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Preprint volume - SIBM

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Pre-print Volume – Introductory lectures<br />

Topic 2: MARINE ORGANISMS AND ECOSYSTEMS AS MODEL SYSTEMS<br />

C. BROWNLEE, G.L. WHEELER, A. CHRACHRI, A.R. TAYLOR,<br />

A. HIGHFIELD, F.J. VERRET, D. SCHROEDER.<br />

Marine Biological Association of the UK, The Laboratory, Citadel Hill, Plymouth PL1 2PB, UK.<br />

cbr@mba.ac.uk<br />

COCCOLITHOPHORE BIOMINERALIZATION:<br />

FROM MOLECULES TO GLOBAL PROCESSES<br />

BIOMINERALIZZAZIONE DEI COCCOLITOFORIDI:<br />

DALLE MOLECOLE AI PROCESSI GLOBALI<br />

Abstract – Coccolithophores are responsible for a major component of biogenic calcite formation in the<br />

oceans. Despite their biogeochemical importance, the molecular and cellular mechanisms of calcification<br />

are poorly understood. However, a deep understanding of the transport and biomineralization processes<br />

underlying coccolithophore biology is essential for understanding their ecological success and for<br />

predicting how they may be affected by or respond or adapt to future ocean acidification scenarios.<br />

Key-words: coccolithophores, calcification, genomics, cell biology.<br />

Introduction - Coccolithophores occur in all of the world’s oceans and are represented<br />

by many different unicellular species. They are characterized by the production of<br />

intricate calcium carbonate (calcite) scales (coccoliths) during at least one phase of<br />

their life cycle. Some species form massive seasonal blooms in temperate oceanic<br />

waters and the reflectance of the calcite in these blooms renders them visible from<br />

space. Estimates suggest that coccolithophores account for approximately half of<br />

global biogenic calcium carbonate production. This group of organisms thus plays an<br />

important role in the biogeochemical cycling of carbon in the oceans. A significant<br />

proportion of the carbon fixed into calcite sinks to the ocean floor where it may form<br />

sediments that give rise to chalk and limestone deposits. This represents a significant<br />

long-term sequestration of inorganic carbon.<br />

Coccolithophores are known to produce two types of calcite scales: Holococcoliths that<br />

have a simple crystal structure, and heterococcliths that are formed by interlocking<br />

calcite crystals made up of calcite crystal elements of complex shape. Significantly,<br />

heterococcolith production has been demonstrated to occur in an intracellular<br />

compartment - the coccolith vesicle (CV). This vesicle is derived from the Golgi body<br />

and encloses the forming coccolith in a membrane-bound isolated compartment,<br />

allowing the chemical composition to be regulated to promote the ordered deposition<br />

of the calcite crystals (Brownlee and Taylor, 2005). A wide rang of experimental<br />

studies have shown that external bicarbonate is the inorganic substrate for calcification.<br />

The intracellular precipitation of carbonate results in the production of protons. It<br />

follows that calcification requires transport of the substrates for calcification<br />

(bicarbonate and calcium) into the cell and removal of the ionic products (protons)<br />

from the cell. Our earlier work has shown that the magnitude of the fluxes involved in<br />

calcification is extremely large, since fixation of inorganic carbon by calcification can<br />

often occur at similar rates to the photosynthetic fixation of carbon.<br />

Some key questions relating to transport processes underlying calcification relate to the<br />

identification of membrane transporters. Do coccolithophore cells have calcificationspecific<br />

transport systems or does calcification recruit the cell’s normal transport<br />

41 st S.I.B.M. CONGRESS Rapallo (GE), 7-11 June 2010<br />

90

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