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Biomineralization: A Structural Perspective
Biomineralization: A Structural Perspective
Biomineralization: A Structural Perspective
com
Journal of
Structural
Biology
Journal of Structural Biology 163 (2008) 229234
www.elsevier.com/locate/yjsbi
Review
Abstract
Biomineralization is an inherently structural subject; the structure of the mineral phase, the structure of the matrix composed of macromolecules and especially the structure of the interphase zone between them. Studies of the dynamics of mineral formation have
revealed that a widespread strategy used by many organisms is to rst form a disordered mineral phase. Only when it is in place and
has adopted its appropriate shape, is it induced to crystallize. Matrix studies have highlighted the importance of a unique group of proteins that are rich in aspartic acid. These are involved in controlling mineral formation. Relating structure to function in mineralized
tissues, often involves an understanding of mechanical properties in terms of not only the hierarchical structure of the tissue, but also
the graded structure that varies from one location to another. Structure is thus in many respects the foundation upon which the eld
of biomineralization rests.
2008 Elsevier Inc. All rights reserved.
Keywords: Biomineralization; Biomaterials; Mineral formation
1047-8477/$ - see front matter 2008 Elsevier Inc. All rights reserved.
doi:10.1016/j.jsb.2008.02.001
230
Fig. 1. Scanning electron micrographs of 4 dierent mineralized tissues. (a) Statoconia from the bullfrog, composed presumably of aragonite (Marmo
et al., 1983) (Micrograph by H.A. Lowenstam). Scale bar: 1 lm. (b). Aragontic nacre in the process of formation on the inner surface of the bivalve Atrina
rigida. (Micrograph by F. Nudelman). Scale bar: 5 lm. (c). Antler-shaped spicules from the ascidian Pyura sacciformis composed of carbonated apatite
(Lowenstam, 1989) (Micrograph by H.A. Lowenstam). Scale bar: 100 lm. (d). Fracture surface of the working stone part of the sea urchin (Paracentrotus
lividus) tooth. The elongated needles of Mgcalcite are embedded in a matrix of nanoscale crystals composed of calcite with very high concentrations of
Mg (Schroeder et al., 1969; Wang et al., 1997) (Micrograph by Y. Ma). Scale bar: 2 lm.
eralized teeth of the chiton, a segmented mollusk, the initially formed mineral phase is not the same as the mature
form, but transforms into the more stable mature phase.
In order to identify this easily overlooked transient phase,
Towe and Lowenstam (1967) took advantage of the fact
that chitons form their teeth using a conveyor belt-like system, where every row of teeth represents a dierent stage of
formation. They could thus isolate the teeth with initial
mineral deposits and then compare this mineral, with the
mature crystalline phase. They found that the rst mineral
deposited was a disordered hydrated iron oxide, ferrihydrite (Fig. 2), and that within a few days after formation
it transformed into the crystalline iron oxide, magnetite.
In fact the mature teeth are magnetic. Chiton magnetite
was the rst example of biogenic magnetite to be discovered (Lowenstam, 1962).
It took 30 more years for the signicance of this paper to
be appreciated. This occurred when it was demonstrated
that members of another phylum, the echinoderms, also
use this transient precursor phase strategy. Beniash et al.
(1997) showed that sea urchin larvae form their calcitic
spicules via a transient disordered precursor phase called
amorphous calcium carbonate (ACC). The last decade
has witnessed an almost exponential increase in the number
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