Nitrogen assimilation by phytoplankton and other microorganisms in the surface waters of the central North Pacific Ocean

Assimilation rates of ¹⁵N-labelled ammonium, urea, and nitrate by plankton in the upper euphotic zone were measured in 2 summer, 2 winter, and 1 spring cruise in the central North Pacific Ocean. Average rates of ammonium plus urea assimilation could not be determined precisely, but were estimated to be 7 to 25 g-at. N m^-3 day^-1. Indirect evidence suggested that non-photosynthetic microorganisms contributed to these rates. Nitrate assimilation was negligible in the upper waters considered in this report (above the chlorophyll maximum and the nutricline). Potential, nitrate-saturated rates were in the range 1 to 8 g-at. N m^-3 day^-1. Seasonal comparison showed lowest rates of both carbon and nitrogen assimilation rates per chlorophyll a in winter.     

Particulate organic carbon and nitrogen in the adjacent seas of the Pacific Ocean

Particulate organic carbon (POC) and nitrogen in sea water were measured in samples collected in the adjacent seas of the Pacific Ocean during the cruises of T. S. Oshoro-Maru (1969, 1970) and the R. V. Hakuho-Maru (KH-70-4, KH-72-1). High values were obtained in the northern North Pacific and the Bering Sea, the concentration of particulate carbon in the upper 50-m layer ranged from 35 to 550 g Cl^-1. In the deep waters of these area, values above 50 g Cl^-1 were frequently observed. The lowest values in the surface layer and deeper layers were obtained in the Japan Sea (23 gCl^-1) and in the South China Sea (7 g Cl^-1) respectively. A consistent minimum was located in the intermediate waters (100–400 m) throughout the entire region studied. Variation with depth was generally irregular with marked peak values in different layers. The POC distribution consited of these peak values and a relatively uniform background concentration. These background values slightly decreased with increasing depth and were different locally. Correlation analysis between carbon concentration and apparent oxygen utilization (AOU) of ambient water for the samples in the Japan Sea and the Sulu Sea showed that there was no systematic decrease of particulate carbon with increasing AOU. In these areas, the carbon concentration scattered in the higher AOU domain ranged from 10–100 g Cl^-1. These observations support the conception that downward transport of particulate matter from the overlying surface layer in the adjacent seas of the Pacific Ocean may be fairly rapid.     

Decomposition of urea associated with photosynthesis of phytoplankton in coastal waters

Decomposition of urea in seawater was studied in Mikawa Bay, a shallow eutrophic bay on the southern coast of central Japan. The urea concentration in seawater ranged from 1.3 to 5.9 μg-at. N/1 and comprised 12 to 40% of the dissolved organic nitrogen. Using ¹⁴C labelled urea, the rate of CO₂ liberation from urea and the incorporation rate of urea carbon into the particulate organic matter were determined. For the surface samples, high rates of CO₂ liberation from urea as well as the incorporation of urea carbon into the particulate organic matter were observed in the light, while much lower rates were obtained in the dark. Incubation experiments with exposure to different light intensities revealed that the rate of CO₂ liberation from urea and the incorporation of urea carbon into particulate organic matter changed with light intensity, showing a pattern similar to that of photosynthesis. The highest liberation and incorporation rates were observed at 12,000 lux. Incubation in light and in dark produced marked decreases and increases, respectively, in urea and ammonia, while no appreciable changes were observed for nitrate and nitrite. It is suggested that urea decomposition associated with photosynthetic activity of phytoplankton is one of the major processes of urea decomposition, and that it plays a significant role in the nitrogen supply for phytoplankton in coastal waters.     

