Each recent report of liquid water existing elsewhere in the Solar System has reverberated through the international press and excited the imagination of humankind. Why? Because in the past few decades we have come to realize that where there is liquid water on Earth, virtually no matter what the physical conditions, there is life. What we previously thought of as insurmountable physical and chemical barriers to life, we now see as yet another niche harbouring 'extremophiles'. This realization, coupled with new data on the survival of microbes in the space environment and modelling of the potential for transfer of life between celestial bodies, suggests that life could be more common than previously thought. Here we examine critically what it means to be an extremophile, and the implications of this for evolution, biotechnology and especially the search for life in the Universe.

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Life in Extreme Environments

The fungal dimension of biodiversity: magnitude, significance, and conservation

Desiccation tolerance of prokaryotes.

More than three-quarters of the Earth's surface is occupied by cold ecosystems, including the ocean depths, and polar and alpine regions. These permanently cold environments have been successfully colonized by a class of extremophilic microorganisms that are known as psychrophiles (which literally means cold-loving). The ability to thrive at temperatures that are close to, or below, the freezing point of water requires a vast array of adaptations to maintain the metabolic rates and sustained growth compatible with life in these severe environmental conditions.

Psychrophilic enzymes: Hot topics in cold adaptation

Calcium oxalate (CaOx) crystals are distributed among all taxonomic levels of photosynthetic organisms from small algae to angiosperms and giant gymnosperms. Accumulation of crystals by these organisms can be substantial. Major functions of CaOx crystal formation in plants include high-capacity calcium (Ca) regulation and protection against herbivory. Ultrastructural and developmental analyses have demonstrated that this biomineralization process is not a simple random physical-chemical precipitation of endogenously synthesized oxalic acid and environmentally derived Ca. Instead, crystals are formed in specific shapes and sizes. Genetic regulation of CaOx formation is indicated by constancy of crystal morphology within species, cell specialization, and the remarkable coordination of crystal growth and cell expansion. Using a variety of approaches, researchers have begun to unravel the exquisite control mechanisms exerted by cells specialized for CaOx formation that include the machinery for uptake and accumulation of Ca, oxalic acid biosynthetic pathways, and regulation of crystal growth.

CALCIUM OXALATE IN PLANTS: Formation and Function

In the frigid desert of the Antarctic dry valleys there are no visible life forms on the surface of the soil or rocks. Yet in certain rock types a narrow subsurface zone has a favorable microclimate and is colonized by microorganisms. Dominant are lichens of unusual organization. They survive not by physiological adaptation to lower temperatures, but by changing their mode of growth, being able to grow between the crystals of porous rocks. Their activity results in mobilization of iron compounds and in rock weatherning with a characteristic pattern of exfoliation. This simple ecosystem lacks both higher consumers and predators.

https://science.sciencemag.org/content/sci/215/4536/1045.full.pdf

Endolithic Microorganisms in the Antarctic Cold Desert

The Atacama along the Pacific Coast of Chile and Peru is one of the driest and possibly oldest deserts in the world. It represents an extreme habitat for life on Earth and is an analog for life in dry conditions on Mars. We report on four years (September 1994-October 1998) of climate and moisture data from the extreme arid region of the Atacama. Our data are focused on understanding moisture sources and their role in creating suitable environments for photosynthetic microorganisms in the desert surface. The average air temperature was 16.5 degrees C and 16.6 degrees C in 1995 and 1996, respectively. The maximum air temperature recorded was 37.9 degrees C, and the minimum was -5.7 degrees C. Annual average sunlight was 336 and 335 W m(-2) in 1995 and 1996, respectively. Winds averaged a few meters per second, with strong fohn winds coming from the west exceeding 12 m s(-1). During our 4 years of observation there was only one significant rain event of 2.3 mm, which occurred near midnight local time. We suggest that this event was a rainout of a heavy fog. It is of interest that the strong El Nino of 1997-1998 brought heavy rainfall to the deserts of Peru, but did not bring significant rain to the central Atacama in Chile. Dew occurred at our station frequently following high nighttime relative humidity, but is not a significant source of moisture in the soil or under stones. Groundwater also does not contribute to surface moisture. Only the one rain event of 2.3 mm resulted in liquid water in the soil and beneath stones for a total of only 65-85 h over 4 years. The paucity of liquid water under stones is consistent with the apparent absence of hypolithic (under-stone) cyanobacteria, the only known primary producers in such extreme deserts.

