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

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The oceans cover more than three quarters of the surface of the Earth and tiny algae in our seas are responsible for half of all photosynthesis on our planet. These single celled organisms, known as phytoplankton, form the basis of marine food webs and their activities can have an enormous impact on the geology of our planet. One group of phytoplankton known as the coccolithophores produce a covering of calcium carbonate plates (coccoliths) and can form vast blooms in the oceans. When the coccolithophores die, their coccoliths settle to the ocean floor, leading to the formation of sedimentary rocks, such as chalks and limestones.In many parts of the ocean the low availability of nutrients (such as nitrogen and phosphorus) limits phytoplankton growth. Competition for nutrients plays an important role in determining which phytoplankton species can grow in different environments. One of the most successful phytoplankton groups in modern oceans is the diatoms, which are fast-growing, making it difficult for many other phytoplankton types to compete with them. However, diatoms need lots of dissolved silicon to make their silica cell walls. In some marine environments, the availability of silicon limits the growth of diatoms, allowing other phytoplankton (which do not need silicon) to grow in their place.It is commonly thought that the calcifying coccolithophores have no requirement for silicon. However, we have recently discovered that some important coccolithophore species actually possess silicon transporters that are similar to those used by diatoms. Remarkably, we found that these coccolithophores use silicon to make their calcium carbonate coccoliths. Therefore the processes of silica formation in diatoms and calcite production in coccolithophores, which were previously believed to be distinct processes, show a completely unexpected link. These findings have important implications for the evolution of biomineralisation in phytoplankton and for the competitive interactions between coccolithophores and diatoms.Not all coccolithophores show a requirement for silicon. We found that the species responsible for the massive coccolithophore blooms, Emiliania huxleyi, does not possess silicon transporters and exhibits no need for silicon in the calcification process. The absence of a requirement for silicon may have enabled bloom-forming species to grow better in areas where silicon is low (e.g. after a diatom bloom). There is therefore a clear need to understand the role of silicon in coccolithophore biology.In this proposal we will address this issue using a combination of laboratory experiments and computational modelling approaches. Firstly, we will use molecular genetic and laboratory experiments to determine which of the major coccolithophore species exhibit a requirement for silicon. We will then select species for detailed physiological analysis, to determine how silicon contributes to the formation of coccoliths and how coccolithophores take up silicon from the surrounding seawater. These studies will allow us to examine the evolutionary history of the requirement for silicon and determine when certain lineages appear to have lost this trait. Using parameters on Si uptake and usage derived from our experimental work, we will use computer simulations to model global coccolithophore distributions and identify environments where the requirement for Si appears to be playing an important role in coccolithophore ecology.The research will provide novel insight into physiology, ecology and evolution of coccolithophores, including information on how and why coccoliths are produced, which is currently poorly understood. The research will also inform us on the evolution of coccolith formation, which will be vitally important if we are to understand how coccolithophores have been influenced by past changes in the Earth's climate and how they may respond to changes in the future.