Increased intrusion of warming Atlantic water leads to rapid expansion of temperate phytoplankton in the Arctic

The Arctic Ocean and its surrounding shelf seas are warming much faster than the global average, which potentially opens up new distribution areas for temperate‐origin marine phytoplankton. Using over three decades of continuous satellite observations, we show that increased inflow and temperature of Atlantic waters in the Barents Sea resulted in a striking poleward shift in the distribution of blooms of Emiliania huxleyi, a marine calcifying phytoplankton species. This species' blooms are typically associated with temperate waters and have expanded north to 76°N, five degrees further north of its first bloom occurrence in 1989. E. huxleyi's blooms keep pace with the changing climate of the Barents Sea, namely ocean warming and shifts in the position of the Polar Front, resulting in an exceptionally rapid range shift compared to what is generally detected in the marine realm. We propose that as the Eurasian Basin of the Arctic Ocean further atlantifies and ocean temperatures continue to rise, E. huxleyi and other temperate‐origin phytoplankton could well become resident bloom formers in the Arctic Ocean.     

Comparison of population growth and photosynthetic apparatus changes in response to different nutrient status in a diatom and a coccolithophore

In many marine ecosystems, diatoms dominate in nutrient‐rich coastal waters while coccolithiophores are found offshore in areas where nutrients may be limiting. In lab‐controlled batch cultures, mixed‐species competition between the diatom Phaeodactylum tricornutum and the coccolithophore Emiliana huxleyi and the response of each species were examined under nitrate (N) and phosphate (P) starvation. Based on the logistic growth model and the Lotka–Volterra competition model, E. huxleyi showed higher competitive abilities than P. tricornutum under N and P starvation. For both species, cell growth was more inhibited by P starvation, while photosynthetic functions (chl a fluorescence parameters) and cellular constituents (pigments) were impaired by N starvation. The decline of photosynthetic functions occurred later in E. huxleyi (day 12) than in P. tricornutum (day 9); this time difference was associated with greater damage of the photosynthetic apparatus in P. tricornutum compared with E. huxleyi. Xanthophyll cycle pigment accumulation and the transformation from diadinoxanthin to diatoxanthin was more active in E. huxleyi than P. tricornutum, under similar N and P starvation. We concluded that E. huxleyi and P. tricornutum have different mechanisms to allocate resources and energy under nutrient starvation. It appears that E. huxleyi has a more economic strategy to adapt to nutrient depleted environments than P. tricornutum. These findings provided additional evidence explaining how N versus P limitation differentially support diatom and coccolithophore blooms in natural environments.     

The 24 hour recovery kinetics from n starvation in Phaeodactylum tricornutum and Emiliania huxleyi

The response of N (nitrate) starved cells of the diatom Phaeodactylum tricornutum and the coccolithophore Emiliania huxleyi to a pulse of new N were measured to investigate rapid cellular and photosynthetic recovery kinetics. The changes of multiple parameters were followed over 24 h. In P. tricornutum, the recovery of F_v/F_m (the maximum quantum yield of PS II) and σ_PSII (the functional absorption cross‐section for PSII) started within the first hour, much earlier than other parameters. Cellular pigments did not recover during the 24 h but the chlorophyll (chl) a/carotenoid ratios increased to levels measured in the controls. Cell division was independent of the recovery of chl a. In E. huxleyi, the recovery of F_v/F_m and σ_PSII started after an hour, synchronous with the increase in cellular organic N and chl a with pigments fully recovered within 14 h. P. tricornutum prioritized the recovery of its photosynthetic functions and cell divisions while E. huxleyi did not follow this pattern. We hypothesize that the different recovery strategies between the two species allow P. tricornutum to be more competitive when N pulses are introduced into N‐limited water while E. huxleyi is adapted to N scarce waters where such pulses are infrequent. These findings are consistent with successional patterns observed in coastal environments. This is one of only a few studies exploring recovery kinetics of cellular functions and photosynthesis after nitrogen stress in phytoplankton. Our results can be used to enhance ecological models linking phytoplankton traits to species diversity and community structure.     