Mechanisms of production and fate of organic phosphorus in the northern Adriatic Sea

In the period from 1980 to 1984 organic phosphorus, nutrients, primary production rates (¹⁴C), chlorophyll a (chl a) standing crops, and basic oceanographic parameters were measured during 23 cruises at six stations in the open waters of the northern Adriatic Sea. These waters are significantly influenced by polluted Po River discharge. Organic phosphorus was correlated with several parameters which characterize phytoplankton activity and organic matter decomposition processes. In the late winter-spring period, organic phosphorus is produced during phytoplankton blooms. It is hypothesized that microzooplankton grazing is the main factor increasing the organic phosphorus concentrations in summer (up to 1.1 mol 1^-1). Fall and winter had much lower values (below 0.3 mol 1^-1) due to remineralization processes and an increased water mass exchange between the northern and central Adriatic regions. The direct contribution of organic phosphorus by freshwater discharge was not found to be significant. The higher organic phosphorus concentrations that can occur in low salinity waters are most likely due to their increased capability to support primary production.     

Primary productivity and sizes of pools of organic carbon in the mixed layer of the ocean

We estimated primary productivity and distributions of carbon in the phytoplankton, micro-zooplankton, and suspended and dissolved matter in various areas of the World Ocean to increase our information about the organic carbon cycle in the surface layer of the sea. Primary productivity ranged from about 0.1 gC m^–2 day^–1 in the Gulf of Mexico to 9 gC m^–2 day^–1 in nutrient-rich water off Peru. Phytoplankton carbon ranged from less than 10 g/l in the former to 750 g/l in the latter and in nutrient-rich water off southwest Africa. Micro-zooplankton carbon usually was less than 50 g/l in all waters, and was dominated by ciliates, copepodids, and copepod nauplii in all areas. Concentrations of particulate carbon ranged from 12 g/l off the east coast of South America to 850 g/l off southwest Africa. Concentrations of dissolved organic carbon varied between 0.5 and 1.5 mg/l in all areas except off Peru, where maximum values of 4.5 mg/l were observed. Turnover rates of carbon by small standing crops of micro-flagellates (1 to 5 longest dimension) and dinoflagellates in nutrient-poor waters were lower than those by large standing crops of diatoms and micro-flagellates in nutrient-rich waters. Concentrations of phytoplankton usually accounted for 20 to 55% and micro-zooplankton for 2 to 30% of the particulate carbon in the surface layer of the sea. Concentrations of dissolved organic carbon were not related to concentrations of particulate carbon in most waters except off Peru, where they appear to be directly related.     

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.     

Urea as a nitrogen source for the phytoplankton in the Oslofjord

Ambient concentrations of urea in the inner Oslofjord, Norway, showed a pronounced yearly cycle in 1980, with values in the range 0.1 to 10.0 μg-at N l^-1; this cycle resemble that of ammonia although urea concentrations were usually lower. The uptake of urea by phytoplankton was investigated using ¹⁵N. Urea was usually a less important N source than NH ₄ ⁺ , and accounted for 0 to 53% (mean 19%) of summed NH ₄ ⁺ +NO ₃ ^- + urea uptake rates from April to October. Absolute as well as relative (specific) uptake rates of urea were higher in the summer (June–August) than at other times. Uptake of urea was inhibited by NH ₄ ⁺ concentrations higher than 1 to 2 μg-at N l^-1. The summed NH ₄ ⁺ +NO ₃ ^- + urea uptake rate was exponentially related to temperature.     

Relationships between microbial distributions and the anaerobic decomposition of organic matter in surface sediments of long Island Sound,USA

