodium [³⁵S] sulfide and subsequent analysis of sulfur products by high-performance liquid chromatography and flow-through scintillation counting showed that sulfide disappeared almost completely within 1 min. Both soluble and insoluble products of sulfide oxidation were produced. The soluble fraction contained sulfate and polysulfides, with no thiosulfate produced. However, the majority of the radioactivity was in the water-insoluble fraction, mostly as elemental sulfur. Whole-worm experiments under pressure showed a rapid removal of³⁵S-sulfide from the incubation water, with sulfide, sulfate, and polysulfides appearing in the blood within 4 h. There was no utilization of thiosulfate by the whole worms, freshly homogenized trophosome tissue, or isolated bacteria.     

Contrasting effects of sulfide and thiosulfate on symbiotic CO2-assimilation of Phallodrilus leukodermatus (Annelida)

Experiments were conducted in order to specify the reductants responsible for the carbon dioxide fixation of the symbiotic sulfur bacteria in the gutless marine oligochaete Phallodrilus leukodermatus (Annelida) from shallow calcareous sediments in Bermuda. Carbon dioxide-uptake rates were suppressed by S⁼ and stimulated by S₂O₃ ⁼. Individuals which hosted bacteria containing reserve energy substances maintained a high short-term CO₂-uptake activity, while bacteria in worm homogenates and in worms treated with an antibiotic (Baypen) did not show any significant metabolic activity. Absolute uptake rates in P. leukodermatus were usually considerably higher than those reported for other animals harbouring prokaryotic sulfuroxidizing symbionts. Utilization of thiosulfate rather than sulfide is compatible with the preferred occurrence of the worms around the redox discontinuity layer and has been confirmed in other thiobiotic animals. Sulfur stored in the symbiotic bacteria appears to be oxidized to sulfate and be excreted when the worms are held under energy-limited conditions. The data emphasize the complexity of the possible metabolic pathways involved in the oxidation of reduced-sulfur compounds by bacterial symbionts in marine invertebrates.     

Effects of sulfide exposure history and hemolymph thiosulfate on oxygen-consumption rates and regulation in the hydrothermal vent crab Bythograea thermydron

The hydrothermal vent crab Bythograea thermydron is exposed to high environmental concentrations of sulfide and low levels of oxygen for extended periods of time. It has previously been shown that hydrogen sulfide is oxidized to the relatively non-toxic thiosulfate (S₂O ₃ ^2– ), which accumulates in the hemolymph. Hemolymph thiosulfate levels in freshly captured crabs vary significantly among crabs from different hydrothermal vent sites as well as between crabs from different microhabitats within the same site. Hemolymph thiosulfate concentrations were not significantly different between crabs captured at the same site 6 mo apart. Hemolymph thiosulfate concentrations ranged from 66 mol 1^–1 in a crab captured at a site with relatively low sulfide venting, to 3206 mol 1^–1 in an individual that was netted from an active smoker vent with much higher sulfide exposure. The differences in hemolymph thiosulfate between sites and the stability of hemolymph thiosulfate in crabs captured at the same site at different times suggest that sulfide exposure is significantly different between sites and that this exposure may not vary significantly over the course of a few months. B. thermydron experimentally exposed to sulfide had high levels of thiosulfate in their hemolymph and increased abilities to regulate oxygen consumption in conditions of low oxygen. This enhancement of regulatory abilities suggests that the previously demonstrated increased hemocyaninoxygen (Hc–O₂) affinity due to elevated thiosulfate may be adaptive in vivo. Average oxygen-consumption rates were much higher in crabs experimentally exposed to sulfide than in unexposed crabs. Crabs injected with isosmotic thiosulfate did not have increased oxygen-consumption rates as did the sulfide-exposed individuals, but did show a similar reduction in P _c (the critical partial pressure of oxygen at which crabs can no longer regulate oxygen consumption). This suggests that it is the sulfide exposure and/or detoxification rather than the elimination of thiosulfate that causes the increase in metabolic rate. Thiosulfate diffuses into dead crabs and into live crabs exposed to 15 mmol S₂O ₃ ^2- l^–1, indicating substantial permeability, and yet live crabs are able to eliminate thiosulfate when incubated in sea water containing 1.5 mmol S₂O ₃ ^2- l^–1, suggesting a process that has an active component.     