Phylogenetics and comparative analysis of the mitochondrial genomes of three violet‐ringed octopuses

Similar morphological characters and little molecular data of Amphioctopus rex, A. neglectus and A. cf. ovulum resulted in their unknown phylogenetic statuses and equivocal relationships. In this study, the complete mitochondrial genomes of these three species collected in Chinese waters were sequenced and compared with each other to clarify the relationships among them. The lengths of the mitochondrial genomes varied from 15,646 bp to 15,814 bp, and the A + T content and GC skew for protein‐coding genes showed little variation. In contrast, both a dendrogram based on codon usage and the gene arrangements of the three octopuses showed that A. rex was more closely related to A. neglectus than to A. cf. ovulum. Five data sets and two methods (maximum likelihood and Bayesian inference) were utilized for the first time to explore the phylogenetic relationships among these three species in Octopodidae. The results indicated that a data set combining protein‐coding genes and RNA genes (PR) was optimal for analysing the relationships among 43 cephalopods. All of the phylogenetic trees divided the cephalopods into 10 taxa and supported the monophyly of Oegopsida, Myopsida, Sepiidae and Octopodidae. In this study, Idiosepiidae was classified as sister to Sepiolidae. Trees constructed using all data sets robustly supported the monophyly of the genus Amphioctopus. Notably, A. rex was more closely related to A. neglectus than to A. cf. ovulum, although these three species share the characteristic of violet rings on dark ocelli.     

Phytoplankton and bacterial assemblages in ballast water of U.S. military ships as a function of port of origin, voyage time, and ocean exchange practices

We characterized the physical/chemical conditions and the algal and bacterial assemblages in ballast water from 62 ballast tanks aboard 28 ships operated by the U.S. Military Sealift Command and the Maritime Administration, sampled at 9 ports on the U.S. West Coast and 4 ports on the U.S. East Coast. The ballast tank waters had been held for 2–176 days, and 90% of the tanks had undergone ballast exchange with open ocean waters. Phytoplankton abundance was highly variable (grand mean for all tanks, 3.21 × 10⁴ viable cells m⁻³; median, 7.9 × 10³ cells m⁻³) and was unrelated to physical/chemical parameters, except for a positive relationship between centric diatom abundance and nitrate concentration. A total of 100 phytoplankton species were identified from the ballast tanks, including 23 potentially harmful taxa (e.g. Chaetoceros concavicornis, Dinophysis acuminata, Gambierdiscus toxicus, Heterosigma akashiwo, Karlodinium veneficum, Prorocentrum minimum, Pseudo-nitzschia multiseries). Assemblages were dominated by chain-forming diatoms and dinoflagellates, and viable organisms comprised about half of the total cells. Species richness was higher in ballast tanks with coastal water, and in tanks containing Atlantic or Pacific Ocean source waters rather than Indian Ocean water. Total and viable phytoplankton numbers decreased with age of water in the tanks. Diversity also generally decreased with water age, and tanks with ballast water age >33 days did not produce culturable phytoplankton. Abundance was significantly higher in tanks with recently added coastal water than in tanks without coastal sources, but highly variable in waters held less than 30 days. Bacterial abundance was significantly lower in ballast tanks with Atlantic than Pacific Ocean source water, but otherwise was surprisingly consistent among ballast tanks (overall mean across all tanks, 3.13 ± 1.27 × 10¹¹ cells m⁻³; median, 2.79 × 10¹¹ cells m⁻³) and was unrelated to vessel type, exchange status, age of water, environmental conditions measured, or phytoplankton abundance. At least one of four pathogenic eubacteria (Listeria monocytogenes, Escherichia coli, Mycobacterium spp., Pseudomonas aeruginosa) was detected in 48% of the ballast tanks, but toxigenic strains of Vibrio cholerae were not detected. For ships with tanks of similar ballasting history, the largest source of variation in phytoplankton and bacteria abundance was among ships; for ships with tanks of differing ballasting histories, and for all ships/tanks considered collectively, the largest source of variation was within ships. Significant differences in phytoplankton abundance, but not bacterial abundance, sometimes occurred between paired tanks with similar ballasting history; hence, for regulatory purposes phytoplankton abundance cannot be estimated from single tanks only. Most tanks (94%) had adequate records to determine the source locations and age of the ballast water and, as mentioned, 90% had had ballast exchange with open-ocean waters. Although additional data are needed from sediments that can accumulate at the bottom of ballast tanks, the data from this water-column study indicate that in general, U.S. Department of Defense (DoD) ships are well managed to minimize the risk for introduction of harmful microbiota. Nevertheless, abundances of viable phytoplankton with maximum dimension >50 μm exceeded proposed International Maritime Organization standards in 47% of the ballast tanks sampled. The data suggest that further treatment technologies and/or alternative management strategies will be necessary to enable DoD vessels to comply with proposed standards.     

