Intracellular magnetite crystal formation by magnetotactic bacteria has emerged as a powerful model for investigating the cellular and molecular mechanisms of biomineralization, a process common to all branches of life. Although magnetotactic bacteria are phylogenetically diverse and their crystals morphologically diverse, studies to date have focused on a few, closely related species with similar crystal habits. Here, we investigate the process of magnetite biomineralization in Desulfovibrio magneticus sp. RS-1, the only reported species of cultured magnetotactic bacteria that is outside of the α-Proteobacteria and that forms bulletshaped crystals. Using a variety of high-resolution imaging and analytical tools, we show that RS-1 cells form amorphous, noncrystalline granules containing iron and phosphorus before forming magnetite crystals. Using NanoSIMS (dynamic secondary ion mass spectroscopy), we show that the iron-phosphorus granules and the magnetite crystals are likely formed through separate cellular processes. Analysis of the cellular ultrastructure of RS-1 using cryo-ultramicrotomy, cryo-electron tomography, and tomography of ultrathin sections reveals that the magnetite crystals are not surrounded by membranes but that the iron-phosphorus granules are surrounded by membranous compartments. The varied cellular paths for the formation of these two minerals lead us to suggest that the iron-phosphorus granules constitute a distinct bacterial organelle.bacterial organelle | biomineralization | magnetotactic bacteria | dynamic secondary ion mass spectroscopy | magnetite B iomineralization, the biologically controlled transformation of inorganic compounds into highly-ordered structures, is performed by organisms in all branches of life, from bacteria to humans. Because biogenic minerals often have superior properties to their chemically synthesized counterparts, understanding biomineralization is important to fields as diverse as inorganic materials synthesis and medicine (1, 2). Among the numerous organisms capable of biomineralization, magnetotactic bacteria (MB), a group of microbes that form intracellular chains of magnetic minerals including magnetite and greigite, have become powerful models for studying biomineralization because of their relatively fast growth rate and genetic tractability (3).Investigation of the cellular ultrastructure of MB has shown that the magnetite crystals are formed within membranous intracellular compartments called magnetosomes, which are present before the formation of magnetite (4, 5). Other studies have shown that when magnetite formation is induced by decreasing oxygen levels or adding iron to iron-deprived cells, multiple crystals in the chain nucleate simultaneously (5, 6). These insights into the kinetics of magnetite biomineralization have been complemented by molecular studies. A genomic island called the magnetosome island (MAI) has been found in all MB studied to date. Loss of the MAI results in an absence of magnetosome membranes and magnetite crystals, and gene...