Environmental constraints on the production and removal of the climatically active gas dimethylsulphide DMS and implications for ecosystem modelling

Jacqueline Stefels • Michael Steinke • Suzanne Turner • Gill Malin • Sauveur Belviso

Received: 14 June 2006/Accepted: 5 August 2006/Published online: 27 April 2007 © Springer Science+Business Media B.V. 2007

Abstract Seawater concentrations of the climate-cooling, volatile sulphur compound dimethylsul-phide (DMS) are the result of numerous production and consumption processes within the marine ecosystem. Due to this complex nature, it is difficult to predict temporal and geographical distribution patterns of DMS concentrations and the inclusion of DMS into global ocean climate models has only been attempted recently. Comparisons between individual model predictions, and ground-truthing exercises revealed that information on the functional

Laboratory of Plant Physiology, University of Groningen, P.O. Box 14, 9750 AA Haren, The Netherlands e-mail: [email protected]

M. Steinke • S. Turner • G. Malin Laboratory for Global Marine and Atmospheric Chemistry, School of Environmental Sciences, University of East Anglia, Norwich NR4 7TJ, UK

S. Belviso

Laboratoire des Sciences du Climat et de l'Environnement, UMR CEA-CNRS 1572, CEN/ Saclay, Bat 709, L'Orme des Merisiers, 91191 Gif-sur-Yvette, France

Present Address: M. Steinke relationships between physical and chemical ecosystem parameters, biological productivity and the production and consumption of DMS and its precursor dimethylsulphoniopropionate (DMSP) is necessary to further refine future climate models. In this review an attempt is made to quantify these functional relationships. The description of processes includes: (1) parameters controlling DMSP production such as species composition and abiotic factors; (2) the conversion of DMSP to DMS by algal and bacterial enzymes; (3) the fate of DMSP-sulphur due to, e.g., grazing, microbial consumption and sedimentation and (4) factors controlling DMS removal from the water column such as microbial consumption, photo-oxidation and emission to the atmosphere. We recommend the differentiation of six phytoplankton groups for inclusion in future models: eukaryotic and prokaryotic picoplankton, diatoms, dinoflagellates, and other phytoflagellates with and without DMSP-lyase activity. These functional groups are characterised by their cell size, DMSP content, DMSP-lyase activity and interactions with herbivorous grazers. In this review, emphasis is given to ecosystems dominated by the globally relevant haptophytes Emiliania huxleyi and Phaeocystis sp., which are important DMS and DMSP producers.

Emiliania huxleyi • Functional groups • Phaeocystis

Department of Biological Sciences, University of Essex, Wivenhoe Park, Colchester CO4 3SQ, UK


Dimethylsulphide (DMS) is a semivolatile organic sulphur compound that accounts for 50-60% of the total natural reduced sulphur flux to the atmosphere, including emissions from volcanoes and from vegetation (Andreae 1990; Bates et al. 1992; Spiro et al. 1992). By providing 95% of the flux to the atmosphere, the oceans are the main source for DMS, with estimates of its emission ranging between 15 and 33 Tg S y^1 (Kettle and Andreae 2000). In the late 1980s, the hypothesis that DMS is involved in the biological regulation of global climate was put forward (Bates et al. 1987; Charlson et al. 1987). It is, however, only recently that DMS has been incorporated in global climate models (e.g., Aumont et al. 2002; Bopp et al. 2003, 2004; Gabric et al. 2004; Kettle and Andreae 2000; Simo and Dachs 2002). After emission to the atmosphere, this volatile sulphur compound is oxidised to sulphur dioxide (SO2) and other products. From SO2, non-sea-salt (nss) sulphate is produced, which can form sulphate (SO4~) particles that act as condensation nuclei for water vapour. These nuclei affect the radiative properties of the atmosphere and clouds, with implications for climate. Higher numbers of condensation nuclei will deflect more incoming solar radiation back into space and thereby reduce the temperature on earth. The hypothesis that this process may modulate the greenhouse effect of increased anthropogenic CO2 input to the atmosphere, was indirectly supported by the modelling results of the effect of anthropogenic SO2 input to the atmosphere (Andreae et al. 2005; Mitchell et al. 1995). Although this study gave a rough indication of the counteracting effects of atmospheric SO2 and CO2, a quantitative understanding of all sources and sinks of atmospheric aerosols is still lacking (Andreae and Crutzen 1997).

