Algal Calcification and Silification

The algae represent major producers of calcium carbonate and silica among the world's biota. Calcification involves the precipitation of  CaCO3 from  Ca2+ and math ions. Algal calcification may account for up to half of global oceanic  CaCO3 production. Silicification is less widespread among algal groups, which transform dissolved silicate to skeletal material. Diatoms play a key role in marine silica cycling. Diatomaceous deposits have long been exploited for building and filling materials, and the low-temperature, low-pressure biogenic formation of silica has potential for biotechnological application in novel industrial processes.

Keywords: calcite; silica; coccolithophores; diatoms; biogeochemistry

Figure 1. Alternative models for the mechanism of external calcifying band production in the giant-celled alga Chara. In (a), calcification is driven by the diffusion of  H+ into the cell in localized regions, producing localized alkalinization of the cell surface and precipitation of  CaCO3. In an adjacent region,  H+ is actively pumped out of the cell producing localized acidification of the cell surface. In (b), calcification is driven by the extrusion of  Ca2+ in exchange for  H+ in the alkaline zone.  CO2 diffusion from the cell's interior provides the carbon source for math formation and  CaCO3 precipitation. In both models,  H+ extrusion in the acidic zone facilitates the production of  CO2 from math which can be used by photosynthesis in the chloroplasts (green).
Figure 2. Model for fluxes of  Ca2 +, math and  H+ during calcification in coccolithophores. (a) Scanning electron micrograph of heterococcoliths on the surface of Coccolithus pelagicus cells (cell diameter = 20 m). (b)  Ca2 + and math uptake into a Golgi-derived compartment leads to the production of  CaCO3 and  H+ during calcite precipitation.  H+ production can be used to counter the alkalinizing effect of  CO2 production from math in the chloroplast (green) and removal of  CO2 by photosynthesis.
Figure 3. (a) Scanning electron micrograph showing complete silica frustules of the diatom Thallassiosira eccentrica (cell diameter = 50 m). (b) Working model of silica biogenesis in a diatom.  Na+-dependent silicate uptake is mediated by specialized silica transporters (SIT) at the plasma membrane. Once inside the cell, the soluble silica is delivered to the silica deposition vesicle (SDV), possibly involving isoforms of the SITs. Silica polycondensation occurs under acidic conditions within the SDV to form insoluble amorphous silica. Soluble silica may be stored in intracellular pools, the size of which is sensitive to silica demand and metabolism. Silifin proteins are thought to play a key role in regulating the formation of insoluble silica within the SDV. Frustulins are a separate group of proteins that together with polysaccharides and lipids are likely to play a role in stabilization of the mature silica wall structure. On maturity, the complete valve is released onto the cell surface by exocytosis.
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 Further Reading
    Bhattacharyya P and Volcani BE (1980) Sodium-dependent silicate transport in the apochlorotic marine diatom Nitzschia alba. Proceedings of the National Academy of Sciences of the USA 77: 6386–6390.
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    book Green JC and Leadbeater BSC (eds) The Haptophyte Algae, pp. 321–334. Oxford: Clarendon Press.
    Mann S (1997) Biomineralization: the formidable part of bioinorganic chemistry. Journal of the Chemical Society, Dalton Transactions 21: 3953–3961.
    Martin-Jézéquel V, Hildebrand M and Brzezinski MA (2000) Silicon metabolism in diatoms: implications for growth. Journal of Phycology 36: 821–840.
    Pickett-Heaps J, Schmid A-MM and Edgar LA (1990) The cell biology of diatom valve formation. Progress in Phycological Research 7: 1–168.
    book Round FE, Crawford RM and Mann DG (1990) The Diatoms: Biology and Morphology of the Genera. Cambridge: Cambridge University Press.
    Siegenthaler U and Sarmiento JL (1993) Atmospheric carbon dioxide and the ocean. Nature 365: 119–125.
    book Simkiss K and Wilbur KM (1989) Biomineralization: Cell Biology and Mineral Deposition. San Diego: Academic Press.
    book Stoermer EF and Smol JP (eds) (1999) The Diatoms: Applications for the Environmental and Earth Sciences. Cambridge: Cambridge University Press.
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Brownlee, Colin, and Taylor, Alison R(May 2002) Algal Calcification and Silification. In: eLS. John Wiley & Sons Ltd, Chichester. http://www.els.net [doi: 10.1038/npg.els.0000313]