Relative rates of the anaerobic decomposition of organic matter in the upper 10 cm of sediment from two stations in central Long Island Sound, USA, were compared. Sediment samples from discrete depth intervas were incubated anoxically and changes in SO ₄ ⁼ , NH ₄ ⁺ , bacterial numbers, extractable adenosine triphosphate (ATP), organic matter, and organic carbon were measured as a function of time and temperature. At both stations (15 and 34 m water-depths, respectively), the calculated rates of SO ₄ ⁼ reduction and NH ₄ ⁺ production decreased exponentially (approximately) with depth below the sediment-water interface. Over the same depth interval, ATP concentrations dropped by a factor of 6 to 7 and bacterial numbers were lower by a factor of 2 to 3. These decreases in SO ₄ ⁼ reduction, NH ₄ ⁺ production, bacterial numbers, and ATP, reflect a change in the physiological state of microbial populations with depth in the sediment and are consistent with the conclusions that the quantity of easily utilizable organic matter changes rapidly below the sediment surface and that food limitation controls the basic depth distribution of microbial activity. The average rates of SO ₄ ⁼ reduction, 29 to 39 mM year^-1 (22°C), in the top 10 cm are similar at both stations studied here, as well as at an additional station from a previous study. In contrast, average NH ₄ ⁺ production differs by a factor of 2 at the two stations, reflecting differences in the C:N ratio of the organic matter supplied to the sediment surface and differences in particle reworking by macrofauna at each site. The apparent activation energy of SO ₄ ⁼ reduction was 19±1 kcal mole^-1 and that of NH ₄ ⁺ production, 18±3 kcal mole^-1. The overall quantity of carbon required to support the calculated average SO ₄ ⁼ reduction rate in the top 10 cm is 23 g C m^-2 year^-1 and represents 36% of all the carbon available to the benthos annually and 11% of the net primary production in the water column. Directly measured fluxes of NH ₄ ⁺ from sediments to overlying water at both stations agree well with those predicted from production rates obtained by the incubation techniques.     

Photochemical reaction of peroxynitrite and carbon dioxide could account for up to 15 % of carbonate radicals generation in surface waters

The carbonate radical (CO ₃ ^?· ) is a photoinduced transient species occurring in surface waters. The carbonate radical can transform both natural compounds and xenobiotics. For instance, it can react with electron-rich substrates such as anilines, phenols and organic sulfur compounds. Here we used the APEX software to assess photochemical reactions, including the formation rates of transient species, based on water chemistry and depth, under summertime irradiation conditions. We found that the reaction between peroxynitrite and carbon dioxide is a potentially significant source of CO ₃ ^?· in sunlit surface waters, and could account for up to 10–15 % of the total CO ₃ ^?· formation. The peroxynitrite pathway to CO ₃ ^?· would be most significant at pH 7–8 and would be enhanced in waters with elevated nitrate and low alkalinity. Therefore, the proposed process could add to the known photochemical sources of CO ₃ ^?· in surface-water environments.     

Direct sampling and in situ observation of a persistent copepod aggregation in the mesopelagic zone of the Santa Barbara Basin

Observations from a one-person submersible (Wasp) in fall, 1982, revealed a persistent aggregation of non-migrating, Stage V copepodites of Calanus pacificus californicus Brodsky in a band 20±3 m thick at a depth of 450 m, about 100 m above the bottom of the Santa Barbara Basin, California. Copepod abundances, calculated from nearest-neighbor distances measured directly from the submersible, yielded maximum densities of 26×10⁶ copepodites m^-3. Quiescent behavior, low laminarinase activity, low protein content, high lipid content and evidence of low excretion rate all suggest that these copepodites were in a state of diapause. Diapausing C. pacificus californicus at other locations along the eastern Pacific coast were also captured in discrete depth plankton tows. Both the submersible observations and the net collections suggest that the dense aggregation of diapausing copepods we observed in the Santa Barbara Basin was a phenomenon associated with seasonal upwelling cycles, and that such aggregations occur during non-upwelling periods when food is scarce in surface waters. Numerous predators, especially the deep sea smelt Leuroglossus stilbius, were observed feeding upon the aggregated copepods; thus, in contrast to the conventional picture of surface-dominated food distribution, deep-water aggregations of C. pacificus californicus may support the mesopelagic community during periods of low food availability in surface waters.     

