NSFGEO-NERC: An unexpected requirement for silicon in coccolithophore calcification: physiological, ecological and evolutionary implications

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Biomineralization by marine phytoplankton has had a profound impact on our planet. The production of special cell wall material, calcite coccoliths by coccolithophores and silica frustules by diatoms, are major drivers in global biogeochemical cycles, but the underlying cellular processes remain poorly understood. It is widely considered that calcification in coccolithophores occurs through a very different process to silicification in diatoms, however some ecologically important coccolithophore lineages possess diatom-like silicon (Si) transport systems and have an absolute requirement for Si during coccolith formation. Importantly, the abundant bloom-forming coccolithophores such as Emiliania huxleyi exhibit no requirement for Si. There is a clear need to understand how these different physiological requirements for dissolved Si have driven the ecology and evolution of the coccolithophores. The project will yield a more complete understanding of the Si requirements of coccolithophores, its role in the calcification process, and the impacts of Si availability on the biogeography of these important bloom forming phytoplankton. The results are expected to strengthen our ability to predict the responses of coccolithophores to short and long-term environmental change, and therefore the consequences for the marine biogeochemical cycles in which they participate. In addition to the scientific outcomes, the project provides independent research opportunities to a diverse pool of undergraduate students, provide interdisciplinary training for graduate students, and facilitate the professional development of post-doctoral researchers. Public engagement in the research is facilitated through participant involvement in regional science festivals, public outreach events, production of educational resources, and targeted K-12 summer camp activities.Calcification in coccolithophores appears to represent a distinct process from silicification in diatoms, another major group of biomineralized phytoplankton. The apparent absence of a requirement for silicon (Si) in coccolithophores has been proposed to play a critical role in their ability to out-compete the otherwise dominant diatoms in areas of low dissolved Si availability. However, the investigators recently demonstrated that some globally important coccolithophores possess diatom-like Si transporters and exhibit an obligate requirement for Si in the calcification process. This discovery has important implications both for phytoplankton ecology and for the evolution of biomineralization. Using a range of physiological, molecular and computational approaches the project will 1) Establish Si requirements of ecologically important coccolithophore groups; 2) Determine the physiological role of Si in coccolithophores; 3) Determine the evolutionary events leading to the differing requirements for Si in calcification; 4) Examine the ecological distribution of Si-requiring coccolithophores, and 5) Determine the impact of the Si requirement on coccolithophore ecology. This project therefore integrates the molecular identification of genes (Si transporters), the physiological role of these transporters, and ecosystem scale models in order to examine how the requirement for Si influences ecosystem functioning and coccolithophore biogeography. The results of this work provides essential data that describes the cellular mechanisms of calcification and the range of physiological diversity between major coccolithophore lineages. The research also explores a previously unforeseen aspect of phytoplankton ecology; examining how the differing requirements for Si in calcifying coccolithophores may have shaped competitive interactions with other phytoplankton over both contemporary and evolutionary timescales. Overall, the research provides novel insights into physiology, ecology and evolution of coccolithophores, including information on how and why coccoliths are produced, which is currently poorly understood. This information is vital in order to understand how coccolithophores have been influenced by past changes in the Earth's climate, and their potential responses to future oceans.