Picophytoplankton account for most of the marine (sub-)tropical phytoplankton biomass and primary productivity. The contribution to biomass among plankton functional types (PFTs) could shift with climate warming, in part as a result of different physiological responses to temperature. To model these responses, Eppley's empirical relationships have been well established. However, they have not yet been statistically validated for individual PFTs. Here, we examine the physiological response of nine strains of picophytoplankton to temperature; three strains of picoprokaryotes and six strains of picoeukaryotes. We conduct laboratory experiments at 13 temperatures between -0.5 and 33 • C and measure the maximum growth rates and the chlorophyll a to carbon ratios. We then statistically validate two hypotheses formulated by Eppley in 1972: The response of maximum growth rates to temperature (1) of individual strains can be represented by an optimum function, and (2) of the whole phytoplankton group can be represented by an exponential function Eppley (1972). We also quantify the temperature-related parameters. We find that the temperature span at which growth is positive is more constrained for picoprokaryotes (13.7-27 • C), than for picoeukaryotes (2.8-32.4 • C). However, the modeled temperature tolerance range ( T) follows an unimodal function of cell size for the strains examined here. Thus, the temperature tolerance range may act in conjunction with the maximum growth rate to explain the picophytoplankton community size structure in correlation with ocean temperature. The maximum growth rates obtained by a 99th quantile regression for the group of picophytoplankton or picoprokaryotes are generally lower than the rates estimated by Eppley. However, we find temperature-dependencies (Q 10 ) of 2.3 and of 4.9 for the two groups, respectively. Both of these values are higher than the Q 10 of 1.88 estimated by Eppley and could have substantial influence on the biomass distribution in models, in particular if picoprokaryotes were considered an independent PFT. We also quantify the increase of the chlorophyll a to carbon ratios with increasing temperature due to acclimation. These parameters provide essential and validated physiological information to explore the response of marine ecosystems to a warming climate using ocean biogeochemistry models.

Several methanogenic pathways in oxic surface waters were recently discovered, but their relevance in the natural environment is still unknown. Our study examines distinct methane enrichments that repeatedly occur below the thermocline during the summer months in the central Baltic Sea. In agreement with previous studies in this region, we discovered differences in the methane distributions between the Western and Eastern Gotland Basin, pointing to in situ methane 5 production below the thermocline in the latter (conc. CH 4 14.1 ±6.1 nM, δ 13 C CH 4 -62.9‰). Through the use of a high resolution hydrographic model of the Baltic Sea, we showed that methane below the thermocline can be transported by upwelling events towards the sea surface thus contributing to the methane flux at the sea/air interface. To quantify zooplankton-associated methane production rates, we developed a sea-going methane stripping-oxidation line to determine methane release rates from copepods grazing on 14 C-labelled phytoplankton. We found that: (1) methane production 10 increased with the number of copepods, (2) dominated zooplankton communities, and (3) methane was only produced on a Rhodomonas sp. diet, but not on a cyanobacteria diet. Furthermore, copepod-specific methane production rates increased with incubation time. The latter finding suggests that methanogenic substrates for water-dwelling microbes are released by cell disruption during feeding, 15 defecation, or diffusion from fecal pellets. In the field, particularly high methane concentrations coincided with stations showing a high abundance of DMSP-rich Dinophyceae. Lipid biomarkers extracted from phytoplankton-and copepod-rich samples revealed that Dinophyceae are a major food source of the T. longicornis dominated zooplankton community, supporting the proposed link between copepod grazing, DMSP release, and the buildup of subthermocline methane enrichments in the central Baltic Sea. 20

Abstract. Several methanogenic pathways in oxic surface waters were recently discovered, but their relevance in the natural environment is still unknown. Our study examines distinct methane (CH4) enrichments that repeatedly occur below the thermocline during the summer months in the central Baltic Sea. In agreement with previous studies in this region, we discovered differences in the methane distributions between the western and eastern Gotland Basin, pointing to in situ methane production below the thermocline in the latter (concentration of CH4 14.1±6.1 nM, δ13C CH4 −62.9 ‰). Through the use of a high-resolution hydrographic model of the Baltic Sea, we showed that methane below the thermocline can be transported by upwelling events towards the sea surface, thus contributing to the methane flux at the sea–air interface. To quantify zooplankton-associated methane production rates, we developed a sea-going methane stripping-oxidation line to determine methane release rates from copepods grazing on 14C-labelled phytoplankton. We found that (1) methane production increased with the number of copepods, (2) higher methane production rates were measured in incubations with Temora longicornis (125±49 fmol methane copepod−1 d−1) than in incubations with Acartia spp. (84±19 fmol CH4 copepod−1 d−1) dominated zooplankton communities, and (3) methane was only produced on a Rhodomonas sp. diet, and not on a cyanobacteria diet. Furthermore, copepod-specific methane production rates increased with incubation time. The latter finding suggests that methanogenic substrates for water-dwelling microbes are released by cell disruption during feeding, defecation, or diffusion from fecal pellets. In the field, particularly high methane concentrations coincided with stations showing a high abundance of DMSP/DMSO-rich Dinophyceae. Lipid biomarkers extracted from phytoplankton- and copepod-rich samples revealed that Dinophyceae are a major food source of the T. longicornis dominated zooplankton community, supporting the proposed link between copepod grazing, DMSP/DMSO release, and the build-up of subthermocline methane enrichments in the central Baltic Sea.

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Picophytoplankton dominate the phytoplankton community in wide ocean areas and are considered efficient in the acquisition of light compared to other phytoplankton groups. To quantify their photophysiological parameters we use three strains of picoprokaryotes and four strains of picoeukaryotes. We measure the acclimated response of the exponential growth rates and chlorophyll a (Chl a) to carbon ratios, as well as the instantaneous response of photosynthesis rates at 5-7 light intensities. We then use a dynamic photosynthesis model (Geider et al. 1997) and extend it with a photoinhibition term. We derive five photophysiological parameters: the maximum rate of photosynthesis (P C m ), the affinity to light (a chl ), the photoinhibition term (b chl ), the respiration rate (resp), and the maximum Chl a to carbon ratio (h max ). We show that P C m is significantly lower for picoprokaryotes than for picoeukaryotes and increases significantly with increasing cell size. In turn, a chl decreases significantly with increasing maximum growth rate (l max ). The latter finding is contrary to a previously reported relationship for phytoplankton, but agrees with theoretical assumptions based on size. The higher efficiency in light acquisition gives picoprokaryotes an advantage in light limited environments at the expense of their maximum growth rate. In addition, our results indicate that the accumulation of long-term damage through photoinhibition during acclimation is not well represented by the dynamic photosynthesis model. Hence, we would recommend to distinguish between the effects of irreversible damage (on a time scale of days) on growth rates and of reversible damage (on a time scale of minutes) on photosynthesis rates.