Bacterial magnetosomes: microbiology, biomineralization
and biotechnological applications
D. Schiiler . R. B. Frankel
Max-Planck-Institut fUr Marine Mikrobiologie, 28 359 Bremen, Germany
Department of Physics, California Polytechnic State University, San Luis Obispo, CA 93407, USA
Abstract Magnetotactic bacteria orient and migrate
along geomagnetic field lines. This ability is based on
intracellular magnetic structures, the magnetosomes,
which comprise nanometer-sized, membrane-bound
crystals of the magnetic iron minerals magnetite (Fe304)
or greigite (Fe3S4)' Magnetosome formation is achieved
by a mineralization process with biological control over
the accumulation of iron and the deposition of the
mineral particle with specific size and orientation within
a membrane vesicle at specific locations in the cell. This
review focuses on the current knowledge about magne
totactic bacteria and will outline aspects of the physi
ology and molecular biology of the biomineralization
process. Potential biotechnological applications of
magnetotactic bacteria and their magnetosomes as well
as perspectives for further research are discussed.
Introduction
The formation of biological hard materials, i.e., bone,
shell and spicule, involves the deposition of specific in
organic minerals on or in an organic matrix in a highly
controlled manner (Lowenstam 1981). Much current
research is directed towards understanding these bio
mineralization processes, which are collectively known
as biologically controlled mineralization, because of
their relevance to the synthesis of advanced materials
with tailored properties (Mann 1993; Moskowitz 1995).
One of the most intriguing examples of biologically
controlled mineralization is the formation of magnetic
nano-crystals in magnetosomes within magnetotactic
bacteria. The formation of magnetosomes in these or
ganisms is a well-documented example of the appar
ently widespread occurrence of magnetic minerals in
the living world. Biomineralization of ferromagnetic
materials, mainly magnetite, has been reported for an
extremely diverse range of organisms including algae
(Torres de Araujo et al. 1986), insects (Maher 1998),
molluscs (Lowenstam 1981), fish (Mann et al. 1988),
birds (Wiltschko and Wiltschko 1995) and even hu
mans (Kirschvink et al. 1992). The discovery of mag
netotactic bacteria by R. Blakemore (1975) stimulated
interdisciplinary research interest among scientists, in
cluding microbiologists, physicists, geologists, chemists,
and engineers. Commercial uses of bacterial magneto
some particles have been suggested, including the
manufacture of magnetic tapes and printing inks,
magnetic targeting of pharmaceuticals, cell separation
and their application as contrast-enhancement agents in
magnetic resonance imaging (Blakemore 1983; Mann
et al. 1990; Matsunaga 1991; Schwartz and Blakemore
1984).
Ecology and phylogeny of magnetotactic bacteria
Magnetotactic bacteria represent a heterogeneous group
of procaryotes with a variety of morphological types
including cocci, rods, vibrios, spirilla and apparently
multicellular forms (Bazylinski et al. 1994; Spring and
Schleifer 1995). They are constituents of natural micro
bial communities in sediments and chemically stratified
water columns and a broad diversity of morphological
forms has been found in many marine and freshwater
habitats. The highest numbers of magneto tactic bacteria
are generally found at the oxic/anoxic transition zone or
redoxocline, which is located in many freshwater habi
tats at the sediment/water interface or just below
(Bazylinski 1995; Stolz 1993). Accordingly, nearly all of
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