Polychlorinated Biphenyls (Pcbs) Pollution in Sea of Japan

Abstract

Deep waters of the Sea of Japan and surface waters of the Pacific Coast of Honshu and the northeast Sea of Japan were analysed for polychlorinated biphenyls (PCBs) pollution. the ΣPCB concentrations in solution in the Sea of Japan (50–3000 m) were between 140 and 1230 fg dm_?3. the space-integrated surface water concentration near the Pacific coast of northern Honshu was 140 fg dm_?3and for the surface water of the Sea of Japan was 230 fg dm_?3. Based on these analyses four water masses were deduced in the Sea of Japan during the summer months. It is shown for the first time in the Sea of Japan that polychlorinated biphenyls are excellent chemical indicators of not only the anthropogenic pollution, but also water masses.     

Activities of sulfur-oxidizing bacteria at the 21°N East Pacific Rise vent site

The incorporation of inorganic and organic carbon into cell material, as well as the activities of carboxylating enzymes (ribulose bisphosphate carboxylase and phosphoenol pyruvate carboxylase), were measured in waters emitted from warm vents at the 21°N ocean spreading site (depth 2 600 m) of the East Pacific Rise. Both obligately and facultatively chemoautotrophic bacteria were present and comprised a significant but variable portion of the total microbial population as indicated by comparisons of microscopic cell counts with liquid enrichments and colony counts on media containing reduced sulfur compounds. The proportion of chemoautotrophic, sulfur-oxidizing bacteria maximally reached 79% of the total microbial population based on ribulose bisphosphate carboxylase activity. Variability of chemoautotrophic activity occurred between vents at different locations, but was also observed in one individual vent. Maximum rates of CO₂ incorporation in warm vent water were similar to levels measured previously in the O₂/H₂S interface of the Black Sea. In shipboard experiments, these rates were virtually unaffected by in-situ pressures (ca 260 atm). Rate measurements at various temperatures, as well as the observed mesophilic characteristics of all isolates obtained, suggest that the microbial, chemoautotrophic activity decreases rapidly as freshly emitted vent water is diluted with cold, ambient, deep-sea water.Contribution No. 6071 of the Woods Hole Oceanographic InstitutionContribution No. 1708 of the Center for Environmental and Estuarine Studies of the University of Maryland     

The gill symbiont of the hydrothermal vent mussel Bathymodiolus thermophilus is a psychrophilic,chemoautotrophic, sulfur bacterium

Certain hydrothermal vent invertebrates, e.g. Riftia pachyptila and Calyptogena magnifica, are clearly established as harboring dense populations of chemoautotrophic sulfur bacteria in specialized tissues. By contrast, the physiological characteristics of the abundant intracellular gill symbiont of the vent mussel Bathymodiolus thermophilus have been questioned. The low activities of enzymes diagnostic for CO₂ fixation (Calvin cycle) and for sulfur-driven energy generation, as measured by other investigators, have been attributed to bacterial contamination of the gill surface. Based on research at the Galápagos Rift hydrothermal vents in 1988 and subsequent laboratory experiments, the current study confirms that the B. thermophilus symbiont is a psychrophile for which thiosulfate and sulfide stimulate CO₂ fixation. It strongly indicates that the symbiont is a chemoautotroph by establishing the following: (1) Sulfide and thiosulfate can stimulate CO₂ fixation by partially purified symbionts by up to 43-fold and 120-fold, respectively; (2) the ribulose-1,5-bisphosphate carboxylase/oxygenase activity of the symbiont is sufficient to account for its sulfide- or thiosulfate-stimulated CO₂ incorporation; (3) the symbiont's molar growth yield on thiosulfate, as judged by CO₂ incorporation, is indistinguishable from that of free-living chemoautotrophs. Due to the high protein-degrading activity of B. thermophilus gill lysate, it is also suggested that host lysis of symbionts plays a more important role in the nutrition of the vent mussel than in R. pachyptila or C. magnifica, for which no comparable protein-degrading activity was found.     

