Strong variability in bacterioplankton abundance and production in central and western Bay of Bengal

With large influx of freshwater that decreases sea-surface salinities, weak wind forcing of <10 m s⁻¹ and almost always warm (>28°C) sea-surface temperature that stratifies and shallows the mixed layer leading to low or no nutrient injections into the surface, primary production in Bay of Bengal is reportedly low. As a consequence, the Bay of Bengal is considered as a region of low biological productivity. Along with many biological parameters, bacterioplankton abundance and production were measured in the Bay of Bengal during post monsoon (September–October 2002) along an open ocean transect, in the central Bay (CB, 88°E) and the other transect in the western Bay (WB). The latter representing the coastal influenced shelf/slope waters. Bacterioplankton abundances (<2 × 10⁹cells l⁻¹) were similar to those reported from the HNLC equatorial Pacific and the highly productive northern Arabian Sea. Yet, the thymidine uptake rates along CB (average of 1.46 pM h⁻¹) and WB (average of 1.40 pM h⁻¹) were less than those from the northwestern Indian Ocean. These abundances and uptake rates were higher than those in the oligotrophic northwestern Sargasso Sea (<7 × 10⁸ cells l⁻¹; av 1.0 pM h⁻¹). Concentrations of chlorophyll a (chl a), primary production rates and total organic carbon (TOC) were also measured for a comparison of heterotrophic and autotrophic production. In the WB, bacterioplankton carbon biomass equaled ∼ 95% of chl a carbon than just 31% in the CB. Average bacterial:primary production (BP:PP) ratios accounted for 29% in the CB and 31% in the WB. This is mainly due to lower primary productivity (PP) in the WB (281 mg C m⁻² d⁻¹) than in the CB (306 mg C m⁻² day⁻¹). This study indicates that bacteria–phytoplankton relationship differs in the open (CB) and coastal waters (WB). Higher abundance and contrastingly low bacterial production (BP) in WB may be because of the riverine bacteria, brought in through discharges, becoming dormant and unable to reproduce in salinities of 28 or more psu. Heterotrophic bacteria appear to utilize in situ DOC rather rapidly and their carbon demand is ∼50% of daily primary production. It is also apparent that allochthonous organic matter, in particular in the western Bay, is important for meeting their carbon demand.     

In situ net primary productivity and photosynthesis of Antarctic sea ice algal, phytoplankton and benthic algal communities

Primary production at Antarctic coastal sites is contributed from sea ice algae, phytoplankton and benthic algae. Oxygen microelectrodes were used to estimate sea ice and benthic primary production at several sites around Casey, a coastal area in eastern Antarctica. Maximum oxygen export from sea ice was 0.95 mmol O₂ m⁻² h⁻¹ (~11.7 mg C m⁻² h⁻¹) while from the sediment it was 6.08 mmol O₂ m⁻² h⁻¹ (~70.8 mg C m⁻² h⁻¹). When the ice was present O₂ export from the benthos was either low or negative. Sea ice algae assimilation rates were up to 3.77 mg C (mg Chl-a)⁻¹ h⁻¹ while those from the benthos were up to 1.53 mg C (mg Chl-a)⁻¹ h⁻¹. The contribution of the major components of primary productivity was assessed using fluorometric techniques. When the ice was present approximately 55–65% of total daily primary production occurred in the sea ice with the remainder unequally partitioned between the sediment and the water column. When the ice was absent, the benthos contributed nearly 90% of the primary production.     

Microbial activity and carbon, nitrogen, and phosphorus content in a subtropical seagrass estuary (Florida Bay): evidence for limited bacterial use of seagrass production