Application of a 15N tracer method to the study of dissolved organic nitrogen uptake during spring and summer in Chesapeake Bay

The dissolved organic nitrogen (DON) pool in marine waters contains a diverse mixture of compounds. It is therefore difficult to accurately estimate planktonic uptake of DON using the limited number of radiolabeled compounds commercially available. We describe a method to estimate DON uptake rates using ¹⁵N-labeled DON recently released from phytoplankton. To make ¹⁵N-labeled DON, we incubated surface water with ¹⁵NH ₄ ⁺ and then isolated the DON, including any recently released DO¹⁵N, with ion retardation resin. This DON was then added to a freshly collected water sample from the same environment to quantify the rate of DON uptake. The technique was applied to investigate rates of DON uptake relative to inorganic nitrogen in the mesohaline Chesapeake Bay during May 1990 and August 1991. The May experiment took place after the spring bloom, and rates of DON uptake [ranging from 0.31 to 0.53 g-atom (g-at) Nl^-1 h^-1] often exceeded rates of NH ₄ ⁺ and NO ₃ ^- uptake combined. The rates of DON uptake at this time were higher than estimated bacterial productivity and were not correlated with bacterial abundance or bacterial productivity. They were, however, correlated with rates of NO ₃ ^- uptake. In May, we estimate that only 7 to 32% of DON uptake was a result of urea utilization. In contrast, in August, when regenerated nutrients predominate in Chesapeake Bay, rates of DON uptake (ranging from 0.14 to 0.51 g-atom Nl^-1 h^-1) were an average of 50% of the observed rates of NH ₄ ⁺ uptake. Consistent with the May experiment, rates of DON uptake were not correlated with bacterial production. A sizable fraction of DON uptake, however, appeared to be due to urea utilization; rates of urea uptake, measured independently, were equivalent to an average of 74% of the measured rates of DON uptake. These findings suggest that, during both periods of study, at least a fraction of the measured DON uptake may have been due to utilization by phytoplankton.     

Population dynamics and production of euphausiids. I. Thysanoessa longicaudata in the North Atlantic Ocean

Adults and furciliae of Thysanoessa longicaudata (Krøyer) taken in continuous plankton records (CPR) in 1966 and 1967 were examined and measured. The measurements were converted to biomass using a length/weight equation, and net production values were calculated from these data. Two generations were spawned annually in the eastern North Atlantic Ocean and only one in the colder western waters and in the Norwegian Sea. The timing of reproduction in the spring was correlated with the mean sea-surface temperature. The highest annual production values (up to 12.24 mg dry weight m^-3 year^-1) were recorded in the eastern areas, where the ratio between production and biomass was usually between 4:1 and 8:1 compared with ratios usually between 2:1 and 3:1 in the western waters. The values for daily production per individual ranged from 3.34 g dry weight day^-1 to 17.54 g day^-1. Mortality in the early stages of development was calculated as about 98.2% of the eggs produced.     

Primary productivity and the flux of dissolved organic matter in several marine environments

Primary productivity and the flux of DO¹⁴C, dissolved saccharides (DSAC) and dissolved free primary amines (DFPA) were followed in the Sargasso Sea, Caribbean and upwelling waters of Peru. Average carbon fixation rates were 42.8, 292.8 and 4791.6 mg C m^-2 d^-1, respectively, with nocturnal respiration rates ranging from 9.8–16.3% of gross photosynthesis for the 3 areas. The release of DO¹⁴C, as a percentage of the total carbon fixed in photosynthesis, was non-detectable in the Sargasso Sea, and 3.2 and 4.4% for the Caribbean and Peruvian phytoplankton communities. Few significant changes in DSAC concentrations were recorded over a 36-h incubation period in the Sargasso Sea and Caribbean stations, whereas light-dependent accumulations of DSAC and DFPA were noted in Peruvian stations which were strongly correlated with total phytoplankton productivity. In the Peruvian stations, the average accumulation rate was 234 mg DSAC-C m^-2h^-1 while the average rate of nocturnal decomposition was 141 mg DSAC-C m^-2h^-1; diurnal and nocturnal rates of DFPA accumulation and decomposition were similar (2 mg DFPA-C m^-2h^-1). These data were used to calculate bacterial production in the upwelling waters of Peru. A general discussion of ¹⁴C-technique and routine analytical techniques for DSAC analysis is presented, as DSAC flux exceeded DO¹⁴C flux by 17-fold in coastal Peruvian stations.     

