Total organic sulfur and dimethylsulfoniopropionate in marine phytoplankton: intracellular variations

Cellular levels of particulate organic sulfur (POS) and dimethylsulfoniopropionate (DMSP) were determined in cultures of five species of marine phytoplankton, Amphidinium carterae, Prorocentrum minimum, Emiliania huxleyii, Dunaliella tertiolecta and Skeletonema costatum. The first three are known producers of DMSP while the latter two produce little or non-detectable amounts of DMSP. In those species which produced significant amounts of DMSP, intracellular levels of POS and DMSP varied for the dinoflagellates (A. carterae and P. minimum) while they remained fairly constant for the prymnesiophyte (E. huxleyii) over the growth cycle (until late stationary phase), and DMSP accounted for the majority of the POS (50 to 100%). In species with low levels of DMSP, intracellular POS and DMSP decreased or remained constant over the growth cycle, and DMSP accounted for a much lower percentage of the POS (0 to 40%). Species with high DMSP concentrations had higher POS levels per unit cell volume as well, and DMSP accounted for substantially higher percentages of the particulate organic carbon (POC). Molar N:S ratios suggest a non-protein origin for much of the sulfur in the species producing DMSP. During late stationary phase, an increasing percentage of the DMSP became extracellular in the dinoflagellate and diatom (S. costatum) cultures, suggesting leakage. In a bacterized algal culture, measured quantities were considerably less than in an axenic counterpart, suggesting consumption.     

Light-dependence of carbon and sulfur production by polar clones of the genus Phaeocystis

Blooms of the marine prymnesiophyte genus Phaeocystis link the oceanic and atmospheric compartments of the carbon and sulfur cycles. Modeling the fluxes of dimethylsulfide from the ocean to the atmosphere has been limited due to a lack of information on functional responses to environmental variables. In this study, the light-dependence of extracellular carbon production and dimethyl sulfide (DMS) production by non-axenic polar clones of Phaeocystis spp. was examined at different growth stages. Comparative experiments were run with non-axenic arctic clones of the diatoms Thalassiossira nordenskioeldii and Skeletonema costatum. A large portion of carbon incorporated by the colonial stage of Phaeocystis spp. is released extracellularly, in particular in stationary colonies. This extracellular production can be modeled as a function of irradiance, as for carbon incorporation. In Phaeocystis spp., cellular and extracellular carbon incorporation represent different uptake rates, indicating the formation of two distinct carbon pools. The release of extracellular carbon by polar Phaeocystis spp. was not a constant fraction of total production over the irradiance range used. We observed little extracellular carbon production by cells at high irradiance, and maximal rates were observed at intermediate irradiance. Newly incorporated carbon that accumulates in the mucilage of the colonial stage of antarctic Phaeocystis sp. during photosynthesis was not reutilized for cellular growth during the dark period, as observed for temperate clones. In contrast, only a minor fraction of the radiocarbon incorporated by the diatoms was released extracellularly for all growth stages. The production of DMS was an order of magnitude higher for Phaeocystis spp. than for diatoms. The chlorophyll-specific production of DMS and DMSP (dimethylsulphoniopropionate, the precursor to DMS) by Phaeocystis spp. showed a hyperbolic response to irradiance, while arctic diatoms (weak or non-producers of DMS), on the other hand, did not show any light-dependency of DMS production. An inverse relationship between DMS and DMSP production in stationary clones of arctic P. pouchetii was observed, but not for the exponentially growing antarctic clone. Stationary colonies also had higher DMS and dissolved DMSP production rates than exponentially growing ones. These relationships can be extrapolated to the field in areas where Phaeocystis spp. dominates.     