Bacterial abundance, production, and extracellular enzyme activity were determined in the shallow water column, in the epiphytic community of Thalassia testudinum, and at the sediment surface along with total carbon, nitrogen, and phosphorus in Florida Bay, a subtropical seagrass estuary. Data were statistically reduced by principle components analysis (PCA) and multidimensional scaling and related to T. testudinum leaf total phosphorus content and phytoplankton biomass. Each zone (i.e., pelagic, epiphytic, and surface sediment community) was significantly dissimilar to each other (Global R = 0.65). Pelagic aminopeptidase and sum of carbon hydrolytic enzyme (esterase, peptidase, and α- and β-glucosidase) activities ranged from 8 to 284 mg N m⁻² day⁻¹ and 113–1,671 mg C m⁻² day⁻¹, respectively, and were 1–3 orders of magnitude higher than epiphytic and sediment surface activities. Due to the phosphorus-limited nature of Florida Bay, alkaline phosphatase activity was similar between pelagic (51–710 mg P m⁻² day⁻¹) and sediment (77–224 mg P m⁻² day⁻¹) zones but lower in the epiphytes (1.1–5.2 mg P m⁻² day⁻¹). Total (and/or organic) C (111–311 g C m⁻²), N (9.4–27.2 g N m⁻²), and P (212–1,623 mg P m⁻²) content were the highest in the sediment surface and typically the lowest in the seagrass epiphytes, ranging from 0.6 to 8.7 g C m⁻², 0.02–0.99 g N m⁻², and 0.5–43.5 mg P m⁻². Unlike nutrient content and enzyme activities, bacterial production was highest in the epiphytes (8.0–235.1 mg C m⁻² day⁻¹) and sediment surface (11.5–233.2 mg C m⁻² day⁻¹) and low in the water column (1.6–85.6 mg C m⁻² day⁻¹). At an assumed 50% bacterial growth efficiency, for example, extracellular enzyme hydrolysis could supply 1.8 and 69% of epiphytic and sediment bacteria carbon demand, respectively, while pelagic bacteria could fulfill their carbon demand completely by enzyme-hydrolyzable organic matter. Similarly, previously measured T. testudinum extracellular photosynthetic carbon exudation rates could not satisfy epiphytic and sediment surface bacterial carbon demand, suggesting that epiphytic algae and microphytobenthos might provide usable substrates to support high benthic bacterial production rates. PCA revealed that T. testudinum nutrient content was related positively to epiphytic nutrient content and carbon hydrolase activity in the sediment, but unrelated to pelagic variables. Phytoplankton biomass correlated positively with all pelagic components and sediment aminopeptidase activity but negatively with epiphytic alkaline phosphatase activity. In conclusion, seagrass production and nutrient content was unrelated to pelagic bacteria activity, but did influence extracellular enzyme hydrolysis at the sediment surface and in the epiphytes. This study suggests that seagrass-derived organic matter is of secondary importance in Florida Bay and that bacteria rely primarily on algal/cyanobacteria production. Pelagic bacteria seem coupled to phytoplankton, while the benthic community appears supported by epiphytic and/or microphytobenthos production.     

Effect of nutrient enrichment in the field on the biomass, growth and calcification of the giant clam Tridacna maxima

Nutrients were added separately and combined to an initial concentration of 10 μM (ammonium) and/or 2 μM (phosphate) in a series of experiments carried out with the giant clam Tridacna maxima at 12 microatolls in One Tree Island lagoon, Great Barrier Reef, Australia (ENCORE Project). These nutrient concentrations remained for 2 to 3 h before returning to natural levels. The additions were made every low tide (twice per day) over 13 and 12 mo periods for the first and second phase of the experiment, respectively. The nutrients did not change the wet tissue weight of the clams, host C:N ratio, protein content of the mantle, calcification rates or growth rates. However, ammonium (N) enrichment alone significantly increased the total population density of the algal symbiont (Symbiodinium sp.: C = 3.6 · 10⁸ cell clam⁻¹, N = 6.6 · 10⁸ cell clam⁻¹, P = 5.7 · 10⁸ cell clam⁻¹, N + P = 5.7 · 10⁸ cell clam⁻¹; and C = 4.1 · 10⁸ cell clam⁻¹, N = 5.1 · 10⁸ cell clam⁻¹, P = 4.7 · 10⁸ cell clam⁻¹, N + P = 4.5 · 10⁸ cell clam⁻¹, at the end of the first and second phases of the experiment, respectively), although no differences in the mitotic index of these populations were detected. The total chlorophyll a (chl a) content per clam but not chlorophyll a per cell also increased with ammonium addition (C = 7.0 mg chl a clam⁻¹, N = 13.1 mg chl a clam⁻¹, P = 12.9 mg chl a clam⁻¹, N + P = 11.8 mg chl a clam⁻¹; and C = 8.8 mg chl a clam⁻¹, N = 12.8 mg chl a clam⁻¹; P = 11.2 mg chl a clam⁻¹, N + P = 11.3 mg chl a clam⁻¹, at the end of the first and second phases of the experiment, respectively). The response of clams to nutrient enrichment was quantitatively small, but indicated that small changes in inorganic nutrient levels affect the clam–zooxanthellae association. Received: 2 June 1997 / Accepted: 9 June 1997     