Physiology and chemical composition of nitrogen-fixing phytoplankton in the central North Pacific Ocean

The magnitude and physiological characteristics of biological nitrogen fixation have been studied in the oligotrophic waters of the North pacific gyre. The filamentous blue-green algae Trichodesmium spp. and Richelia intracellularis were the important nitrogen-fixing phytoplankton. Most of the nitrogen fixation occurs in the upper 40 m of the water column, with detectable fixation as deep as 90 m, which corresponds to about the 1 % light depth. There was no evidence of photoinhibition of nitrogen fixation, although CO₂ reduction was depressed slightly at the highest light levels. The rate of nitrogen fixation in the water column varied throughout the day, being highest in mid-morning and in late afternoon. Relatively high fixation rates were also found during periods of darkness. Elevated oxygen concentrations had a marked inhibitory effect on rates of nitrogen fixation, a pO₂ of 0.4 atm causing a 75% inhibition. Data from studies of nitrogen fixation and assimilation rates of ¹⁵N-labelled nitrate, ammonium, and urea indicate that nitrogen fixation furnished about 3% of the total daily fixed nitrogen requirement for phytoplankton growth. Studies with isolated colonies of Trichodesmium spp. indicated that 100% of their nitrogen requirement was met by nitrogen fixation. Chemical composition of the Trichodesmium colonies showed that the C:N ratio was 4.1 and that their phosphorus content relative to carbon or nitrogen was much lower than that of the total particulate material in the water column. Elevated ratios of carbon: adenosine triphosphate (ATP) also suggest that phosphorus deficiency may be limiting the growth of Trichodesmium. The magnitude of nitrogen fixation in the gyre is seasonally dependent, with high rates in late summer and autumn. At these times the water column is stratified, with phosphate and nitrate barely detectable in the upper 100 m. Our data suggest that during these months of stratification, biological fixation of nitrogen amounts to about 33 g-at N/m²/day.     

Clearance rates of the salp Thalia democratica fed naturally occurring particles

Net-captured aggregate stages of the salp Thalia democratica were fed naturally occurring particles in 2-liter bottles onboard ship. The goal was to compare clearance rates of net-captured T. democratica fed natural particles to clearance rates previously determined in laboratoryborn T. democratica fed monocultures of phytoplankton. On a cruise from 19–29 April, 1979, clearance rates were estimated by monitoring the decrease in concentration of particles with a multi-channel Coulter Counter TA II over a particle size range of 2 to 32 m equivalent spherical diameter (E.S.D.). Clearance rates were slightly lower than those measured in the laboratory, decreasing 3-fold over an 18-fold increase in the initial food concentration. Ingestion rates were slightly higher than those measured in the laboratory, increasing about 6-fold over the same range of food concentration. Within the particle concentration range examined (0.08 to 1.34 mm³l^-1), there was no lower threshold of food concentration below which clearance of particles ceased. There was no effect of the concentration of T. democratica on clearance rate over a range of 2.5 to 11.5 salps l^-1. The mean hourly ration was 2.0% of body carbon.     

