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Previous studies of the subsurface biosphere have deduced average cellular doubling times of hundreds to thousands of years based upon geochemical models. We have directly constrained the in situ average cellular protein turnover or doubling times for metabolically active micro-organisms based on cellular amino acid abundances, D/L values of cellular aspartic acid, and the in vivo aspartic acid racemization rate. Application of this method to planktonic microbial communities collected from deep fractures in South Africa yielded maximum cellular amino acid turnover times of ~89 years for 1 km depth and 27 °C and 1-2 years for 3 km depth and 54 °C. The latter turnover times are much shorter than previously estimated cellular turnover times based upon geochemical arguments. The aspartic acid racemization rate at higher temperatures yields cellular protein doubling times that are consistent with the survival times of hyperthermophilic strains and predicts that at temperatures of 85 °C, cells must replace proteins every couple of days to maintain enzymatic activity. Such a high maintenance requirement may be the principal limit on the abundance of living micro-organisms in the deep, hot subsurface biosphere, as well as a potential limit on their activity. The measurement of the D/L of aspartic acid in biological samples is a potentially powerful tool for deep, fractured continental and oceanic crustal settings where geochemical models of carbon turnover times are poorly constrained. Experimental observations on the racemization rates of aspartic acid in living thermophiles and hyperthermophiles could test this hypothesis. The development of corrections for cell wall peptides and spores will be required, however, to improve the accuracy of these estimates for environmental samples., (© 2013 John Wiley & Sons Ltd.)

Although the notion of an early origin and diversification of life on Earth during the Archaean eon has received increasing support in geochemical, sedimentological and palaeontological evidence, ambiguities and controversies persist regarding the biogenicity and syngeneity of the record older than Late Archaean. Non-biological processes are known to produce morphologies similar to some microfossils, and hydrothermal fluids have the potential to produce abiotic organic compounds with depleted carbon isotope values, making it difficult to establish unambiguous traces of life. Here we report the discovery of a population of large (up to about 300 mum in diameter) carbonaceous spheroidal microstructures in Mesoarchaean shales and siltstones of the Moodies Group, South Africa, the Earth's oldest siliciclastic alluvial to tidal-estuarine deposits. These microstructures are interpreted as organic-walled microfossils on the basis of petrographic and geochemical evidence for their endogenicity and syngeneity, their carbonaceous composition, cellular morphology and ultrastructure, occurrence in populations, taphonomic features of soft wall deformation, and the geological context plausible for life, as well as a lack of abiotic explanation falsifying a biological origin. These are the oldest and largest Archaean organic-walled spheroidal microfossils reported so far. Our observations suggest that relatively large microorganisms cohabited with earlier reported benthic microbial mats in the photic zone of marginal marine siliciclastic environments 3.2 billion years ago.

Three morphotypes of microbial mats are preserved in rocks deposited in shallow-water facies of the 3.42 Ga Buck Reef chert (BRC). Morphotype alpha consists of fine anastomosing and bifurcating carbonaceous laminations, which loosely drape underlying detrital grains or form silica-filled lenses. Morphotype beta consists of meshes of fine carbonaceous strands intergrown with detrital grains and dark laminations, which loosely drape coarse detrital grains. Morphotype gamma consists of fine, even carbonaceous laminations that tightly drape underlying detrital grains. Preservation of nearly uncompacted mat morphologies and detrital grains deposited during mat growth within a well-characterized sedimentary unit makes quantitative correlation between morphology and paleoenvironment possible. All mats are preserved in the shallowest-water interval of those rocks deposited below normal wave base and above storm wave base. This interval is bounded below by a transgressive lag formed during regional flooding and above by a small condensed section that marks a local relative sea-level maximum. Restriction of all mat morphotypes to the shallowest interval of the storm-active layer in the BRC ocean reinforces previous interpretations that these mats were constructed primarily by photosynthetic organisms. Morphotypes alpha and beta dominate the lower half of this interval and grew during deposition of relatively coarse detrital carbonaceous grains, while morphotype gamma dominates the upper half and grew during deposition of fine detrital carbonaceous grains. The observed mat distribution suggests that either light intensity or, more likely, small variations in ambient current energy acted as a first-order control on mat morphotype distribution. These results demonstrate significant environmental control on biological morphogenetic processes independent of influences from siliciclastic sedimentation.