Diversity of anaerobic halophilic microorganisms
Life in the presence of high salt concentrations is compatible with life in the absence of oxygen. Halophilic and halotolerant anaerobic prokaryotes are found both in the archaeal and in the bacterial domain, and they display a great metabolic diversity. Many of the representatives of the Halobacteriales (Archaea), which are generally considered aerobes, have the potential of anaerobic growth. Some can use alternative electron acceptors such as nitrate, fumarate, dimethylsulfoxide or trimethylamine-N-oxide Halobacterium salinarum can also grow fermentatively on L-arginine, and bacteriorhodopsin-containing cells may even grow anaerobically, energized by light. Obligatory anaerobic halophilic methanogenic Archaea also exist. The bacterial domain contains many anaerobic halophiles, including sulfate reducers. There is also a group of specialized obligatory anaerobic Bacteria, phylogenetically clustering in the low G+C branch of the Firmicutes. Most representatives of this group (order Haloanaerobiales , families Haloanaerobiaceae and Halobacteroidaceae ) are fermentative, using a variety of carbohydrates and amino acids. One species combines the potential for anaerobic growth at high salt concentrations with a preference for high temperatures. Others are homoacetogens; Acetohalobium arabaticum can grow anaerobically as a chemolithotroph, producing acetate from hydrogen and CO 2 . The Haloanaerobiales accumulate high concentrations of K + and Cl - in their cytoplasm, thereby showing a strategy of salt adaptation similar to that used by the Halobacteriales . Recently a new representative of the Haloanaerobiales was isolated from bottom sediments of the Dead Sea (strain DSSe1), which grows anaerobically by oxidation of glycerol to acetate and CO 2 while reducing selenate to selenite and elementary selenium. Other electron acceptors supporting anaerobic growth of this strain are nitrate and trimethylamine- N -oxide. The versatility of life at high salt concentrations with respect to the variety of substrates used, the types of dissimilatory metabolism, and the diversity of potential electron acceptors has important implications for the potential for life in hostile environments lacking oxygen and high in salt, implications that may also be relevant to astrobiology.