Mesozooplankton communities in the Aegean and Black Seas: a comparative study

An attempt was made for a comparative study of the mesozooplankton communities between the Aegean and Black Seas. These areas are in connected through the Sea of Marmara, the Bosporus and the Dardanelles straits. The comparison was mainly based on data collected both in coastal and offshore waters, by the Hellenic Centre for Marine Research (Greece), the Shirshov Institute of Oceanology (Russia), the Institute of Biology of Southern Seas (Ukraine) and the Institute of Oceanology (Bulgaria). Our data revealed important differentiation in terms of species composition and interannual fluctuations of zooplankton quantity. Among the common copepod species, those presenting an eurytherm and/or euryhaline character (e.g. Acartia clausi, Paracalanus parvus, Penilia avirostris, Podon polyphemoides) were found dominant in the Black Sea and common in the Aegean Sea. Decreased dissimilarity of species composition seems to exist between the North Aegean Sea and the Black Sea, as well as between the coastal areas of the Black Sea and those affected by pollution in the Aegean Sea. A common feature of both seas is the existence of spatial differentiation in terms of the quantity of zooplankters due to anthropogenic and/or natural influences. The pattern of this differentiation seems to vary interannually in the Black Sea, depending either on the abundance of mesozooplankton predators (indigenous or invasive species) or on global climatic changes. Although the Black Sea is generally richer in zooplankton than the Aegean Sea, the significant decrease in the quantity of zooplankton from the early 1980s to the mid-1990s, due to the above factors, resulted in quasi-similar values in both areas. Black Sea mesozooplankton has revealed strong interannual variability both quantitatively and qualitatively, whereas no significant variability was detected in the Aegean Sea. This difference could be attributed both to variant anthropogenic forcing on these ecosystems and to the sensitive character of the semi-enclosed Black Sea.Communicated by M.E. Vinogradov, Moscow     

Multiple trophic resources for a chemoautotrophic community at a cold water brine seep at the base of the Florida Escarpment

The biological community that surrounds the hypersaline cold water brine seeps at the base of the Florida Escarpment is dominated by two macrofaunal species: an undescribed bivalve of the family Mytilidac and a vestimentiferan worm, Escarpia laminata. These animals are apparently supported by the chemoautotrophic fixation of carbon via bacterial endosymbionts. Water column and sediment data indicate that high levels of both sulfide and methane are present in surface sediments around the animals but absent from overlying waters. Stable isotopic analyses of pore water indicate that there are two sources of sulfide: the first is geothermal sulfide carried in groundwater leaching from the base of the escarpment, and the second is microbial sulfide produced in situ. The vestimentiferan E. laminata, and the mytilid bivalve (seep mussel) live contiguously but rely on different substrates for chemoautotrophy. Enzyme assays, patterns of elemental sulfur storage and stable isotopic analyses indicate that E. laminata relies on sulfide oxidation and the seep mussel on methane oxidation for growth.     

New denitrifying bacteria isolated from Red Sea sediments

Sediments of various newly discovered deeps in the Red Sea were analyzed for the occurrence of denitrifying bacteria. The samples were collected in 1972 during the Valdivia cruises. Among the 27 different samples investigated, 17 revealed both coccoid and rod-shaped bacteria when enriched in complex nutrient broth (with 10% NaCl). Denitrifiers were recorded abundantly in the sediments, their population decreasing from some 10⁶/g in the surface material to only a few in the subsediment. A total of 16 pure cultures of denitrifying bacteria were isolated from the Suakin-and Thetis-deeps and studied morphologically, physiologically and biochemically. Genetics (molar percentage of guanine plus cytosine, % GC) and numerical taxonomy were included to reveal relationships and improve taxonomic classification. Fifteen isolates were described as Gram-negative, aerobic and facultative anaerobic (with NO ₃ ^- as H⁺-acceptor), polarly flagellated rods (Pseudomonas spp.); one was an inmotile, Gram-positive, facultative anaerobic coccus. None of the 15 Pseudomonas-isolates could be identified with one of the denitrifying species so far described and recognized. The strains should be regarded as hitherto undescribed denitrifying marine bacteria.Supported by the Bundesministerium für Forschung und Technologie, Bonn, Germany (FRG).     

Etude des eaux rouges dues à la prolifération des bactéries photosynthétiques sulfo-oxydantes dans l'étang du Prévost,lagune saumâtre méditerranéenne

Dystrophic crises or malaígues with red water, were observed in summer, in Mediterranean brackish lagoons near Montpellier, France. During July 1976, the photosynthetic sulfide-oxidizing bacteria causing this phenomenon were isolated from the Prévost lagoon. The most important genera isolated from this red water wereChromatium, Thiocystis andThiocapsa (Chromatiaceae): the predominant species wasThiocapsa roseopersicina. Water and sediment samples from the same lagoon were collected during the winter season (October 1976 and January 1977) and were enriched with organic matter. The samples were incubated in aquaria under artificial light (800 to 1000 lux) at room temperature (ca. 22°C). Eight to 10 days later, red water developed which was similar to that observedin situ in summer. These red waters develop under conditions of anaerobiosis and H₂S production. Such conditions are necessary for the growth of Chromatiaceae. Excessive deposists of organic matter in the lagoon water lead to increased oxygen consumption and overproduction of hydrogen sulfide, which can be oxidized by photosynthetic sulfideoxidizing bacteria. This bacteria bloom (causing the red water) removes the hydrogen sulfide, thus re-establishing aerobiotic conditions in the water of the Prévost lagoon.     

