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The ability of anaerobic microorganisms to proliferate under extreme conditions is of widespread importance for microbial physiology, remediation, industry, and evolution. The halophilic alkalithermophiles are a novel group of polyextremophiles. Tolerance to alkaline pH, elevated NaCl concentrations, and high temperatures necessitates mechanisms for cytoplasmic pH acidification; permeability control of the cell membrane; and stability of proteins, the cell wall, and other cellular constituents to multiple extreme conditions. Although it is generally assumed that extremophiles growing at more than one extreme combine adaptive mechanisms for each individual extreme, adaptations for individual extremes often counteract each other. However, in alkaline, hypersaline niches heated via intense solar irradiation, culture-independent analyses have revealed the presence of an extensive diversity of aerobic and anaerobic microorganisms belonging to Bacteria and Archaea that survive and grow under multiple harsh conditions. Thus, polyextremophiles must have developed novel adaptive strategies enabling them to grow and proliferate under multiple extreme conditions. The recent isolation of two novel anaerobic, halophilic alkalithermophiles, Natranaerobius thermophilus and Halonatronum saccharophilum, will provide a platform for detailed biochemical, genomic, and proteomic experiments, allowing a greater understanding of the novel adaptive mechanisms undoubtedly employed by polyextremophiles. In this review, we highlight growth characteristics, ecology, and phylogeny of the anaerobic halophilic alkalithermophiles isolated. We also describe the bioenergetic and physiological problems posed by growth at the multiple extreme conditions of alkaline pH, high NaCl concentration, and elevated temperature under anoxic conditions and highlight recent findings and unresolved problems regarding adaptation to multiple extreme conditions.

The vertical and seasonal distribution and diversity of archaeal sequences was investigated in a hypersaline, stratified, monomictic lake, Solar Lake, Sinai, Egypt, during the limnological development of stratification and mixing. Archaeal sequences were studied via phylogenetic analysis of 16S rDNA sequences as well as denaturing gradient gel electrophoresis analysis. The 165 clones studied were grouped into four phylogenetically different clusters. Most of the clones isolated from both the aerobic epilimnion and the sulfide-rich hypolimnion were defined as cluster I, belonging to the Halobacteriaceae family. The three additional clusters were all isolated from the anaerobic hypolimnion. Cluster II is phylogenetically located between the genera Methanobacterium and Methanococcus. Clusters III and IV relate to two previously documented groups of uncultured euryarchaeota, remotely related to the genus Thermoplasma. No crenarchaeota were found in the water column of the Solar Lake. The archaeal community in the Solar Lake under both stratified and mixed conditions was dominated by halobacteria in salinities higher than 10%. During stratification, additional clusters, some of which may possibly relate to uncultured halophilic methanogens, were found in the sulfide- and methane-rich hypolimnion.

Methanohalophilus zhilinae, a new alkaliphilic, halophilic, methylotrophic species of methanogenic bacteria, is described. Strain WeN5T (T = type strain) from Bosa Lake of the Wadi el Natrun in Egypt was designated the type strain and was further characterized. This strain was nonmotile, able to catabolize dimethylsulfide, and able to grow in medium with a methyl group-containing substrate (such as methanol or trimethylamine) as the sole organic compound added. Sulfide (21 mM) inhibited cultures growing on trimethylamine. The antibiotic susceptibility pattern of strain WeN5T was typical of the pattern for archaeobacteria, and the guanine-plus-cytosine content of the deoxyribonucleic acid was 38 mol%. Characterization of the 16S ribosomal ribonucleic acid sequence indicated that strain WeN5T is phylogenetically distinct from members of previously described genera other than Methanohalophilus and supported the partition of halophilic methanogens into their own genus.