Spring sea ice photosynthesis,primary productivity and biomass distribution in eastern Antarctica, 2002–2004

While it is known that Antarctic sea ice biomass and productivity are highly variable over small spatial and temporal scales, there have been very few measurements from eastern Antarctic. Here we attempt to quantify the biomass and productivity and relate patterns of variability to sea ice latitude ice thickness and vertical distribution. Sea ice algal biomass in spring in 2002, 2003 and 2004 was low, in the range 0.01–8.41 mg Chl a m⁻², with a mean and standard deviation of 2.08 ± 1.74 mg Chl a m⁻² (n = 199). An increased concentration of algae at the bottom of the ice was most pronounced in thicker ice. There was little evidence to suggest that there was a gradient of biomass distribution with latitude. Maximum in situ production in 2002 was approximately 2.6 mg C m⁻² h⁻¹ with assimilation numbers of 0.73 mg C (mg Chl a)⁻¹ h⁻¹. Assimilation numbers determined by the ¹⁴C incubations in 2002 varied between 0.031 and 0.457 mg C (mg Chl a)⁻¹ h⁻¹. Maximum fluorescence quantum yields of the incubated ice samples in 2002 were 0.470 ± 0.041 with E _k indices between 19 and 44 μmol photons m⁻² s⁻¹. These findings are consistent with the shade-adapted character of ice algal communities. In 2004 maximum in situ production was 5.9 mg C m⁻² h⁻¹ with an assimilation number of 5.4 mg C (mg Chl a)⁻¹ h⁻¹. Sea ice biomass increased with ice thickness but showed no correlation with latitude or the time the ice was collected. Forty-four percent of the biomass was located in bottom communities and these were more commonly found in thicker ice. Surface communities were uncommon.     

Production and biomass accumulation of periphytic diatoms growing on glass slides during a 1-year cycle in a subtropical estuarine environment (Bay of Paranaguá, southern Brazil)

Production rates, chlorophyll concentrations and general composition of periphytic diatom communities growing on glass slides were studied in relation to environmental parameters during one seasonal cycle in the Bay of Paranaguá, southern Brazil. Slides were routinely submersed at 1, 2 and 3 m depth and recovered weekly for microscopic examinations, analyses of chlorophyll, cell counts and in situ photosynthetic incubations using the Winkler titration method. Water samples were also collected at surface and bottom layers for determinations of temperature, salinity, nutrients and chlorophyll in the water. The periphytic community was mainly formed by epipelic and epipsammic species, dominated by Navicula phyllepta, Cylindrotheca closterium, Navicula spp. and Amphora sp. Weekly chlorophyll a and cell accumulations on slides varied from <1–32 mg m⁻² and up to 31 × 10⁸cells m⁻², respectively. Photosynthetic rates varied from <1 to 35 mg oxygen mg chlorophyll a ⁻¹ h⁻¹, with higher values in summer. Daily production varied from 5 to 3,600 mg oxygen m⁻² day⁻¹ (<0.01–1.4 g carbon m⁻² day⁻¹). Multiple regression analysis revealed that vertical differences in light conditions and grazing pressure jointly affected the influence of temperature on the seasonal patterns of cell densities and chlorophyll concentrations according to depth. Received: 27 April 2000 / Accepted: 16 August 2000     

Phytoplankton growth and microzooplankton grazing rates in a restricted Mediterranean lagoon (Bizerte Lagoon,Tunisia)