Manganese in natural waters II. Evidence for microbiological oxidation of Mn (II)

The oxidation of Mn(II) to Mn(IV) in natural waters has been investigated. Various hypotheses have been put forward for the mechanism of oxidation of Mn(II) of which only biological oxidation was found to occur in the aquatic system studied. The effect of adsorption, usually associated with the oxidation of Mn(II) was not observed in our study. The oxidation kinetics can be divided into three periods. The first period is the induction time corresponding to the growth of microorganisms that act as oxidising agents for Mn² _aq ⁺. During this stage Mn² _aq ⁺ remains constant. This period is followed by a decrease in Mn² _aq ⁺ concentration following zero order rate law. Finally in the third period the decrease in Mn² _aq ⁺ follows a first order rate law. The microorganisms acting as oxidising agents for Mn(II) have been identified to be filamentous Gallionella type bacteria and Leptothrix discophora.     

Effect of inorganic phosphorus on the growth rate of diatoms

The effect is studied of different inorganic phosphorus concentrations on changes in growth rates of 7 diatom species isolated from the plankton of the Black Sea. All species examined increase their cell-division rate with increasing phosphorus concentration in the medium. The phosphorus concentrations have been determined above which division rate is not limited by phosphorus content in the medium. The non-limiting concentration amounts to V/V _m =0.9. In the species studied, non-limiting concentrations ranged from 1 to 30 g P/1. The highest values were obtained for relatively large-sized species. The ratio of cell surface to volume tended to be inversely related to the growth-rate-limiting concentration. Minimal values of phosphorus content in cells have been calculated to range from 0.6 to 4.0·10^–17 g.at–P/^–3. Based on a comparison of phosphate levels in the Black Sea with experimentally-derived rate-limiting concentrations, it is concluded that phosphorus does not limit the reproduction rate of phytoplankton in the more productive regions of the Black Sea or in the lower strata of the euphotic zone.     

Sulfide tolerance and detoxification in shallow-water marine fishes

Hydrogen sulfide is a potent inhibitor of aerobic respiration. Sulfide is produced in sediments, and many species of fish live in association with the bottom. Tolerance tests, enzyme assays, and chromatography of sulfur compounds in thirteen species of shallow-water marine fishes (collected in San Diego, California, USA in 1987–1988) indicate adaptations to sulfide that vary with habitat and lifestyle. Tidal-marsh inhabitants, like Gillichthys mirabilis and Fundulus parvipinnis, have higher tolerance to sulfide (96 h LC₅₀ at 525 to 700 M) relative to outer-bay and open-coast inhabitants (surviving <12 h at much lower concentrations). The cytochrome c oxidase of all species shows high activity and susceptibility to sulfide poisoning, with 50% inhibition at 30 to 500 nM in various tissues. The two marsh species are able to survive at sulfide concentrations already inhibitory to their cytochrome c oxidase and fatal to other species. All species detoxify sulfide by oxidizing it to thiosulfate. All have sulfide-oxidizing activity in the blood, spleen, kidney, liver and gills, which correlates significantly with heme content. Thiosulfate appears in the tissues of sulfide-exposed fish and builds up to high concentrations (up to 2 mM) with stronger and longer exposure. Unexposed fish contain little or no thiosulfate. Sulfide is barely detectable in the tissues, even in high-sulfide exposure tests. We suggest that fish blood, in having high sulfide-oxidizing activity and no cytochrome c oxidase, can act as a short-term first line of defense against sulfide, and thus minimize the amount that reaches the vital organs. The results of this study indicate that sulfide is a significant environmental factor influencing the ecological distribution of marine fishes.     