EMILIANIA HUXLEYI (PRYMNESIOPHYCEAE): NITROGEN‐METABOLISM GENES AND THEIR EXPRESSION IN RESPONSE TO EXTERNAL NITROGEN SOURCES1

The availability and composition of dissolved nitrogen in ocean waters are factors that influence species composition in natural phytoplankton communities. The same factors affect the ratio of organic to inorganic carbon incorporation in calcifying species, such as the coccolithophore Emiliania huxleyi (Lohman) W. W. Hay et H. Mohler. E. huxleyi has been shown to thrive on various nitrogen sources, including dissolved organic nitrogen. Nevertheless, assimilation of dissolved nitrogen under nitrogen‐replete and ‐limited conditions is not well understood in this ecologically important species. In this study, the complete amino acid sequences for three functional genes involved in nitrogen metabolism in E. huxleyi were identified: a putative formamidase, a glutamine synthetase (GSII family), and assimilatory nitrate reductase. Expression patterns of the three enzymes in cells grown on inorganic as well as organic nitrogen sources indicated reduced expression levels of nitrate reductase when cells were grown on NH₄⁺ and a reduced expression level of the putative formamidase when growth was on NO₃^?. The data reported here suggest the presence of a nitrogen preference hierarchy in E. huxleyi. In addition, the gene encoding for a phosphate repressible phosphate permease was more highly expressed in cells growing on formamide than in cells growing on inorganic nitrogen sources. This finding suggests a coupling between phosphate and nitrogen metabolism, which might give this species a competitive advantage in nutrient‐depleted environments. The potential of using expression of genes investigated here as indicators of specific nitrogen‐metabolism strategies of E. huxleyi in natural populations of phytoplankton is discussed.     

Transcriptome analysis of functional differentiation between haploid and diploid cells of Emiliania huxleyi, a globally significant photosynthetic calcifying cell

Background  

Eukaryotes are classified as either haplontic, diplontic, or haplo-diplontic, depending on which ploidy levels undergo mitotic cell division in the life cycle. Emiliania huxleyi is one of the most abundant phytoplankton species in the ocean, playing an important role in global carbon fluxes, and represents haptophytes, an enigmatic group of unicellular organisms that diverged early in eukaryotic evolution. This species is haplo-diplontic. Little is known about the haploid cells, but they have been hypothesized to allow persistence of the species between the yearly blooms of diploid cells. We sequenced over 38,000 expressed sequence tags from haploid and diploid E. huxleyi normalized cDNA libraries to identify genes involved in important processes specific to each life phase (2N calcification or 1N motility), and to better understand the haploid phase of this prominent haplo-diplontic organism.     