Organic solute accumulation in osmotically-stressed Enteromorpha intestinalis

The role of organic solutes in the osmotic adjustment processes of the marine macroalga Enteromorpha intestinalis (L.) Link was investigated in 1986, using fresh samples collected from mid-shore rock pools at Tayport, Fife, Scotland. Natural-abundance ¹³C nuclear magnetic-resonance spectroscopy revealed -dimethylsulphoniopropionate (DMSP) to be the only major low molecular weight organic osmolyte present. However, on transfer to a hypersaline medium (300% sea water; 100%=35 S), tissue sucrose and proline levels increased markedly, while DMSP remained constant. Recovery of optimal photosynthetic activity and increases in inorganic ion levels occurred over a similar time scale to the changes in sucrose and proline (within 48 h), indicating that these two organic solutes are involved in hyperosmotic adjustment in E. intestinalis while DMSP is not. Freshly-collected plants transferred to 300% sea water medium in the dark showed no significant increases in organic osmolytes. In contrast, starch-enrichment (16 d continuous illumination) led to enhanced synthesis of sucrose and proline in the light and in darkness, but tissue DMSP levels showed no variation throughout. These observations suggest that DMSP is not involved in short-term osmoacclimation in E. intestinalis.     

Succession and growth limitation of phytoplankton in the Gulf of Bothnia (Baltic Sea)

A one year field study of four stations in the Gulf of Bothnia during 1991 showed that the biomass was ca. two times, and primary productivity ca. four times, lower in the north (Bothnian Bay) than in the south (Bothnian Sea) during the summer. Nutrient addition experiments indicated phosphorus limitation of phytoplankton in the Bothanian Bay and the coastal areas in the northern Bothnian Sea, but nitrogen limitation in the open Bothanian Sea. A positive correlation between the phosphate concentration and the production/biomass ratio of phytoplankton was demonstrated, which partly explained the differences in the specific growth rate of the phytoplankton during the summer. Differences in photosynthetic active radiation between the stations also showed a covariation with the primary productivity. The relative importance of nutrient or light limitation for photosynthetic carbon fixation could not, however, the conclusively determined from this study. Marked differences in phytoplankton species composition from north to south were also observed. The number of dominating species was higher in the Bothnian Sea than in the Bothnian Bay. The distribution of some species could be explained as due to nutrient availability (e.g. Nodularia spumigena, Aphanizomenon sp.), while salinity probably limits the distribution of some limnic as well as marine species. The potentially toxic phytoplankton N. spumigena, Dinophysis acuminata and Chrysochromulina spp. were common in the Bothnian Sea but not in the Bothnian Bay. The pico- and nanoplankton biomass during late summer was higher than previously reported due to a revised carbon/volume ratio.     

Assessment of simultaneous uptake of nitrogenous nutrients (15N) and inorganic carbon (13C) by natural phytoplankton populations

The simultaneous uptake of nitrogenous nutrients and inorganic carbon was measured in shipboard incubations of natural phytoplankton populations, using tracer additions of ¹³C-bicarbonate and ¹⁵N-labelled nitrogenous substrates. From March 1991 through March 1992, three stations on the Scotian Shelf (eastern Canada) were sampled monthly at ten depths in the euphotic zone. Additions of labelled nitrogen compounds ranged between 0.5 and 98% of ambient concentrations. Most of the C/N (at/at) uptake ratios were lower than the Redfield ratio, suggesting that nitrogen was not limiting. The fixation of carbon with and without addition of nitrate, ammonium or urea was generally similar. Some samples presented significant differences in carbon uptake rate between the four treatments, but these differences were not related to nitrogen enrichment (percent or nitrogen species). Given these results, the double-labelling method appears to be a reliable tool for measuring the simultaneous uptake of carbon and nitrogen by natural phytoplankton.     