Phytoplankton growth and microzooplankton grazing were investigated in the restricted Bizerte Lagoon in 2002 and 2004. The 2002 study, carried out at one station from January to October, showed significant seasonal variations in phytoplankton dynamics. High growth rates (0.9–1.04 day⁻¹), chlorophyll a (Chl a) concentrations (6.6–6.8 μg l⁻¹) and carbon biomass (392–398 μg C l⁻¹) were recorded in summer (July), when several chain-forming diatoms had intensively proliferated and dominated the carbon biomass (74%). In 2004, four stations were studied during July, a period also characterized by the high proliferation of several diatoms that made up 70% of the algal carbon biomass. In 2004, growth rates (0.34–0.45 day⁻¹) and biomass of algae (2.9–5.4 μg Chl a l⁻¹ and 209–260 μg C l⁻¹) were low, which may be related to the lower nutrient concentrations recorded in 2004. Microzooplankton >5 μm were mainly composed of heterotrophic dinoflagellates and ciliates. Microzooplankton biomass peaked during summer (2002 320–329, 2004 246–361 μg C l⁻¹), in response to the enhanced phytoplankton biomass and production. The grazer biomass was dominated by ciliates (71–76%) in July 2002 and by heterotrophic dinoflagellates (52–67%) in July 2004. Throughout the year and at different stations, microzooplankton grazed actively on phytoplankton, removing 26–58% of the Chl a and 57-84% of the primary production. In 2002, the highest grazing impact was observed on the large algae (>10 μm) during the period of diatom dominance. These results have a significant implication for carbon export to depth. Indeed, the recycling of most of the diatom production by the microbial food web in the upper water column would reduce the flux of material to the seafloor. This should be considered when modeling the carbon cycling in coastal environments and under conditions of diatom dominance. During both studies, ciliates had higher growth rates (0.5–1.5 day⁻¹) and a higher carbon demand (165–470 μg C l⁻¹ day⁻¹) than dinoflagellates (0.1–0.5 day⁻¹, 33–290 μg C l⁻¹ day⁻¹). Moreover, when grazer biomass was dominated by ciliates (in July 2002), herbivory accounted for 71–80% of the C ingested by microzooplankton while it accounted only for 14–23% when dinoflagellates dominated the grazer biomass (in July 2004). These results suggest that, in contrast to findings from open coastal waters, ciliate species of the restricted Bizerte Lagoon were more vigorous grazers of the large algae (diatoms) than were dinoflagellates.     

Flux capacities and acclimation costs in Trichodesmium from the Gulf of Mexico

Phytoplankton function and acclimation are driven by catalytic protein complexes that mediate key physiological transformations, including generation of photosynthetic ATP and reductant, and carbon and nitrogen fixation. Quantitation of capacities for these processes allows estimation of rates for key ecosystem processes, and identification of factors limiting primary productivity. We herein present molar quantitations of PSI, PSII, ATP synthase, RuBisCO and the Fe protein of nitrogenase of Trichodesmium collected from the Gulf of Mexico, in comparison to determinations for a range of cyanobacteria growing in culture. Using these measurements, estimates were generated for Trichodesmium capacities for carbon fixation of 1–3.4 g C g chl a ⁻¹ h⁻¹ and nitrogen fixation of 0.06–0.17 g N g chl a ⁻¹ h⁻¹, with diel variations in capacities. ATP synthase levels show that ATP synthesis capacity is sufficient to support these levels of carbon and nitrogen fixation, and that ATP synthase levels change over the day in accordance with the ATP demands of nitrogenase and RuBisCO activity. Levels of measured complexes indicate that Trichodesmium manifests n-type diel light acclimation through rapid changes in RuBisCO:PSII, supported by significant investment of cellular nitrogen. The plasticity in the levels and stoichiometry of these core complexes show that changes in the abundance of core protein complexes are an important component of acclimation and regulation of metabolic function by Trichodesmium populations. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Grazing of phytoplankton by copepods in eastern Antarctic coastal waters

Chlorophyll a, primary productivity and grazing by copepods on phytoplankton were measured in the upper water column during the summer of 1994/1995 at a coastal site near Davis Station, East Antarctica. Chlorophyll a was at a maximum in mid-December, then dropped markedly as the coastal fast ice melted and broke‐out. Phytoplankton biomass increased again from mid‐ to late‐February. Copepods accounted for at least 65% of zooplankton biomass in the water column before sea ice break‐out, whereas larval polychaetes and ctenophores dominated after ice break‐out. Oncaeacurvata was the numerically dominant species throughout the study. The highest grazing rate (8.7 mg C␣m⁻³␣d⁻¹) was recorded on 21 December when O.␣curvata accounted for 64% of the total. Grazing had decreased markedly by 28 December (0.9 mg C m⁻³ d⁻¹); again O. curvata accounted for over 50% of the total ingested. Copepod grazing increased after ice break-out until the last experiment on 20 February (⋍5 mg C␣m⁻³␣d⁻¹). The main species responsible for grazing during this period were O. curvata, Oithonasimilis, Calanoidesacutus and unidentified copepod nauplii. It was estimated that copepods removed between 1 and 5% of primary productivity. Received: 11 October 1996 / Accepted: 22 October 1996     

Phytoplankton production patterns in Massachusetts Bay and the absence of the 1998 winter–spring bloom