Effects of the diatom-Emiliana huxleyi succession on photosynthesis, calcification and carbon metabolism by size-fractioned phytoplankton

Changes in the species composition, photosynthesis, calcification and size-fractionated carbon metabolism by natural phytoplankton assemblages were monitored in three mesocosms under different nutrient conditions during May 1993. In the 3 enclosures, the decline of the diatom-dominated assemblages was followed by the development of a bloom of the coccolithoporid Emiliania huxleyi. Highest growth of E. huxleyi was observed in the mesocosm with a high N : P ratio, suggesting this species is a good competitor at low phosphate concentrations. The transition from diatom- to E. huxleyi-dominated assemblages brought about a sharp reduction of the phytoplankton standing stock and carbon-specific photosynthetic rate. The relative contribution of the smaller size fraction to total photosynthesis increased as the succession progressed. Calcification rate and E. huxleyi cell-specified calcite production were highest during the early stages of development of the E. huxleyi bloom. Distinct changes in the patterns of ¹⁴C allocation into biomolecules were noticed during the diatom-E. huxleyi succession. The diatom-dominated assemblage showed high relative ¹⁴C incorporation into low molecular weight metabolites (LMWM), whereas proteins and, specially, lipids accounted for the largest proportion of carbon incorporation in the E. huxleyi bloom. The patterns of photoassimilated carbon metabolism proved to be strongly dependent on cellular size, as protein relative synthesis was significantly higher in the smaller than in the larger size fraction, irrespective of the nutrient regime and the successional stage. These results are discussed in relation to the ecological and physiological features of small phytoplankton.     

Different photochemical responses of phytoplankters from the large shallow Taihu Lake of subtropical China in relation to light and mixing

The maximum quantum yield of photosystem II was estimated from variable chlorophyll a fluorescence in samples of phytoplankton collected from the Taihu Lake in China to determine the responses of different phytoplankters to irradiance and vertical mixing. Meteorological and environmental variables were also monitored synchronously. The maximum quantum yield of three phytoplankton groups: cyanobacteria, chlorophytes, and diatoms/dinoflagellates, showed a similar diurnal change pattern. F _v/F _m decreased with a significant depth-dependent variation as irradiance increased during the morning and increased as irradiance declined in the afternoon. Furthermore, the rates of F _v/F _m depression were dependent upon the photon flux density, whereas the rates of recovery of F _v/F _m were dependent upon the historical photon density. Moreover, photoinhibition affected the instantaneous growth rates of phytoplankton. Although at noon cyanobacteria had a higher photoinhibition value (up to 41%) than chlorophytes (32%) and diatoms/dinoflagellates (34%) at the surface, no significant difference in diurnal growth rates among the three phytoplankton groups were observed indicating that cyanobacteria could photoacclimate better than chlorophytes and diatoms/dinoflagellates. In addition, cyanobacteria had a higher nonphotochemical quenching value than chlorophytes and diatoms/dinoflagellates at the surface at noon, which indicated that cyanobacteria were better at dissipating excess energy. The ratios of enclosed bottle samples F _v/F _m to free lake samples F _v/F _m showed different responses for the three phytoplankton groups to irradiance and vertical mixing when wind speed was approximately constant at about 3.0 m s⁻¹. When wind speed was lower than 3.0 m s⁻¹, cyanobacteria accumulated mainly at the surface and 0.3 m, because of their positive buoyancy, where diurnal growth rates of phytoplankton were relatively higher than those at 0.6 m and 0.9 m. Chlorophytes were homogenized completely by vertical mixing, while diatoms/dinoflagellates avoided active high irradiance by moving downward at noon, and then upward again when irradiance decreased. These results explain the dominance of cyanobacteria in Taihu Lake. Handling editor: L. Naselli-Flores     

Genetic delineation between and within the widespread coccolithophore morpho‐species Emiliania huxleyi and Gephyrocapsa oceanica (Haptophyta)