Impact of UV-B radiation on uptake of 15N-ammonia and 15N-nitrate by phytoplankton of the Wadden Sea

Natural marine phytoplankton populations from the German Wadden Sea and unialgal cultures of the haptophycean Phaeocystis pouchetii were tested in 1989 under controlled UV-B stress conditions. Assimilation of ¹⁵N-nitrate in phytoplankton consisting mainly of P. pouchetii, or in pure cultures of this alga, was found to be very sensitive to enhanced UV-B dosage in comparison ¹⁵N-ammonia uptake. In contrast, in phytoplankton samples containing Ceratium spp., Coscinodiscus sp., Noctiluca sp. or others, rate of ¹⁵NO₃ ^- uptake was higher and only slightly affected by UV-B irradiance compared to the P. pouchetii sample. UV-B inhibitory effect on uptake of inorganic nitrogen by P. pouchetii was more pronounced under strong white-light conditions and after a UV-B pre-illumination period of several hours than under low white light. Pools of glutamine and alanine decreased after UV-B exposure. Results are discussed with reference to the damaging effects of white light and UV-B on nitrogen metabolism.     

The effect of nitrogen deficiency,methionine and inhibitors of methionine metabolism on the DMSP contents of Tetraselmis subcordiformis (Stein)

The unicellular alga Tetraselmis (Platymonas) subcordiformis (Prasinophyceae) displayed a two-phase increase in DMSP (dimethylsulfoniopropionate) contents in response to nitrogen deficiency. In the first phase, DMSP pools increased by 75% within 24 h. The second increase in DMSP was observed after 14 d and was accompanied by cyst formation. Methionine metabolism was involved in DMSP accumulation. A 2.6-fold increase of the DMSP pool could be induced by supplementing a complete ASP (artifical seawater Provasoli) medium with 100 M L-methionine. Inhibition of methionine utilization by T. subcordiformis caused by the addition of S-adenosylhomocystein, a competitive inhibitor of S-adenosylmethionine-dependent transmethylases, resulted in a 24% increase of DMSP contents after 7 h. DMSP accumulation after hyperosmotic shock was significantly delayed when methionine liberation from proteins was inhibited by a set of protease inhibitors. A model for the mechanism of DMSP accumulation is presented on the basis of these data and additional information from the literature. It is proposed that methionine availability in T. subcordiformis determines the rate of DMSP synthesis.     

Interspecific differences in the photosynthetic carbon metabolism of marine phytoplankton

Six species of marine phytoplankton of different sizes and taxonomic categories were grown in microcosms under identical experimental conditions; the species cultured were: Pavlova lutheri (Prymnesiophyceae), Dunaliella tertiolecta (Chlorophyceae), Phaeodactylum tricornutum (Baciollariophyceae), Eutreptiella sp. (Euglenophyceae), Alexandrium tamarense (Dinophyceae), and Phaeocystis pouchetii (Prymnesiophyceae). The photosynthetic carbon metabolism of these phytoplankton was studied throughout the exponential and lag phases of growth after nutrient depletion. The relative incorporation of carbon into protein was positively correlated with phytoplankton growth, while carbon assimilation into low molecular weight metabolites (LMWM) and storage products, i.e., lipid and polysaccharides, generally increased under nutrient-limiting conditions. Clear taxonspecific differences were observed in the proportions of carbon incorporated into cell constituents. A significant linear relationship was consistently found between the relative synthesis of protein to LMWM, and both the production normalised to chlorophyll (P:B) and the phytoplankton growth rate. However, ANCOVA revealed significant, interspecific differences in these relationships.     

Carbon and nitrogen dynamics during growth and degradation of phytoplankton under natural surface irradiance

Under conditions of natural irradiance, the development and decline of a flagellate-dominated phytoplankton population was followed in a coastal North Atlantic pond over a 3 d period in summer 1986. Irradiance negatively affected phytoplankton biomass estimated as chlorophyll a, which decreased during the day at photosynthetically available radiation (PAR) levels above 600 to 1000 mol m^-2s^-1; chlorophyll a increased at PAR values below this threshold. In addition, an inverse relationship was found between changes in chlorophyll a and changes in dissolved inorganic nitrogen, indicating synthesis of nitrogenous biomass mainly at night and degradation mainly during the day, with intense exchanges of material between the particulate and dissolved nitrogen fractions. The natural abundance of ¹³C in particulate matter increased initially, and then remained constant, and was controlled mainly by the ratio -carboxylases activity: ribulose biphosphate carboxylase activity. The hypothesis that the latter enzyme is broken down under high irradiance and is partly responsible for increases in external dissolved nitrogen was rejected.     

