Anhydrobiosis: the extreme limit of desiccation tolerance

Extreme habitats give rise to strong stressors that lead organisms to die or to possess specific adaptations to those stressors. One of the most widespread adaptations is quiescence, a common term for several strategies, including anhydrobiosis, a highly stable state of suspended animation due to complete desiccation pending recovery by rehydration. Anhydrobiosis is widespread in nature in a wide taxonomic variety among bacteria, protists, metazoans and plants. Using as model organisms, mainly tardigrades, micrometazoans able to enter anhydrobiosis in any phase of their life cycle from egg to adult, this review presents the response to desiccation from molecules to cells and organisms. Particular emphasis has been done with studies devoted to elucidate phenomena such as the long-term resistance in a desiccated state, the extraordinary resistance to chemical and physical extremes, the morphological, physiological, biochemical, and molecular constraints allowing organisms to enter and to survive anhydrobiosis, and the evolutionary meaning of life without water