Emiliania huxleyi and Gephyrocapsa oceanica are abundant coccolithophore morpho‐species that play key roles in ocean carbon cycling due to their importance as both primary producers and cal‐cifiers. Global change processes such as ocean acidification impact these key calcifying species. The physiology of E. huxleyi, a developing model species, has been widely studied, but its genetic delineation from G. oceanica remains unclear due to a lack of resolution in classical genetic markers. Using nuclear (18S rDNA and 28S rDNA), mitochondrial (cox1, cox2, cox3, rpl16, and dam), and plastidial (16S rDNA, rbcL, tufA, and petA) DNA markers from 99 E. huxleyi and 44 G. oceanica strains, we conducted a multigene/multistrain survey to compare the suitability of different markers for resolving phylogenetic patterns within and between these two morpho‐species. The nuclear genes tested did not provide sufficient resolution to discriminate between the two morpho‐species that diverged only 291Kya. Typical patterns of incomplete lineage sorting were generated in phylogenetic analyses using plastidial genes. In contrast, full morpho‐species delineation was achieved with mitochondrial markers and common intra‐morpho‐species phylogenetic patterns were observed despite differing rates of DNA substitution. Mitochondrial genes are thus promising barcodes for distinguishing these coccolithophore morpho‐species, in particular in the context of environmental monitoring.     

Seasonal dependence and functional implications of macrophyte–phytoplankton allelopathic interactions

Invasive plant species such as Ludwigia hexapetala might have a competitive advantage if they produce allelopathically active compounds against primary producers. Both phytoplankton and plant community structure may be affected due to different, species‐specific sensitivity to allelochemicals. Moreover, such allelopathic interactions could vary over the year depending on (i) the plant's phenological stage and (ii) the abilities of the native macrophytes to suppress—or the non‐native macrophytes to stimulate—the non‐native macrophyte population. We tested the allelopathic effects of aqueous leaf extracts of L. hexapetala on the photosynthetic activity of three target phytoplankton strains (Scenedesmus communis, a toxic Microcystis aeruginosa strain and a non‐toxic Microcystis aeruginosa strain) over three seasons of development (spring, summer and autumn). We also tested seasonal allelopathic effects of aqueous leaf extracts of both L. hexapetala (i.e. the non‐native invasive species) and the native Mentha aquatica on L. hexapetala seed germination. Finally, we identified three main secondary compounds present in the aqueous leaf extracts of L. hexapetala and we tested each individual compound on the phytoplankton's photosynthetic activity and on L. hexapetala seed germination. We observed marked seasonal and species‐specific patterns of L. hexapetala allelopathy on phytoplankton. The photosynthetic activities of S. communis and the toxic M. aeruginosa strain were stimulated by L. hexapetala aqueous leaf extracts in autumn and spring, respectively, whereas the non‐toxic M. aeruginosa strain was strongly inhibited in these two seasons. In summer, photosynthesis of all phytoplankton strains was inhibited. The germination rate of L. hexapetala seeds was stimulated by both L. hexapetala and M. aquatica aqueous leaf extracts, especially in summer, concomitant with the strong negative effects observed on the three phytoplankton strains. Three flavonoid glycosides (myricitrin, prunin and quercitrin) were identified as the main secondary compounds present in the L. hexapetala aqueous leaf extracts. The photosynthetic activity of S. communis was slightly stimulated by the three compounds. The photosynthetic activity of the toxic M. aeruginosa strain was stimulated by myricitrin and quercitrin, whereas that of the non‐toxic M. aeruginosa strain was inhibited by prunin. Finally, the germination rate and the germination velocity of L. hexapetala seeds were stimulated by myricitrin and prunin. These findings suggest that L. hexapetala could favour the photosynthetic activity of toxic cyanobacteria in spring and reduce their photosynthetic activity in summer, potentially leading to drastic changes in the phytoplankton communities and therewith ecological functioning of invaded ponds. Moreover, the stimulation of its seed germination could give a strong competitive advantage to L. hexapetala, thus promoting its invasiveness.     