Change in the uptake and cellular Si:N ratio in diatoms responding to the ambient Si:N ratio and growth phase

A change in the Si:N ratio of diatom cells during growth was examined for Chaetoceros socialis and Thalassiosira sp., with different initial silicate to nitrate (Si:N) ratios in the media. During exponential growth, C. socialis assimilated silicate and nitrate with a molar ratio of 0.5, independent of the ratio in the media, but after the depletion of nitrate, silicate continued to be taken up, and the Si:N ratio in the stationary phase increased to 2 as a function of the Si:N ratios in the media. In contrast, the ratio of silicate to nitrate taken up by Thalassiosira sp. increased with an increase in the Si:N ratio in the media. The Si:N ratio in the cells during the stationary phase increased in response to an increase in this ratio in the media. The Si:chl  a ratio also increased with the increase in the initial Si:N ratio in the media, while the N:chl  a ratio did not change to a great extent, indicating the changes in the cellular Si:N ratio was derived from changes in the Si content of the cells. These results indicated that the cellular Si:N ratio changed with the Si:N ratio in the medium, and the Si:N uptake ratio during the growth phase was different depending on diatom species. Thus, the dominance of different diatom species may affect nutrient composition and dynamics in the ocean.     

Ammonium regeneration by microzooplankton in the Oslofjord

Concentrations of dissolved inorganic nitrogen compounds above the pycnocline in the Oslofjord are very low in the summer, with turnover times of the inorganic N pools of no more than a few hours. To investigate the possibility that continued phytoplankton growth in the summer depends on ammonium excretion by microzooplankton, rates of NH ₄ ⁺ regeneration and assimilation were measured by a ¹⁵N isotope dilution method. Daytime regeneration rates at 0–2 m depth were 0–28% of the calculated assimilation rates at ambient NH ₄ ⁺ concentrations. Regeneration was faster during a dinoflagellate bloom in August than in mixed diatom-dinoflagellate blooms in June and September. Most of the NH ₄ ⁺ appeared to be produced by juvenile copepods, rotifers, tintinnids, and heterotrophic dinoflagellates in the size fraction 45–200 m.     

Effect of food type and ration on growth of juvenile Capitella sp. I (Annelida: Polychaeta): macro- and micronutrients

Growth rates of juvenile Capitella sp. I were determined on different rations of six food types: Gerber's mixed cereal, TetraMin fish flakes, benthic diatoms, Ulva sp., spring detritus, and summer detritus. A simple growth model estimated maximum growth rate and maintenance ration for each food. There were differences in the growth response among foods relative to nitrogen content. As juveniles increased in size, differences in growth between foods became more pronounced. For all juveniles, growth rates were correlated with levels of the amino acids histidine, phenylalanine, threonine, and valine, and the polyunsaturated fatty acid 20:5w3; correlations with histidine and phenylalanine levels were the most significant. Regressions of growth rates as a function of these two amino acids suggest a daily maintenance ration of 300 pg histidine and phenylalanine mg^-1 nitrogen biomass. Juvenile worms grew on spring detritus but not on summer detritus, indicating the probable importance of micronutrients (polyunsaturated fatty acids, amino acids) for growth of juvenile Capitella sp. I in the field.     

Dimethylsulfoniopropionate and dimethylsulfide in the marine flatworm Convoluta roscoffensis and its algal symbiont