The seasonal productivity cycle and factors controlling annual variation in the timing and magnitude of the winter–spring bloom were examined for several locations (range: 42°20.35′–42°26.63′N; 70°44.19′–70°56.52′W) in Boston Harbor and Massachusetts Bay, USA, from 1995 to 1999, and compared with earlier published data (1992–1994). Primary productivity (mg C m⁻² day⁻¹) in Massachusetts Bay from 1995 to 1999 was generally characterized by a well-developed winter–spring bloom of several weeks duration, high but variable production during the summer, and a prominent fall bloom. The bulk of production (mg C m⁻³ day⁻¹) typically occurred in the upper 15 m of the water column. At a nearby Boston Harbor station a gradual pattern of increasing areal production from winter through summer was more typical, with the bulk of production restricted to the upper 5 m. Annual productivity in Massachusetts Bay and Boston Harbor ranged from a low of 160 g C m⁻² year⁻¹ to a high of 787 g C m⁻² year⁻¹ from 1992 to 1999. Mean annual productivity was higher (mean=525 g C m⁻² year⁻¹) and more variable near the harbor entrance than in western Massachusetts Bay. At the harbor station productivity varied more than 3.5-fold (CV=40%) over an 8 year sampling period. Average annual productivity (305–419 g C m⁻² year⁻¹) and variability around the means (CV=25–27%) were lower at both the outer nearfield and central nearfield regions of Massachusetts Bay. Annual productivity in 1998 was unusually low at all three sites (<220 g C m⁻² year⁻¹) due to the absence of a winter–spring phytoplankton bloom. Potential factors influencing the occurrence of a spring bloom were investigated. Incident irradiance during the winter–spring period was not significantly different (P > 0.05) among years (1995–1999). The mean photic depth during the bloom period was significantly deeper (P < 0.05) in 1998, signifying greater light availability with depth. Nutrients were also in abundance during the winter–spring of 1998 with stratified conditions not observed until May. In general, the magnitude of the winter–spring bloom in Massachusetts Bay from 1995 to 1999 was significantly correlated with winter water temperature (r ²=0.78) and zooplankton abundance (r ²=0.74) over the bloom period (typically February–April). The absence of the 1998 bloom was associated with higher than average water temperature and elevated levels of zooplankton abundance just prior to, and during, the peak winter–spring bloom period. Received: 3 July 2000 / Accepted: 6 December 2000     

Contrasting life histories and secondary production of populations of Munnopsurus atlanticus (Isopoda: Asellota) from two bathyal areas of the NE Atlantic and the NW Mediterranean

The isopod Munnopsurus atlanticus occupies bathyal depths in both the Bay of Biscay (NE Atlantic; between 383 and 1022 m) and in the Catalan Sea (Northwestern Mediterranean; between 389 and 1859 m). The species was dominant in both assemblages, reaching bathymetric peaks of abundance on the upper part of the continental slope (400 m depth) in the Bay of Biscay and at ˜600 m in the Catalan Sea. Both the Atlantic and the Mediterranean populations are bivoltines. Demographic analysis of the Bay of Biscay population revealed the production of two generations per year with different potential longevity (5 mo for G₁ and 11 mo for G₂). The mean cohort-production interval (CPI) was estimated at 8 mo, and results of the demographic analysis were also used to estimate production for the Catalan Sea populations. Mean annual density (D) and biomass (B) were higher in the Bay of Biscay (D = 356.7 individuals 100 m⁻²; B = 0.803 mg dry wt m⁻² yr⁻¹) than in the Mediterranean (D = 16.3 individuals 100 m⁻²; B = 0.078 mg dry wt m⁻² yr⁻¹). Also, mean annual production was an order of magnitude higher in the Atlantic (between 4.063 and 4.812 mg dry wt 100 m⁻² yr⁻¹ depending on the method used) than in the Catalan Sea (between 0.346 and 0.519 mg dry wt 100 m⁻² yr⁻¹). M. atlanticus feeds on a wide variety of benthic and pelagic food sources. In both study areas, phytodetritus was not important in the diet of M. atlanticus. In contrast, gut-content data suggested an indirect coupling with phytoplankton production in both areas via foraminiferans. The life history and the recorded production are considered in respect to both the dynamics and levels of primary production and the total mass flux in the respective study areas. Differences in the secondary production of both populations seemed to be more consistently explained by differences in total mass flux than by differences in the primary production levels; this is also consistent with the variety of food sources exploited by M. atlanticus. Received: 22 February 1999 / Accepted: 3 February 2000     