Abiotic stress modifies the synthesis of alpha‐tocopherol and beta‐carotene in phytoplankton species

We performed laboratory experiments to investi‐gate whether the synthesis of the antioxidants α‐tocopherol (vitamin E) and β‐carotene in phytoplankton depends on changes in abiotic factors. Cultures of Nodularia spumigena, Phaeodactylum tricornutum, Skeletonema costatum, Dunaliella tertiolecta, Prorocentrum cordatum, and Rhodomonas salina were incubated at different tempe‐ratures, photon flux densities and salinities for 48 h. We found that abiotic stress, within natural ecological ranges, affects the synthesis of the two antioxidants in different ways in different species. In most cases antioxidant production was stimulated by increased abiotic stress. In P. tricornutum KAC 37 and D. tertiolecta SCCAP K‐0591, both good producers of this compound, α‐tocopherol accumulation was negatively affected by environmentally induced higher photosystem II efficiency (F_v/F_m). On the other hand, β‐carotene accumulation was positively affected by higher F_v/F_m in N. spumigena KAC 7, P. tricornutum KAC 37, D. tertiolecta SCCAP K‐0591 and R. salina SCCAP K‐0294. These different patterns in the synthesis of the two compounds may be explained by their different locations and functions in the cell. While α‐tocopherol is heavily involved in the protection of prevention of lipid peroxidation in membranes, β‐carotene performs immediate photo‐oxidative protection in the antennae complex of photosystem II. Overall, our results suggest a high variability in the antioxidant pool of natural aquatic ecosystems, which can be subject to short‐term temperature, photon flux density and salinity fluctuations. The antioxidant levels in natural phytoplankton communities depend on species composition, the physiological condition of the species, and their respective strategies to deal with reactive oxygen species. Since α‐tocopherol and β‐carotene, as well as many other nonenzymatic antioxidants, are exclusively produced by photo‐synthetic organisms, and are required by higher trophic levels through dietary intake, regime shifts in the phytoplankton as a result of large‐scale environmental changes, such as climate change, may have serious consequences for aquatic food webs.     

Phylogeographic characteristics of hypervariable regions in the mitochondrial genome of a cosmopolitan,bloom‐forming raphidophyte,Heterosigma akashiwo

The global distribution of phytoplankton is defined by many events, including long‐term evolutionary processes and shorter time span processes (e.g., global climate change). Furthermore, human‐assisted, unintentional dispersion, including the transport of live fish and spat for aquaculture, and transfer of aquatic microorganisms contained in ship ballast water, may aid the spread of phytoplankton. To understand the phylogeographic history of a species, the development of useful molecular markers is crucial. We previously reported a hypervariable mitochondrial gene in the cosmopolitan bloom‐forming alga, Heterosigma akashiwo. In this study, we identified two additional hypervariable segments in the H. akashiwo mitochondrial genome, one a protein coding sequence, and the other an intergenic region, by comparing the whole mitogenomes of strains obtained from various geographic origins. Interestingly, the newly identified hypervariable protein coding sequence was a paralogue of the previously identified gene, and both sequences showed tendencies to correlate with latitude of geographic origin. However, the hypervariable intergenic sequence did not show a clear correlation with origin. Our results demonstrated that the protein coding sequences may serve as useful tools for understanding the phylogeographic history of H. akashiwo, and they may crucially function in adaptation of the species to the environment.     

Eco‐energetic consequences of evolutionary shifts in body size

Size imposes physiological and ecological constraints upon all organisms. Theory abounds on how energy flux covaries with body size, yet causal links are often elusive. As a more direct way to assess the role of size, we used artificial selection to evolve the phytoplankton species Dunaliella tertiolecta towards smaller and larger body sizes. Within 100 generations (c. 1 year), we generated a fourfold difference in cell volume among selected lineages. Large‐selected populations produced four times the energy than small‐selected populations of equivalent total biovolume, but at the cost of much higher volume‐specific respiration. These differences in energy utilisation between large (more productive) and small (more energy‐efficient) individuals were used to successfully predict ecological performance (r and K) across novel resource regimes. We show that body size determines the performance of a species by mediating its net energy flux, with worrying implications for current trends in size reduction and for global carbon cycles.     

