ALBERT

Notes: Lilypad stromatolites, up to 3 m long and 1·5 m wide, were found to be actively growing in the shallow marginal waters of Frying Pan Lake and its outflow channel. These stromatolites, composed of Phormidium (〉 90%), Fischerella, and a variety of other microbes, develop through a series of distinct growth stages. Dark green microbial mats cover the floor of the outflow channel and give rise to columns of various sizes and shapes in the shallower marginal waters. Once the columns reach the water level, the mats spread laterally to form a lilypad stromatolite. The lilypads are characterized by a raised, dark green rim, 4–5 mm high, that encircles a flat interior covered with a distinctive orange-red mat. The microbes forming the columns and lilypad plate are being actively silicified. The stromatolites are formed of: (i) flat-lying Phormidium filaments (P-laminae), (ii) upright filaments of Phormidium that are commonly associated with Fischerella (U-laminae), and (iii) mucus, diatoms and pyrite framboids (M-laminae). P-laminae dominate most of the columns, with tripartite cycles of P-, U-, to M-laminae being found mostly in the upper parts of the stromatolites. The transition from the P- to U-laminae is marked by a change in the growth pattern of the Phormidium and branching of Fischerella, which was probably triggered by a change in environmental conditions. In the Frying Pan Lake outflow channel, this change may be related to fluctuations in water level and flow rates that are caused by periods of heavy rain, seasonal changes, long-term variations in rainfall, and/or the unique 40-day hydrological cycle that exists between Frying Pan Lake and Inferno Crater, which is a nearby hydrothermal crater lake.

Notes: Silicified deposits, such as sinters, occur in several modern geothermal environments, but the mechanisms of silicification (and crucially the role of microorganisms in their construction) are still largely unresolved. Detailed examination of siliceous sinter, in particular sections of microstromatolites growing at the Krisuvik hot spring, Iceland, reveals that biomineralization contributes a major component to the overall structure, with approximately half the sinter thickness attributed to silicified microorganisms. Almost all microorganisms observed under the scanning electron microscope (SEM) are mineralized, with epicellular silica ranging in thickness from 〈 5 μm coatings on individual cells, to regions where entire colonies are cemented together in an amorphous silica matrix tens of micrometres thick. Within the overall profile, there appears to be two very distinct types of laminae that alternate repeatedly throughout the microstromatolite: ‘microbial’ layers are predominantly consisting of filamentous, intact, vertically aligned, biomineralized cyanobacteria, identified as Calothrix and Fischerella sp.; and weakly laminated silica layers which appear to be devoid of any microbial component. The microbial layers commonly have a sharply defined base, overlying the weakly laminated silica, and a gradational upper surface merging into the weakly laminated silica. These cyclic laminations are probably explained by variations in microbial activity. Active growth during spring/summer allows the microorganisms to keep pace with silicification, with the cell surfaces facilitating silicification, while during their natural slow growth phase in the dark autumn/winter months silicification exceeds the bacteria’s ability to compensate (i.e. grow upwards). At this stage, the microbial colony is probably not essential to microstromatolite formation, with silicification presumably occurring abiogenically. When conditions once again become favourable for growth, recolonization of the solid silica surface by free-living bacteria occurs: cell motility is not responsible for the laminations. We have also observed that microbial populations within the microstromatolite, some several mm in depth, appear viable, i.e. they still have their pigmentation, the trichomes are not collapsed, cell walls are unbroken, cytoplasm is still present and they proved culturable. This suggests that the bulk of silicification occurred rapidly, probably while the cells were still alive. Surprisingly, however, measurements of light transmittance through sections of the microstromatolite revealed that photosynthetically active light (PAL) only transmitted through the uppermost 2 mm. Therefore the ‘deeper’ microbial populations must have either: (i) altered their metabolic pathways; (ii) become metabolically inactive; or (iii) the deeper populations may be dominated by different microbial assemblages from that of the surface. From these collective observations, it now seems unequivocal that microstromatolite formation is intimately linked to microbial activity and that the sinter fabric results from a combination of biomineralization, cell growth and recolonization. Furthermore, the similarities in morphology and microbial component to some Precambrian stromatolites, preserved in primary chert, suggests that we may be witnessing contemporaneous biomineralization processes and growth patterns analogous to those of the early Earth.

Notes: Transmission electron microscopy examination of bacterial cells, growing naturally in freshwater and marine environments, reveals that they can precipitate a variety of iron minerals. The development of these authigenic mineral phases may be either ‘biologically controlled’, whereby the cell regulates mineral formation, or ‘biologically induced’, with biominerals commonly generated as secondary by-products of microbe-environment interactions. With the vast majority of bacteria biomineralisation is a two-step process; initially metals are electrostatically bound to the anionic surfaces of the cell wall and surrounding organic polymers, where they subsequently serve as nucleation sites for crystal growth. Because of its relatively high activity in aqueous solutions, iron is preferentially bound to reactive organic sites. As the latter stages of mineralisation are inorganically driven, the type of iron mineral formed is inevitably dependent on the available counter-ions, and hence, the chemical composition of the waters in which the microorganisms are growing.

Notes: The spontaneous precipitation of amorphous iron hydroxide and ferric hydroxysulfate has generally been considered to be an inorganic process involving the oxidation of ferrous iron with or without the presence of sulfate. However, our study of bacterial communities growing in an acid mine drainage lagoon sediment has confirmed that microorganisms were also capable of facilitating this mineral precipitation. Transmission electron microscopy revealed that bacteria growing at the surface had iron-rich capsules, along with detectable amounts of Zn, Ti, Mn and K incorporated into the mineralised matrix. In the subsurface, more cells were associated with granular, fine-grained mineral precipitates, composed almost exclusively of iron and sulfur. Pore water profiles indicated that no discernible sulfate reduction had taken place, suggesting that these authigenic minerals were ‘ferric hydroxysulfate’, and not iron sulfide. Energy dispersive X-ray spectroscopy further indicated that the subsurface minerals had variable composition, with the Fe:S ratio decreasing with depth from 3.5:1 at 15 cm to 1.9:1 at 30 cm. This indicates the high reactivity of ferric hydroxide for dissolved sulfate. Because iron reduction was limited to sediment depths between 3–10 cm, it is conceivable that these minerals are not amenable to bacterial reduction, and hence, the ability of bacteria to bind and form such precipitates may provide a natural solution to cleansing acidified waters with a high dissolved metal content.