Currently, anthropogenic SO2 production exceeds natural SO2 production by a factor of 3 (Bates et al. 1992), but the impact of the former on aerosol production is largely confined to industrialised areas of the Northern Hemisphere. The oceans, on the other hand, cover approximately 70% of the Earth's surface and much of this area is remote from man-made atmospheric contaminants. Consequently, the exchange of marine DMS is of high regional importance and may affect climate globally. Since the publication of a global inventory of DMS data by

Kettle and co-authors (Kettle et al. 1999), it has become possible to include DMS in global climate models. Indeed, recent model calculations have shown that in the Southern Hemisphere, where anthropogenic sulphate emission is low, DMS plays a major role in the production of atmospheric nss-sulphate (Gondwe et al. 2003). Gondwe et al. calculated that the contribution of DMS to the total (global) atmospheric nss-sulphate burden is 18% and that it shows significant regional and temporal differences; e.g., in the Southern Hemisphere its annual contribution is 43% and over the Southern Ocean it is in excess of 80% during summer.

In one of the early attempts to add DMS in a global ocean climate model, Bopp et al. (2003) showed that a doubling of the atmospheric CO2 concentration resulted in a reduction of the DMS flux at low latitudes and in enrichment at mid-latitudes. Thus, depending on the sign of the chance in DMS flux, the subsequent climate forcing by sulphur products could either alleviate or amplify the greenhouse effect (Bopp et al. 2004). The inclusion of DMS in the model was achieved by coupling the production of dimethylsulphoniopropionate (DMSP) and its conversion to DMS with the trophic status of the ecosystem, which in turn was based on the silica ratio, defined as the local simulated production of biogenic silica relative to the maximum production and is related to the proliferation of diatoms. It is computed from local silica concentrations and a variable Si:C ratio, which depends on the silica concentration. Although simplistic in its ecological approach, this study showed that an increased CO2 concentration doesnot necessarily result in increased DMS production that may counterbalance the greenhouse effect, as suggested by the Charlson-Lovelock-Andreae-Warren (CLAW) hypothesis (Charlson et al. 1987). Clearly, an improved understanding of the biological processes is necessary to address the role of DMS in climate feedback mechanisms.

The production of DMS is almost exclusively through biogenic processes and shows strong seasonal and latitudinal variation (Kettle et al. 1999). DMS mainly results from the enzymatic cleavage of DMSP, a compound that is produced in several groups of marine phytoplankton. A complex network of production and consumption pathways of both DMSP and DMS involves most of the microbial food web (Fig. 1) and determines the concentration of

DMS in surface water and consequently its flux to the atmosphere (Malin and Kirst 1997). Physical and chemical ecosystem parameters all affect this network, potentially resulting in dramatic shifts in the DMS flux to the atmosphere. Although our knowledge on the qualitative aspects of the marine sulphur cycle has improved considerably during the past two decades, it is still difficult to quantify the effects of controlling factors on the various pathways.

Ecosystem modelling provides a tool for investigating how the DMS concentration and subsequently its flux to the atmosphere are regulated and what the most critical processes are. In a recent review on DMS and DMSP ecosystem models, Vezina (2004) concluded that although all current models will greatly benefit from improvements to the underlying ecosystem model, the quantitative understanding of the processes that drive variations in DMS and DMSP

Biomes Concept Map Answer Key

Fig. 1 Schematic representation of the processes and pools involved in the marine biogeochemical cycling of DMSP and DMS. Dominant role of functional groups in the different processes is indicated by coloured ellipses: green, phytoplank-ton; blue, zooplankton; red, bacteria; black, abiotic factors.

CCN, cloud-condensation nuclei; DOM, dissolved organic material; DMSO, dimethyl sulphoxide; MeSH, methanethiol; MPA, mercaptopropionate; MMPA, methylmercaptopropio-nate; MSA, methanesulphonic acid

Fig. 1 Schematic representation of the processes and pools involved in the marine biogeochemical cycling of DMSP and DMS. Dominant role of functional groups in the different processes is indicated by coloured ellipses: green, phytoplank-ton; blue, zooplankton; red, bacteria; black, abiotic factors.