PHYTOPLANKTON SELENIUM REQUIREMENTS: THE CASE FOR SPECIES ISOLATED FROM TEMPERATE AND POLAR REGIONS OF THE SOUTHERN HEMISPHERE1

A series of laboratory culture experiments was used to investigate the effect of selenium (Se, 0–10 nM) on the growth, cellular volume, photophysiology, and pigments of two temperate and four polar oceanic phytoplankton species [coccolithophore Emiliania huxleyi (Lohmann) W. W. Hay et H. P. Mohler, cyanobacterium Synechococcus sp., prymnesiophyte Phaeocystis sp., and three diatoms—Fragilariopsis cylindrus (Grunow) Kriegar, Chaetoceros sp., and Thalassiosira antarctica G. Karst.]. Only Synechoccocus sp. and Phaeocystis sp. did not show any requirement for Se. Under Se‐deficient conditions, the growth rate of E. huxleyi was decreased by 1.6‐fold, whereas cellular volume was increased by 1.9‐fold. Se limitation also decreased chl a (2.5‐fold), maximum relative electron transport rate (1.9‐fold), and saturating light intensity (2.8‐fold), suggesting that Se plays a role in photosynthesis or high‐light acclimation. Pigment analysis for Antarctic taxa provided an interesting counterpoint to the physiology of E. huxleyi. For all Se‐dependent Antarctic diatoms, Se limitation decreased growth rate and chl a content, whereas cellular volume was not affected. Pigment analysis revealed that other pigments were affected under Se deficiency. Photoprotective pigments increased by 1.4‐fold, while diadinoxanthin:diatoxanthin ratios decreased by 1.5‐ to 4.9‐fold under Se limitation, supporting a role for Se in photoprotection. Our results demonstrate an Se growth requirement for polar diatoms and indicate that Se could play a role in the biogeochemical cycles of other nutrients, such as silicic acid in the Southern Ocean. Se measurements made during the austral summer in the Southern Ocean and Se biological requirement were used to discuss possible Se limitation in phytoplankton from contrasting oceanographic regions.     

Coccolith volume of the Southern Ocean coccolithophore Emiliania huxleyi as a possible indicator for palaeo‐cell volume

Coccolithophores are a key functional phytoplankton group and produce minute calcite plates (coccoliths) in the sunlit layer of the pelagic ocean. Coccoliths significantly contribute to the sediment record since the Triassic and their geometry have been subject to palaeoceanographic and biological studies to retrieve information on past environmental conditions. Here, we present a comprehensive analysis of coccolith, coccosphere and cell volume data of the Southern Ocean Emiliania huxleyi ecotype A, subject to gradients of temperature, irradiance, carbonate chemistry and macronutrient limitation. All tested environmental drivers significantly affect coccosphere, coccolith and cell volume with driver‐specific sensitivities. However, a highly significant correlation emerged between cell and coccolith volume with V_coccolith = 0.012 ± 0.001 * V_cell + 0.234 ± 0.066 (n = 23, r² = .85, p < .0001, σ_est = 0.127), indicating a primary control of coccolith volume by physiological modulated changes in cell volume. We discuss the possible application of fossil coccolith volume as an indicator for cell volume/size and growth rate and, additionally, illustrate that macronutrient limitation of phosphorus and nitrogen has the predominant influence on coccolith volume in respect to other environmental drivers. Our results provide a solid basis for the application of coccolith volume and geometry as a palaeo‐proxy and shed light on the underlying physiological reasons, offering a valuable tool to investigate the fossil record of the coccolithophore E. huxleyi.