Spring and summer growth rates of subarctic Pacific phytoplankton assemblages determined from carbon uptake and cell volumes estimated using epifluorescence microscopy

In order to determine whether phytoplankton growth rates were normal or depressed, total plant carbon (g l^–1) and in situ production rates (g C l^–1 d^–1) were measured for phytoplankton assemblages at Weathership Station P (50°N; 145°W) and at 53°N; 145°W in the subarctic Pacific in May and August 1984. Plant carbon, estimated from cell volumes determined using epifluorescence microscopy, was distributed as follow: 28% in the <2 m fraction, 38% in the 2 to 5 m size fraction, and the remainder in size classes >5 m. Carbon-specific growth rates (k), as doublings d^–1, were calculated for the phytoplankton assemblages as a whole at each sampling depth down to 100 m for three days in May and for four days in August. The populations in the upper part of the euphotic zone showed average doubling rates of 1 d^–1 and thus appeared to be growing at rates normally expected for the prevailing conditions of light and temperature. The low chlorophyll concentrations (0.3 to 0.4 mg chl a m^–3) characteristically found in this oceanic region do not seem to be due to very slow growth of algal populations.Contribution No. 1695 of the School of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Abundance and production rates of floating diatom mats (Rhizosolenia castracanei and R. imbricata var. shrubsolei) in the Eastern Pacific Ocean

Abundance and production rates were measured on freefloating mats composed of the diatoms Rhizosolenia castracanei and R. imbricata var. shrubsolei in the California Current and boundary waters of the central North Pacific during October, 1980. Mats ranged from 3.0 to 10.6 cm in maximum length and had a mean volume of 6.4 ml. Production rates of the diatom mats averaged 4.0 g C colony^-1 h^-1, more than 10³ times higher than that of an equal volume of surrounding water. However, because of their low density at these sites, about 1 mat m^-3, diatom mats contributed only about 1% of the total primary production. The two large Rhizosolenia species comprised almost all of the phytoplankton biomass within the mats. Rhizosolenia species rarely occurred in seawater between mats, where the phytoplankton community was dominated by a diverse array of nannoplankton. The Rhizosolenia species in the mats appeared to be in healthy condition and contained intracellular bacteria. The very high production rates of the colonies indicate high nutrient demand and, since these particular diatoms lack the symbiotic cyanobacterium Richelia intracellularis, which is implicated in N-fixation in other species of Rhizosolenia, our results indicate that other nutrient sources must be present. We discuss the potential role of the intracellular bacteria in nitrogen fixation.     

Surface-film microbial populations: diel amino acid metabolism,carbon utilization,and growth rates

Microheterotrophic dissolved free amino acid (DFAA) utilization, and microbial community and bacterial community carbon production and growth were studied using ³H-labeled organics as tracers in marine surface-film and subsurface (10 cm) waters off Baja California in November 1983. DFAA utilization was generally more rapid during the day (0.14 to 0.38 nM h^-1) than at night (0.04 to 0.14 nM h^-1) in surface-film and subsurface waters, but the percent of utilized amino acid which was respired was always greater during the night (22 to 57%) compared to the day (14 to 18%). Utilization of DFAA-carbon was estimated to range from 0.3 to 5.3 g C l^-1 d^-1 for all stations studied. In six of the 8 samples examined, the percentage of microbial carbon accounted for by the bacterial component of the population (1.4 to 5.9%) was strikingly similar to the percentage of microbial carbon production accounted for by bacterial carbon production (1.9 to 5.1%). In all of these six samples, total microbial specific-growth rates and bacterial specific-growth rates were approximately equivalent (0.9 to 2.2 d^-1 for the microbial community; 0.7 to 1.9 d^-1 for bacteria). The two exceptions were samples apparently influenced by transient flagellate populations migrating into the surface or subsurface waters at night. These observations support the conclusion that surface films contain unique and highly active microbial populations.     