 A distinct smell of dimethylsulfide (DMS) was noted at the edge of the intertidal mudflat of Marennes-Oléron Bay, at the French Atlantic coast, where dense populations of the marine flatworm Convoluta roscoffensis Graff (Platyhelminthes: Turbellaria) were present. DMS is the cleavage product of dimethylsulfoniopropionate (DMSP). DMSP was shown to be present in high amounts in sediment containing the flatworm as well as in axenic cultures of the symbiotic alga Tetraselmis sp. that was isolated from the flatworm. In untreated sediment samples containing C. roscoffensis the concentration of DMS was as high as ∼55 μmol l⁻¹ sediment, and in samples that were fixed with glutaraldehyde the concentration of DMS was even three orders of magnitude higher (∼66 mmol l⁻¹ sediment). This rapid cleavage of DMSP to DMS in fixed samples was unexpected. Pure DMSP was stable in glutaraldehyde, and it was therefore concluded that a DMSP-lyase was responsible for cleavage in the field samples. The isolated symbiotic alga, Tetraselmis sp., did not show DMSP-lyase activity, indicating that DMSP-lyase may have been present in the flatworm, although the role of bacteria could not be excluded. The Chl a-specific DMSP content of C. roscoffensis (∼200 mmol g⁻¹) was much higher than that of Tetraselmis sp. (∼30 mmol g⁻¹). Possibly, DMSP was not only present in the symbiotic alga, but was also incorporated in the body tissue of the flatworm. It remains unclear what the function of DMSP is in C. roscoffensis. In Tetraselmis sp., but not in C. roscoffensis, DMSP increased with increasing salinity. It was concluded that salinity probably does not play an important role in the dynamics of DMSP and DMS in sediment containing C. roscoffensis. Received: 21 January 2000 / Accepted: 29 August 2000     

Photosystem II photoinhibition and altered kinetics of photosynthesis during nutrient-dependent high-light photoadaptation in Gonyaulax polyedra

Gonyaulax poledra Stein was transferred at different cell densities from increasingly nutrient-limited low-light (LL, 80 E m^-2 s^-1) batch-cultures to high-light (HL, 330 E m^-2 s^-1) growth conditions. Several age-dependent differences in HL-adaptation strategies were apparent. Short-term (3h) susceptibility to photosynthetic photoinhibition increased with culture age, with light-limited rates of photosynthesis exhibiting greater photosuppression than light-saturated rates at all stages of growth. These shortterm changes were not accompanied by photobleaching of chlorophyll but were directly related to age-dependent photoinactivation of Photosystem II electron-transport rates. The capacity of electron transport by Photosystem I was only slightly affected. Prolonged exposure of LL log-phase cells to HL conditions did induce photobleaching of chlorophyll associated with increased cell volume, a transient decrease of organic carbon and nitrogen content, enhanced cellular-, carbon-and chlorophyll-based rates of light-saturated photosynthesis (P _max) and suppressed cellular rates of light-limited photosynthesis. As a result, the density of LL log-phase cells doubled and their cellular photosynthetic performance nearly tripled within 1 d of HL exposure while cellular respiratory demands remained unchanged. By contrast, prolonged HL incubation of LL stationary populations induced a transitory burst in cell division and a large reduction in cell volume, leading to a short-term increase in volume-based organic carbon and nitrogen content. Despite reduced cell volume and lowered carbon demand, the cellular-, carbon-and chlorophyll-based rates of P _max in nondividing populations fell by 64, 48 and 27%, respectively, over a 4 d exposure to HL, while light-limited rates were almost fully suppressed within 1 d and chlorophyll a content was reduced by 56%. As a result, the photosynthetic performance of LL-aged cells declined immediately under HL conditions. Addition of inorganic nutrients to LL stationary cultures at the time of HL transfer led to immediate and complete suppression of photosynthesis and cell lysis within 1 d. Addition of nutrients following transfer to HL induced cell responses intermediate to those described for LL log and aged cells exposed to HL. Results support the view that declining nutrient-status impairs HL photoadaptive responses in phytoplankton populations and that the rate and pattern of photoadaptive responses may be used as physiological growth indicators in field studies. The study was conducted from March 1981 to May 1983.     