Synchronous recruitment and growth pattern of planktonic larvae of the amphioxus Branchiostoma belcheri in the Seto Inland Sea, Japan

Planktonic larvae of the amphioxus Branchiostoma belcheri were collected in the western part of the Seto Inland Sea, Japan, in order to document their seasonal occurrence, recruitment, and growth patterns. The larvae appeared from mid-July to early October and their size ranged from 0.6 to 5.5 mm. Three distinct cohorts were observed during the summers, indicating multiple, synchronous spawning within the population. The length increment pattern of a cohort demonstrates slow growth for at least a week during the early larval stage. July 10, 20, and 30 are the inferred dates when spawnings started to produce the three cohorts in 2001. Favorable temperature range for spawning was 21–23°C. Bias in population structure due to advection can be assumed to be negligible; therefore the length increment of the cohort, about 0.05 mm day⁻¹, is regarded as the larval growth rate after the initial slow-growth stage. The planktonic stage of the first cohort in 2001 is estimated to have lasted around 55 days. The relative proportions of the second and third cohorts in samples from surface and mid-depth waters suggest downward migration of the planktonic larvae, as they age.     

Interrelationships between sulfate reducing and methane producing bacteria in coastal sediments with intense sulfide production

Coastal sediments receiving different amounts of organic carbon through sedimentation were investigated with respect to sulfate reduction and methanogenic activity. Sampling was carried out at sediment temperatures of 7° and 15°C. Sulfate-reducing and methanogenic bacteria were found at all depths. Sulfate reduction decreased with depth and the highest sulfide concentrations were found a few centimeters below the sediment surface, up to 15 mM at 15°C and pH 7.1. In the same segments a maximum in the methane concentration was also found, 0.91 mM. The high sulfide concentration inhibited the methane formation from acetate but not from carbon dioxide. In the organic rich sediment sulfate reduction was limited by the diffusion of SO ₄ ⁺ into the sediment and methane production from acetate by sulfide diffusion out of the sediment. When electron acceptor concentration limits sulfate reduction, thermodynamic calculations show that the utilization of electron donors more reduced than acetate is favored. In the sediment with the high carbon-input, acetate predominated at 15°C whereas in the low carbon-input sediment hardly any short chain organic acids were detected. The possibility of a shift in sulfate reduction from acetate oxidation to acetate production is discussed.     

Microbial activities in the emitted hydrothermal waters of the Galápagos rift vents

In-situ measurement of chemolithotrophic and some heterotrophic microbial activities were made in the immediate vicinity of actively discharging hydrothermal vents of the Galápagos Rift region at depths of 2 500 to 2 600 m. The CO₂-assimilation or chemosynthesis productivity in the emitted vent waters, freshly mixed with oxygenated ambient seawater of 2°C, was minor compared to the bacterial biomass produced within the subsurface vent system prior to emission. Uptake of acetate and glucose indicated the presence of mixotrophic or facultatively chemolithotrophic bacteria in the emitted vent waters in agreement with isolations. Demonstration of ribulose bisphosphate carboxylase and phosphoenol pyruvate carboxylase in cultures of thiobacilli isolated from these vent water supports the notion that chemoautotrophic sulfur-oxidizing bacteria are one of the sources of primary production in the form of particulate organic carbon for filtering organisms in the deep sea hydrothermal environment. The rates of bacterial metabolic activities in emitted vent water are too low for the amount of invertebrate biomass and the rate of its growth and maintenance. Therefore, the larger portion of chemosynthetic sustenance of deep sea vent ecosystems appears to be based on symbiotic associations between bacteria and invertebrates and on surface attached bacteria.     

Geographical differences in early growth of walleye pollock Theragra chalcogramma,estimated by back-calculation of otolith daily growth increments

The relationship between otolith length and body length (total length) was examined in walleye pollock Theragra chalcogramma (Pallas), collected from Pacific Ocean waters adjacent to Hokkaido, Japan, and in larvae reared from fertilized eggs. A linear relationship was found between log-transformed body length and otolith length data with two inflection points at ca. 11 mm and 100 mm total length. This relationship was found to be applicable also to samples from the Japan Sea, Bering Sea and Okhotsk Sea. The early growth pattern estimated by back-calculation of otolith increments of 1-yr fish from Funka (Uchiura) Bay (Hokkaido) accorded with that obtained from size-at-age data of 0 yr fish collected in the preceding year. Differences were found in comparisons of the back-calculated early growth pattern between samples from waters adjacent to Hokkaido, the Pacific Ocean, the Japan Sea and the Okhotsk Sea. The growth curve of the Okhotsk samples was markedly different from others, showing rapid initial growth up to about 100 d after hatching and attaining a small body size in the first year.