CCN, cloud-condensation nuclei; DOM, dissolved organic material; DMSO, dimethyl sulphoxide; MeSH, methanethiol; MPA, mercaptopropionate; MMPA, methylmercaptopropio-nate; MSA, methanesulphonic acid

[DMS(P)] quotas and microbial yields is still too limited. Such insights are needed to inform laboratory and field studies and aid us in the development of more robust DMS(P)-modules within ecosystem models. During the past decade, many excellent reviews have been written on several aspects of the marine sulphur cycle. One of the emerging pictures is that this cycle is not only of interest for global climate, but that DMS and DMSP are compounds which are central to the microbial food web in their own right. The purpose of this review is not so much to reiterate these reviews, but rather to use pertinent information from them in an attempt to assist the development of parameterisations for DMSP and DMS modelling.

In this review, much attention has been given to two specific algal haptophyte taxa: Phaeocystis sp. and Emiliania huxleyi. These algae are well known as prolific producers of DMS and DMSP and their blooms can cover extensive areas in neritic and open ocean waters, respectively. Due to an increased interest to define the phytoplankton realm in models in more detail, we have tried to find unifying processes, but most published information is from Phaeocystis and E. huxleyi. Since our main goal is to provide an understanding of the complexity of the system, we have chosen to describe the various processes independently, even though this may have resulted in some repetition of observations. The level of detail may not be equal throughout the paper, but is a reflection of the current state of knowledge, our judgment of the potential impact of a specific process on the marine sulphur cycle and the assignment of different processes and functional groups in recent (complex) ecosystem models (e.g., Archer et al. 2004). We have tried to be as concise as possible, without losing information necessary for a holistic description. In order to evaluate the relative importance of individual pathways, we have provided the reader with an educated guess of the quantitative aspects, whenever possible. Obviously, when describing the different processes in detail, one comes across many gaps in knowledge. We have therefore taken the opportunity to highlight these gaps and make recommendations for future research.

Factors controlling DMSP production

A direct coupling of DMSP production with primary production would be ideal for modelling. However, there is no straightforward relationship since DMSP production is confined to a limited number of algal taxa. A further complicating factor is that the physiological conditions of the algal cells affect DMSP production. As a result, there is no definitive global relationship between algal biomass parameters such as chlorophyll-a and algal DMSP. Many ecosystem models are expressed in pools of carbon or nitrogen, whereas many global climate models are more often expressed in units of carbon. These models would benefit from conversion factors to describe the particulate DMSP pool, hence we choose to estimate DMSP:C ratios (on a molar basis) from literature data whenever possible.

Species composition

From an ecosystem perspective, species composition is the factor that affects community-DMSP production the most. Keller et al. (1989) made an extensive inventory of the DMSP content of 123 clones of marine phytoplankton, analysed during mid-exponential growth in nutrient replete media, and concluded that the major production of DMSP is found in a limited number of species, which mainly belong to the classes of Haptophyceae (=Prymnesi-ophyceae) and Dinophyceae (dinoflagellates). However, some members of the Chrysophyceae and Bacillariophyceae (diatoms) can also produce significant amounts of DMSP. In order to use this knowledge in models, we have recalculated published data of cellular DMSP concentrations to provide DMSP-to-carbon (DMSP:C) ratios, that can be used to estimate DMSP production in blooms of different taxonomic groups (Table 1). In addition, we have provided DMSP-to-chlorophyll-a (DMSP:chl-a) ratios, since several global models use satellite-derived chlorophyll-a data multiplied with a trophic status factor, as a proxy for particulate DMSP (Anderson et al. 2001; Aumont et al. 2002; Bopp et al. 2003; Simo and Dachs 2002). The carbon-to-chlorophyll-a conversion factor we used for this calculation (60 g/g) is typical for cultures that grow under nutrient-replete conditions and saturating light intensities (Geider 1987). Variations in abiotic factors in the field will, however, have a strong impact on this ratio and, as will be discussed in the following sections, on the DMSP:C and DMSP:chl-a ratios.

Table 1 Mean DMSP:C ratios, proportion of cell carbon composed of DMSP and DMSP:chlorophyll-a ratios in species groups, with the standard deviation in brackets. Data are recalculated from published data. Carbon per cell was calculated from cell volumes, according to the formula given by

Menden-Deuer and Lessard (2000): diatoms: pgC/ cell = 0.288 x cell volume (pm3)0 811 ; all other algae: pgC/

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