DMSP-lyase activity in five marine phytoplankton species: its potential importance in DMS production

Activity of DMSP-lyase, which cleaves dissolved DMSP (henceforth DMSP_d-lyase), was examined in five axenically cultured phytoplankton species, including both DMSP-producing and non-DMSP-producing species. High DMSP_d-lyase activity was found in two DMSP producers, Heterocapsa triquetra strain NIES-7 and Scrippsiella trochoidea strain NIES-369 (Dinophyceae). The DMS production rates at 100 nM DMSP_d were 0.5 fmol cell⁻¹ min⁻¹ for H. triquetra and 0.3 fmol cell⁻¹ min⁻¹ for S. trochoidea. In a non-DMSP producer, Heterosigma akashiwo strain NIES-6 (Raphidophyceae), the DMSP_d-lyase activity was not found. Two DMSP-producing Prymnesiophyceae species, Isochrysis galbana strain CCMP-1323 and Gephyrocapsa oceanica strain NIES-353, did not show any obvious activity either, in contrary to other authors' findings on Phaeocystis sp., another DMSP-producing Prymnesiophyceae species. The comparison of the DMSP_d-lyase activity of the two Dinophyceae species with bacterial DMSP consumption and DMS production activity in Tokyo Bay showed that the DMSP_d-lyase activity of H. triquetra and S. trochoidea could be an important mechanism for DMS production during their blooms. Received: 9 April 1999 / Accepted: 10 December 1999     

Dimethylsulfide and the algaPhaeocystis pouchetii in antarctic coastal waters

The concentrations of dimethylsulfide (DMS), dimethylsulfoniopropionate (DMSP) and dimethylsulfoxide (DMSO) were measured in water collected from the Southern Ocean 10 km offshore from Davis Station, Antarctica, during the period May 1987 to January 1988, inclusive. During winter and spring, when the sea-ice was up to 1.9 m thick, DMS, DMSP and DMSO concentrations were low (0.2 to 1.5 nM), as were phytoplankton numbers. The maximum concentration of the sulfur compounds generally occurred in the top 10 m of the water column. DMS levels rose dramatically from early December onwards, reaching a peak of 290 nM at a depth of 15 m in January. This concentration is higher than reported elsewhere in the ocean. These high concentrations occurred at the same time as a bloom of the algaPhaeocystis pouchetii. A significant correlation occurred between DMS concentration and cell numbers of the alga. The ratio of DMS concentration to the number of cells of the alga was considerably higher than reported for blooms of this species elsewhere in the ocean. Up to 10% of the total global flux of DMS to the atmosphere may emanate from antarctic seas.     

Spatial variation in dimethylsulfoniopropionate (DMSP) production in <Emphasis Type="Italic">Ulva lactuca</Emphasis> (Chlorophyta) from the Northeast Pacific

Although dimethylsulfoniopropionate (DMSP) has a variety of functions in marine macroalgae including that of a cryoprotectant, an osmolyte, a way to remove excess sulfur and energy, an antioxidant, and an allelopathic precursor, the latter two functions are believed to be the most important in Ulva lactuca L. (=U. fenestrata) in intertidal populations on the coast of Washington state, USA. The present study found significant variation in DMSP concentrations among U. lactuca collected in May 2005 from six sites ranging from 47°54.45′N (Possession Point, Whidbey Island, WA, USA) to 48°30.55′N (Shannon Point Beach, Anacortes, WA, USA), and also among individuals within sites, and among tissues (basal tissues near the holdfast, middle of the blades, and tips). Concentrations ranged from 37 to 224 μmol g⁻¹ fresh mass (FM). In several 10-day experiments between July 2001 and August 2004 with U. lactuca collected from several places on the coast of Washington, the effects of nutrient level (DIN), light intensity and wavelength, and grazing by the herbivorous gastropod Lacuna vincta, were examined. None of these manipulations resulted in DMSP concentrations that differed significantly from controls, and variance in DMSP concentrations within each experiment was very low. Although DMSP concentrations in U. lactuca may be affected by factors not tested in these experiments, it is also possible that the observed spatial differences reflect constitutive genotypic or phenotypic differences among geographically separated U. lactuca populations or among cryptic Ulva species. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Growth and development of Calanus spp. (Copepoda) during spring phytoplankton succession in the North Sea