Life cycle of <Emphasis Type="Italic">Pseudocalanus acuspes</Emphasis> Giesbrecht (Copepoda,Calanoida) in the Central Baltic Sea: II. Reproduction,growth and secondary production

The population dynamics of Pseudocalanus acuspes in the Central Baltic Sea were studied from March 2002 to May 2003 on a monthly basis. All stages were present year round with a stage shift from nauplii to older copepodite stages over the course of the year. Biomass, estimated from prosome length, peaked between May and September with maximum recorded values of 594 and 855 mg C m⁻² in May 2002 and 2003, respectively. Differences in biomass between stations up to a factor of 20 were observed especially in April/May and October. Mean egg production rate (EPR) showed a seasonal course and was highest in April 2002 and 2003 with 3.6 and 2.1 eggs f⁻¹ day⁻¹, respectively, corresponding to a mean weight-specific egg production rate (SEPR) of 0.13 and 0.04. Egg production seems to be limited by food from May on. Stage durations determined from moulting experiments turned out to be extremely long. Maximum growth rates based on stage durations of 15–25 days at 4°C in May and July 2003 amounted for 0.03–0.05 day⁻¹ in CI-CIV. Comparing these rates with rates derived from temperature–development relationships for P. acuspes from the literature resulted in five times higher growth rates for the latter case. Secondary production reached values up to 9.1 mg C m⁻² day⁻¹ (method for continuously reproducing populations) and 10.5 mg C m⁻² day⁻¹ (increment summation).     

Differences in productivities between the Great Australian Bight and the Gulf of Carpentaria,Australia, in summer

Phytoplankton standing crop (chlorophyll a) and primary productivity were recorded, and zooplankton biomass was estimated in the two large bays of Australia, the Great Australian Bight on the south coast (December, 1965) and the Gulf of Carpentaria on the north coast (December, 1968). In the Gulf of Carpentaria, the phytoplankton standing crop (average, 27.3 mg chlorophyll a m^-2) and primary productivity (average, 133.1 mg C m^-2 h^-1), as well as zooplankton biomass (average, 305.3 mg wet weight m^-3) are much higher than in the Great Australian Bight (12.1 mg chlorophyll a m^-2, 18.2 mg C m^-2 h^-1, 7.1 mg wet weight m^-3, respectively). The unexpectedly low productivity values in the Great Australian Bight are attributable to environmental conditions of this bay, which obtains neither replenishment of nutrients from the land nor receives upwelling of deep water.     

Salps in the Lazarev Sea,Southern Ocean: I. Feeding dynamics

Feeding dynamics of the Antarctic salps Ihlea racovitzai and Salpa thompsoni were studied in the Lazarev Sea in fall 2004, summer 2005–2006 and winter 2006. Pigment concentrations in the guts of both species were positively correlated with ambient surface chlorophyll a (chl a). No evidence was found for salp clogging even at dense surface concentrations of up to 7 μg chl a L⁻¹. However, gut pigment concentrations had a lower range than ambient pigment concentrations, suggesting that salps increased retention times of ingested material in low-food environments. For medium-sized I. racovitzai and S. thompsoni, estimated individual daily rations reached 7–10 and >100% of body carbon in winter and summer, respectively. Daily respiratory needs of I. racovitzai and S. thompsoni accounted for 28 and 22% of daily carbon assimilation based on pigment ingestion rates in winter, and for 2 and 1% in summer, respectively. The grazing impact of the salp populations on the phytoplankton standing stock was negligible during all seasons due to generally low salp densities. Fatty acid trophic biomarkers in the salps suggest high year-round contributions of flagellates and modest contributions of diatoms to the salp’s diet. These markers showed low seasonal variability for I. racovitzai. The more pronounced seasonality of trophic markers in S. thompsoni were likely related to their generally deeper residence depth in winter linked to a seasonal alternation of sexual and asexual generations.     

