The genus Geobacillus comprises endospore-forming obligate thermophiles. These bacteria have been isolated from cool soils and even cold ocean sediments in anomalously high numbers, given that the ambient temperatures are significantly below their minimum requirement for growth. Geobacilli are active in environments such as hot plant composts, however, and examination of their genome sequences reveals that they are endowed with a battery of sensors, transporters and enzymes dedicated to hydrolysing plant polysaccharides. Although they appear to be relatively minor members of the plant biomass-degrading microbial community, Geobacillus bacteria have achieved a significant population with a worldwide distribution, probably in large part due to adaptive features of their spores. First, their morphology and resistance properties enable them to be mobilized in the atmosphere and transported long distances. Second, their longevity, which in theory may be extreme, enables them to lie quiescent but viable for long periods of time, accumulating gradually over time to achieve surprisingly high population densities.
Introduction: an environmental enigmaGeobacillus bacteria are rod-shaped, aerobic or facultatively anaerobic, endospore-forming microbes. Depending on the strain, the temperature range for growth can extend as low as 35 u C or as high as 80 u C. But for most isolates, temperatures between about 45 and 70 u C are required (Nazina et al., 2001). These characteristics make Geobacillus bacteria (geobacilli) attractive to the biotechnology industry as sources of thermostable enzymes (de Champdoré et al., 2007; Karagüler et al., 2007), as platforms for biofuel production (Cripps et al., 2009;Taylor et al., 2009) and as potential components of bioremediation strategies (Markossian et al., 2000;Obojska et al., 2002;Perfumo et al., 2007). Obligate thermophiles, however, face a significant challenge outside the lab: they must survive on a planet with an average annual land surface temperature that has only ranged between 7 and 10 uC over the past three centuries (Rohde et al., 2013). It would be logical to assume, then, that Geobacillus would only be found in substantial numbers in the warmest regions of the planet, such as equatorial deserts or naturally occurring geothermal and hydrothermal hotspots. The truth, however, is rather more complicated.As Fig. 1 illustrates, people have been able to detect Geobacillus nearly everywhere they have thought to look. (This figure is based on a survey of 146 mostly recent references describing the isolation of Geobacillus in natural or human-made environments; for full details, see Table S1, available in Microbiology Online.) One can see at a glance that these bacteria have been isolated from sources on all seven continents as well as the Pacific Ocean and the Mediterranean Sea. What may not be obvious from a twodimensional map is that there has been much diversity in the third dimension as well. Geobacillus has been isolated from the Bolivian Andes at an altitude of 3653 m (Ma...