Vertical fluxes and ecological significance of organic materials during the phytoplankton bloom during austral summer in Breid Bay,Antarctica

A mooring system consisting of an in situ fluorometer at a depth of 50 m and a time-series sediment trap at a depth of 110 m was deployed at the sediment trap site (70°11.536'S; 24°18.679'E; water depth: 300 m) in Breid Bay, Antarctica in austral summer from 28 December 1985 to 13 February 1986. Sinking particles, consisting of diatoms (mainly Thalassiosira antarctica), were analyzed for organic materials, stable carbon and nitrogen isotopes. Vertical fluxes of organic carbon and nitrogen were determined to be within the ranges of 12.3 to 116 mg C m^-2 d^-1 and 1.79 to 15.4 mg N m^-2 d^-1, respectively, with maxima in the middle of January 1986, after which time the organic carbon and nitrogen fluxes tended to decrease with a steep gradient. High values of ¹³C were found in the organic matter of the sinking particles collected before the middle of January, indicating that the organic matter was derived from the diatoms in the logarithmic phase of their growth. Increased abundance of glucose was found in the water-extractable carbohydrate, which was one of the sinking particles collected in the middle of January. This fact clearly indicated that the diatoms were no longer in the growth phase but rather in the stationary phase, because reserved glucan as well as various cellular organic materials were reportedly accumulated within the algal cells in the stationary phase. Fifteen species of protein amino acids with trace amounts of -alanine, -aminobutyric acid and ornithine were found in the sinking particles upon acid hydrolysis, but the amino acid composition of these samples had not been affected much by biological agents. The essential amino acid index was calculated for the sinking particles collected in the course of the sediment trap experiment. The indices obtained indicated that the sinking particles collected in Breid Bay were more ecologically significant than the sinking and suspended particles from deep waters.     

Diurnal gut pigment rhythm and metabolic rate of<Emphasis Type="Italic"> Calanus euxinus</Emphasis> in the Black Sea

The vertical distribution, diel gut pigment content and oxygen consumption of Calanus euxinus were studied in April and September 1995 in the Black Sea. Gut pigment content of C. euxinus females was associated with diel vertical migration of the individuals, and it varied with depth and time. Highest gut pigment content was observed during the nighttime, when females were in the chlorophyll a (chl a) rich surface waters, but significant feeding also occurred in the deep layer. Gut pigment content throughout the water column varied from 0.8 to 22.0 ng pigment female^–1 in April and from 0.2 to 21 ng pigment female^–1 in September 1995. From the diel vertical migration pattern, it was estimated that female C. euxinus spend 7.5 h day^–1 in April and 10.5 h day^–1 in September in the chl a rich surface waters. Daily consumption by female C. euxinus in chl a rich surface waters was estimated by taking into account the feeding duration and gut pigment concentrations. Daily carbon rations of female C. euxinus, derived from herbivorous feeding in the euphotic zone, ranged from 6% to 11% of their body carbon weight in April and from 15% to 35% in September. Oxygen consumption rates of female and copepodite stage V (CV) C. euxinus were measured at different temperatures and at different oxygen concentrations. Oxygen consumption rates at oxygen-saturated concentration ranged from an average of 0.67 g O₂ mg^–1 dry weight (DW) h^–1 at 5°C to 2.1 g O₂ mg^–1 DW h^–1 at 23°C for females, and ranged from 0.48 g O₂ mg^–1 DW h^–1 at 5°C to 1.5 g O₂ mg^–1 DW h^–1 at 23°C for CVs. The rate of oxygen consumption at 16°C varied from 0.62 g O₂ mg^–1 DW h^–1 at 0.65 mg O₂ l^–1 to 1.57 g O₂ mg^–1 DW h^–1 at 4.35 mg O₂ l^–1 for CVs, and from 0.74 g O₂ mg^–1 DW h^–1 at 0.57 mg O₂ l^–1 to 2.24 g O₂ mg^–1 DW h^–1 at 4.37 mg O₂ l^–1 for females. From the oxygen consumption rates, daily requirements for the routine metabolism of females were estimated, and our results indicate that the herbivorous daily ration was sufficient to meet the routine metabolic requirements of female C. euxinus in April and September in the Black Sea.Communicated by O. Kinne, Oldendorf/Luhe