Carbon and nitrogen exchange between sandy beach clams (Donax serra) and kelp beds in the Benguela coastal upwelling region

Carbon consumption and nitrogen requirements were estimated for populations of the sandy beach bivalve Donax serra on nine beaches of the west coast of South Africa. Subtidal populations composed mainly of adult clams were responsible for the bulk of standing stock (3538 g C m⁻¹), annual carbon consumption (13 444 g C m⁻¹ yr⁻¹), faeces production (6478 g C m⁻¹ yr⁻¹ ) and nitrogen regeneration (2525 g N m⁻¹ yr⁻¹). Kelp detritus, bacteria and kelp consumers' faeces available in the water column surpass several times the carbon and nitrogen requirements of intertidal and subtidal clam populations. Individual Donax serra pop ulations, in turn, may regenerate up to 3.2% of the total nitrogen requirements of all primary producers from kelp beds and 14% of the requirements of phytoplankton. These high standing stocks of clams are presumably supported mainly by organic matter originating from kelp which, in contrast to phytoplankton, is in constant supply and comprises the largest proportion of the annual production of particulate organic matter on this coast. Wide and shallow continental shelves with gentle slopes probably limit the penetration of upwelled waters to the nearshore waters, decreasing the influence of external inputs and increasing the importance of internal flows of nutrients and carbon within the nearshore zone. In this context, sandy beaches, rocky shores and kelp beds may be more closely interlinked compartments of a larger ecosystem encompassing the whole nearshore than traditionally thought. Received: 28 August 1996 / Accepted: 7 October 1996     

UV- and salinity-induced oxidative effects in the marine diatom <Emphasis Type="Italic">Cylindrotheca closterium</Emphasis> during simulated emersion

The diatom Cylindrotheca closterium was exposed to transient light- and osmotic conditions as occur during its tidal emersion. The objective was to analyze how this simulated emersion contributes to the production of active oxygen species (AOS) and via this, to oxidative cell damage. Light- and salinity conditions were varied in factorial combination: low light (no UVB) or high light (unweighted UVB-dose rates of respectively 0.01; 0.07; 0.24; 1.03 W m⁻²) at normal (30 psu) or high salinity (60 psu). UVB (0.01–0.24 W m⁻²) and high salinity had a significant, negative effect on the photosynthetic efficiencies ΔF/F _m’ (steady-state quantum yield) and F _v/F _m (maximum yield). UVB at 1.03 W m⁻² (15 kJ m⁻² d⁻¹) almost arrested electron transport. At ecologically relevant UVB levels, i.e. below 0.24 W m⁻² (≈3.4 kJ m⁻² d⁻¹) with UVB:PAR<0.4:100 (PAR photosynthetically active radiation) only dynamic photoinhibition was observed (protection via heat dissipation). Non-photochemical quenching was positively correlated with the de-epoxidation of diadinoxanthin (DD) to diatoxanthin (DT). A decreasing ratio DT/(DD+DT) after 4 h of UVB at >0.07 W m⁻² and at 60 psu indicated a reversal of the diatom xanthophyll cycle (diminished photoprotection) which may be caused by an enhanced AOS production. Oxidative stress and -damage to C. closterium cells were assessed applying fluorescent indicator dyes, via confocal microscopy and quantitative image analysis. AOS production rates (cellular DCF fluorescence) were stimulated by UV, and were ~50% higher at 60 psu. AOS production decreased with an increasing pre-exposure (0–4 h) to normal UVB (0.24 W m⁻²), which indicated a stimulation of the antioxidative defence. Non-protein thiols (indicator CMF) and glutathione pools (HPLC-analyzed) decreased with UVB-dose rates (0.01–0.24 W m⁻²), most likely due to AOS-mediated thiol oxidation. Hypersalinity (60 psu) and UVB (0.01–0.24 W m⁻²) caused membrane depolarization (dye DIBAC₄(3)) and phospholipid hydrolysis (phospholipase A₂ dye: bis-BODIPY FL-C₁₁-PC). AOS production may have diminished the membrane polarity, and peroxidized the membrane lipids (HPLC-analyzed malondialdehyde) which enhanced PLA₂ activity. The dyes indicated an increased oxidative (lipid) damage at a 15% inhibition of photosynthesis in this diatom, at UVB levels and salinities that can be expected in situ during its periodic tidal emersion.     