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Temperature and moisture conditions for life in the extreme arid region of the Atacama desert: four years of observations including the El Nino of 1997-1998

Endolithic unicellular blue-green algae occur under the surface of orthoquartzite rocks in the dry valleys of southern Victoria Land, Antarctica. This report of primary producers in the Antarctic desert ecosystem suggests that, in future efforts to detect life in extraterrestrial (for example, martian) environments, scientists should consider the possible existence of endolithic life forms.

https://science.sciencemag.org/content/sci/193/4259/1247.full.pdf

Endolithic Blue-Green Algae in the Dry Valleys: Primary Producers in the Antarctic Desert Ecosystem

In the McMurdo Dry Valleys of Antarctica, microorganisms colonize the pore spaces of exposed rocks and are thereby protected from the desiccating environmental conditions on the surface. These cryptoendolithic communities have received attention in microscopy and culture-based studies but have not been examined by molecular approaches. We surveyed the microbial biodiversity of selected cryptoendolithic communities by analyzing clone libraries of rRNA genes amplified from environmental DNA. Over 1,100 individual clones from two types of cryptoendolithic communities, cyanobacterium dominated and lichen dominated, were analyzed. Clones fell into 51 relatedness groups (phylotypes) with ≥98% rRNA sequence identity (46 bacterial and 5 eucaryal). No representatives of Archaea were detected. No phylotypes were shared between the two classes of endolithic communities studied. Clone libraries based on both types of communities were dominated by a relatively small number of phylotypes that, because of their relative abundance, presumably represent the main primary producers in these communities. In the lichen-dominated community, three rRNA sequences, from a fungus, a green alga, and a chloroplast, of the types known to be associated with lichens, accounted for over 70% of the clones. This high abundance confirms the dominance of lichens in this community. In contrast, analysis of the supposedly cyanobacterium-dominated community indicated, in addition to cyanobacteria, at least two unsuspected organisms that, because of their abundance, may play important roles in the community. These included a member of the α subdivision of the Proteobacteria that potentially is capable of aerobic anoxygenic photosynthesis and a distant relative of Deinococcus that defines, along with other Deinococcus-related sequences from Antarctica, a new clade within the Thermus-Deinococcus bacterial phylogenetic division.

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Microbial Diversity of Cryptoendolithic Communities from the McMurdo Dry Valleys, Antarctica

Endolithic microorganisms (those living inside rocks) occur in hot and cold deserts and exist under extreme environmental conditions. These conditions are discussed on a comparative basis. Quantitative estimates of biomass are comparable in hot and cold deserts. Despite the obvious differences between the hot and cold desert environment, survival strategies show some common features. These endolithic organisms are able to ‘switch’ rapidly their metabolic activities on and off in response to changes in the environment. Conditions in hot deserts impose a more severe environmental stress on the organisms than in the cold Antarctic desert. This is reflected in the composition of the microbial flora which in hot desert rocks consist entirely of prokaryotic microorganisms, while under cold desert conditions eukaryotes predominate.

E. Imre Friedmann

Papers

Endolithic Microorganisms in the Antarctic Cold Desert

Temperature and moisture conditions for life in the extreme arid region of the Atacama desert: four years of observations including the El Nino of 1997-1998

Endolithic Blue-Green Algae in the Dry Valleys: Primary Producers in the Antarctic Desert Ecosystem

Microbial Diversity of Cryptoendolithic Communities from the McMurdo Dry Valleys, Antarctica

Endolithic microbial life in hot and cold deserts