The formation of marine phosphatic mineral deposits remains incompletely understood, despite decades of research. The involvement of bacteria in this process has long been suspected, and both modern and ancient associations between bacteria and phosphorites have been recorded. Only recently has a specific bacterial metabolic process associated with the formation of phosphorites been discovered. Recent studies demonstrate that polyphosphate utilization by sulfide-oxidizing bacteria results in the rapid precipitation of apatite – providing at least a partial mechanism to explain the close spatial correlation between accumulations of sulfide-oxidizing bacteria and modern phosphorites. Possible fossilized bacteria are known from ancient phosphatic mineral deposits. Potentially, the fossilized cells represent the remains of bacteria that induced the formation of those phosphorites. However, robust criteria for the recognition of these bacteria have yet to be identified.

Marine phosphate-rich sedimentary deposits (phosphorites) are important geological reservoirs for the biologically essential nutrient phosphorous. Phosphorites first appear in abundance approximately 600 million years ago, but their proliferation at that time is poorly understood. Recent marine phosphorites spatially correlate with the habitats of vacuolated sulfide-oxidizing bacteria that store polyphosphates under oxic conditions to be utilized under sulfidic conditions. Hydrolysis of the stored polyphosphate results in the rapid precipitation of the phosphate-rich mineral apatite-providing a mechanism to explain the association between modern phosphorites and these bacteria. Whether sulfur bacteria were important to the formation of ancient phosphorites has been unresolved. Here, we present the remains of modern sulfide-oxidizing bacteria that are partially encrusted in apatite, providing evidence that bacterially mediated phosphogenesis can rapidly permineralize sulfide-oxidizing bacteria and perhaps other types of organic remains. We also describe filamentous microfossils that resemble modern sulfide-oxidizing bacteria from two major phosphogenic episodes in the geologic record. These microfossils contain sulfur-rich inclusions that may represent relict sulfur globules, a diagnostic feature of modern sulfide-oxidizing bacteria. These findings suggest that sulfur bacteria, which are known to mediate the precipitation of apatite in modern sediments, were also present in certain phosphogenic settings for at least the last 600 million years. If polyphosphate-utilizing sulfide-oxidizing bacteria also played a role in the formation of ancient phosphorites, their requirements for oxygen, or oxygen-requiring metabolites such as nitrate, might explain the temporal correlation between the first appearance of globally distributed marine phosphorites and increasing oxygenation of Neoproterozoic oceans.

Sedimentary phosphorites comprise a major phosphorus (P) ore, yet their formation remains poorly understood. Extant polyphosphate-metabolizing bacterial communities are known to act as bacterial phosphate-pumps, leading to episodically high dissolved phosphate concentrations in pore waters of organic-rich sediment. These conditions can promote the precipitation of amorphous precursor phases that are quickly converted to apatite-usually in carbonate fluorapatite form [Ca (PO ,CO ) F ]. To assess the mechanisms underpinning the nucleation and growth of sedimentary apatite, we sampled P-rich sediments from the Namibian shelf, a modern environment where phosphogenesis presently occurs. The P-rich fraction of the topmost centimetres of sediment mainly consists of pellets about 50-400 μm in size, which in turn are comprised of micron-sized apatite particles that are often arranged into radial structures with diameters ranging from 2 to 4 μm, and morphologies that range from rod-shapes to dumbbells to spheres that resemble laboratory-grown fluorapatite-gelatin nanocomposites known from double-diffusion experiments in organic matrices. The nucleation and growth of authigenic apatite on the Namibian shelf is likely analogous to these laboratory-produced precipitates, where organic macromolecules play a central role in apatite nucleation and growth. The high density of apatite nucleation sites within the pellets (>10 particles per cm ) suggests precipitation at high pore water phosphate concentrations that have been reported from the Namibian shelf and may be attributed to microbial phosphate pumping. The intimate association of organic material with the apatite could suggest a possible role of biological substrata, such as exopolymeric substances (EPS), in the nucleation of apatite precursors. Importantly, we do not observe any evidence that the apatite particles are actual phosphatized microbes, contradicting some earlier studies. Nevertheless, these results further evidence the potential importance of microbially derived (extracellular) organic matter as a template for phosphatic mineral nucleation in both recent and ancient phosphorites.

The oxygenation of Earth's atmosphere allowed for the diversification of metabolisms to include those that rely on oxygen and its derivatives. For example, chemolithotrophic oxidation of sulfide and iron both require oxygen or nitrate as terminal electron acceptors. A growing number of oxygen-utilizing chemolithotrophs are known to accumulate intracellular polyphosphate as an energy reserve that allows them to adapt to the fluctuating redox conditions in their distinctive-gradient habitats. Polyphosphate is also thought to play an important role in the formation of phosphatic mineral deposits. Here we present fossil evidence of iron-oxidizing bacteria preserved as filamentous iron oxides within phosphatic Paleoproterozoic stromatolites. The filaments include twisted stalks similar to those produced by modern iron-oxidizing bacteria that are known to metabolize polyphosphate and inhabit steep redox gradients. Fossil iron-oxidizing bacteria preserved within some of the oldest known phosphorites serve as indicators of O 2 -Fe(II) gradients that may have supported microbially mediated phosphogenesis via polyphosphate metabolism and/or an active iron redox pump.

Certain phosphatic grains preserved in the rock record are interpreted as microfossils representing a diversity of microorganisms from bacteria to fossil embryos. In addition to bona fide primary biological features, phosphatic microfossils and fossil embryos commonly exhibit features that result from abiotic precipitation or diagenetic alteration. Distinguishing between abiotic and primary biological features can be difficult, and some features thought to represent biological tissue could instead be artifacts that are unrelated to the original morphology of a preserved organism. Here, we present experimentally generated, abiotically produced mineral precipitates that morphologically resemble biologically produced features, some of which may be observed in the rock record or noted in extant organisms, including embryos. These findings extend the diversity of biomorphic features known to result from abiotic precipitation.