Life history and production of the mysid <Emphasis Type="Italic">Orientomysis robusta</Emphasis>: high <Emphasis Type="Italic">P/B</Emphasis> ratio in a shallow warm-temperate habitat of the Sea of Japan

Although mysids play important roles in marine food chains, studies on their production are scarce, especially for warm-water species. We investigated life history and production of Orientomysis robusta in a shallow warm-temperate habitat of the Sea of Japan. Its spawning and recruitment occurred throughout the year; 19 overlapping cohorts were recognizable over an annual cycle. The summer cohorts recruited in July–September exhibited rapid growth, early maturity, small brood size, and small body size. A converse set of life history traits characterized the autumn–winter cohorts recruited in October–March. The spring cohorts recruited in April–June had intermediate characteristics of both cohorts. Life spans were 19–33, 21–48, and 69–138 days for summer, spring, and autumn–winter cohorts, respectively, and mortality rates were high for spring and summer cohorts, especially during June–August but were low for autumn–winter cohorts. Production calculated from the summation of growth increments was 488.8 mg DW m⁻² year⁻¹ with an annual P/B ratio of 21.26. The short life span seems to be responsible for such an extremely high P/B ratio. A method not requiring recognition and tracking cohorts gave similar values (534.0 mg DW m⁻² year⁻¹ and 20.49). The close agreement in production values between the two methods indicates our estimates are valid.     

Nitrate storage by phytoplankton in a coastal upwelling environment

The storage of nitrate by phytoplankton cells during the early phases of upwelling was studied in coastal stations off northern Spain (southern Bay of Biscay) between 1990 and 1994. In this region, a persistent upwelling during summer is characterised by intermittent pulses of variable intensity, and increased nutrient concentrations in the surface layer. The main effect of an upwelling pulse on phytoplankton distribution is the shifting of the chlorophyll a and primary production maxima to near the surface. When the upwelling relaxes, thermal stratification of the water column occurs, and a distinct subsurface chlorophyll maximum develops below the production maximum. An accumulation of intracellular nitrate characterized the early phases of upwelling (mean = 2.73 μmol N m⁻³), maximum concentrations being attained at depths where biomass and production values were moderate. In contrast, phytoplankton cells from non-upwelling situations contained significantly lower concentrations of intracellular nitrate (mean = 0.17 μmol N m⁻³). The variations in the intracellular pool of nitrate may result from the differential allocation of resources within the cell as a result of variations in the energy available, since the uptake and assimilation of nitrate is a relatively expensive process involving several enzymatic systems. We hypothesize that nitrate storage by phytoplankton cells is characteristic of early phases of upwelling and is linked to patterns of carbon fixation. Average nitrogen budgets for upwelling and non-upwelling situations indicate that intracellular nitrate reserves are not responsible for maintaining high phytoplankton growth rates, since they only account for <2% of daily primary production during upwelling events. Received: 28 August 1996 / Accepted 3 December 1996     

Seasonal variations in the growth rates of euphausiids (<Emphasis Type="Italic">Thysanoessa inermis</Emphasis>, <Emphasis Type="Italic">T. spinifera</Emphasis>, and <Emphasis Type="Italic">Euphausia pacifica</Emphasis>) from the northern Gulf of Alaska

The euphausiids Thysanoessa inermis (Kroyer 1846), Thysanoessa spinifera (Holmes 1900), and Euphausia pacifica (Hansen 1911) are key pelagic grazers and also important prey for many commercial fish species in the Gulf of Alaska (GOA). To understand the role of the euphausiids in material flows in this ecosystem their growth rates were examined using the instantaneous growth rate (IGR) technique on the northern GOA shelf from March through October in 2001–2004. The highest mean molting increments (over 5% of uropod length increase per molt) were observed during the phytoplankton bloom on the inner shelf in late spring for coastal T. inermis, and on the outer shelf in summer for T. spinifera and more oceanic E. pacifica, suggesting tight coupling with food availability. The molting rates were higher in summer and lower in spring, for all species and were strongly influenced by temperature. Mean inter-molt periods calculated from the molting rates, ranged from 11 days at 5°C to 6 days at 8°C, and were in agreement with those measured directly during long-term laboratory incubations. Growth rate estimates depended on euphausiid size, and were close to 0 in early spring, reaching maximum values in May (0.123 mm day⁻¹ or 0.023 day⁻¹ for T. inermis) and July (0.091 mm day⁻¹ or 0.031 day⁻¹ for T. spinifera). The growth rates for E. pacifica remained below 0.07 mm day⁻¹ (0.016 day⁻¹) throughout the season. The relationship between T. inermis weight specific growth rate (adjusted to 5°C) and ambient chlorophyll-a concentration fit a Michaelis–Menten curve (r ² = 0.48) with food saturated growth rate of 0.032 day⁻¹ with half saturation occurring at 1.65 mg chl-a m⁻³, but such relationships were not significant for T. spinifera or E. pacifica.     

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