Experiments were carried out to determine growth and development rates of the herbivorous copepod Calanus finmarchicus (Gunnerus) under natural conditions during the phytoplankton spring bloom in the northern North Sea. From 28 April to 25 May 1983 copepodite stages I, IV and V were incubated for a 3-d period on board a ship in vessels with naturally occurring phytoplankton or cultured algae as food. Highest rates of growth and development were achieved while the diatom Chaetoceros sp. was the dominant phytoplankton organism. These rates decreased considerably when this chain-forming diatom was succeeded after one week by the small-celled diatom Thalassiosira conferta. Again one week later, during the bloom of the succeeding colonial microflagellate Corymbellus aureus, copepodite stage IV still managed to maintain moderate rates of growth and development, but these rates dropped to almost zero in CV, suggesting the start of a resting stage. Nevertheless, brood collected from this generation and from Calanus helgolandicus (Claus) was raised in the laboratory to the adult stage at high speed. Since temperature and the total phytoplankton concentration in the sea remained almost constant it seems that the retardation and arrestment of growth and development were an immediate response to a qualitative change of the food composition related to the successive blooms of different algal species.     

Nocturnal synthesis and diurnal degradation of phytoplankton biomass in surface waters

Natural populations of phytoplankton were collected near the Bay of Bourgneuf, France, in spring 1982, and were subjected to natural surface irradiance outdoors. They exhibited exponential growth on time scales of a week, but significant decreases in biomass indicators such as chlorophyll a and particulate nitrogen were observed during daytime. At night, these decreases were more than compensated by increases in the same biomass variables, which could double over 12 h of darkness. These features are characteristic of phytoplankton populations in surface waters which cannot escape high irradiances, and may be representative of situations in incubation bottles held at fixed depths near the surface. Under such conditions, a decrease in biomass during daytime should not necessarily be interpreted as irreversible damage unless growth measurements are carried out over the following night hours to check for possible recovery.     

Production of DMSP and DMS during a mesocosm study of an Emiliania huxleyi bloom: influence of bacteria and Calanus finmarchicus grazing

We investigated the influence of bacteria and metazooplankton on the production of dimethylsulfoniopropionate (DMSP) and dimethylsulfide (DMS) during blooms of Emiliania huxleyi (Lohmann) Hay and Mohler in seawater mesocosms. The phytoplankton succession was marked by the rapid collapse of an initial Skeletonema costatum (Greville) Cleve bloom followed by a small E. huxleyi bloom. The collapse of the diatom bloom was accompanied by an increase in concentrations of dissolved DMSP (DMSP_d) and bacterial abundance and activity (as determined by the thymidine incorporation technique). The increase in bacterial activity was followed by a rapid decrease in DMSP_d concentrations which remained low for the rest of the experiment, even during the subsequent collapse of the E. huxleyi blooms. The absence of DMSP_d and DMS peaks during the declining phase of the E. huxleyi blooms was attributed to the high bacterial activity prevailing at that time. The influence of metazooplankton grazing on DMSP and DMS production was investigated by adding moderate (24 mg dry weight m^-3) and high (520 mg dry weight m^-3) concentrations of Copepodite Stage V and adults of Calanus finmarchicus to two of four filtered (200 m mesh net) enclosures during the E. huxleyi blooms. The addition of C. finmarchicus, even in high concentrations, had no apparent effect on the dynamics of E. huxleyi, suggesting that the copepods were not grazing significantly on nanophytoplankton. The addition of copepods in high concentrations favored an accumulation of chlorophyll a and particulate DMSP. These results suggest that copepods were preying on the herbivorous microzooplankton which, in turn, was controlling the biomass of nanophytoplankton. DMS production was also enhanced in the enclosure with maximum metazooplankton biomass, suggesting that the grazing of C. finmarchicus on microzooplankton containing DMSP may contribute to DMS production. These results provide strong support to the emerging idea that bacteria and metazooplankton grazing play a dominant role in determining the timing and magnitude of DMS pulses following phytoplankton blooms.