Drying to equilibrium with the air kills nearly all animals and flowering plants, including livestock and crops. This makes drought a key ecological problem for terrestrial life and a major cause of human famine. However, the ability to tolerate complete desiccation is widespread in organisms that are either <5路mm long or found mainly where desiccation-sensitive organisms are scarce. This suggests that there is a trade-off between desiccation tolerance and growth. Recent molecular and biochemical research shows that organisms tolerate desiccation through a set of mechanisms, including sugars that replace water and form glasses, proteins that stabilize macromolecules and membranes, and anti-oxidants that counter damage by reactive oxygen species. These protections are often induced by drying, and some of the genes involved may be homologous in microbes, plants and animals. Understanding how mechanisms of desiccation tolerance may constrain growth might show how to undo the constraint in some economically important macroorganisms and elucidate the muchstudied but elusive relationship between tolerance of stress and productivity.Key words: animal, desiccation, drought, growth, microbe, plant, productivity, tolerance, trade-off.
SummaryThe potential of -100路MPa. These thresholds also clearly separate desiccation-sensitive from -tolerant species (Alpert, 2005): there is a gap in the minimum water contents that different living things can survive. Except for a small proportion of seeds (Tweddle et al., 2003), almost all species tested either die if dried to 20% water content, and are thus desiccationsensitive, or survive drying to 10% water content and thus tolerate desiccation. It should be noted that 'desiccation tolerance' has sometimes been used to mean tolerance of partial desiccation by organisms that die if they desiccate completely, as in the literature on insects and intertidal algae. Here, the term will be used to mean tolerance of complete desiccation, defined as drying to equilibrium with moderately to very dry air, or to 10% water content or less.A prime secret of desiccation tolerance appears to be sugars (Alpert and Oliver, 2002). Certain sugars, mainly nonreducing disaccharides, may take the place of water in preventing the aggregation of macromolecules and the disintegration of membranes as cells dry. Many tolerant plants accumulate high concentrations of the familiar disaccharide sucrose (Vicre et al., 2004a). Many desiccation-tolerant animals and microbes and also some plants synthesize the disaccharide trehalose (Wingler, 2002;Breeuwer et al., 2003;Elbein et al., 2003;Crowe et al., 2005). In tandem with specific proteins (Goyal et al., 2005b), these sugars probably stabilize drying cells both by direct interaction with macromolecules and membranes and by reversibly immobilizing cytoplasm in an extremely slow-flowing liquid, a glass (Buitink and Leprince, 2004). Interestingly, sugar glasses also tend to form at the threshold of 10% water content, at least in seeds (Walters et al., 2005).Researchers ...