The microfossil record suggests that cyanobacteria or cyanobacteria-like prokaryotes were present on the primitive Earth in the Archaean era more than 3.5 billion years ago (28). The exquisite preservation of these microfossils is thought to reflect the intrinsic stability of the extracellular polysaccharide (EPS) and its ability to bind heavy metals as well as resist degradation (13). Extant cyanobacteria dominate the microbial populations of many extreme environments including soda lakes (Spirulina, Cyanospira), the nutrient-poor open ocean (Trichodesmium), thermal springs (Synechococcus and Mastigocladis), and the cold dry polar deserts (Chroococcidiopsis) (35). In these environments the cyanobacteria produce copious amounts of EPSs in the form of sheaths, slimes, and capsules. Very little is known about the diversity, mode of synthesis, structure, or properties of these biopolymers (19). A recent review emphasized the potential role of EPSs in the desiccation tolerance of prokaryotes (23). However, much further research is needed to resolve the specific mechanisms which biopolymers contribute to such a complex process.The terrestrial cyanobacterium Nostoc commune has a marked capacity for desiccation tolerance and can survive storage at Ϫ400 MPa (0% relative humidity) for centuries (23). The cells produce large amounts of an unusual excreted polysaccharide that contributes in at least four ways to the marked stabilization of cells during prolonged storage in the air-dried state, at low or high temperatures. First, the glycan inhibits fusion of membrane vesicles during desiccation and freezedrying (10) and acts as an immobilization matrix for a range of secreted enzymes which remain fully active after long-term air-dried storage (11,27,32). Second, the glycan provides a structural and/or molecular scaffold with rheological properties which can accommodate the rapid biophysical and physiological changes in the community upon rehydration and during recovery from desiccation. The glycan swells from brittle dried crusts to cartilaginous structures within minutes of rehydration. Third, the glycan matrix contains both lipid-and watersoluble UV radiation-absorbing pigments which protect the cell from photodegradation (12). Fourth, although epiphytes colonize the surfaces of Nostoc colonies, there is no penetration of the glycan due in part to a silicon-and calcium-rich pellicle and inherent resistance of the glycan to enzymatic breakdown. Preliminary structural work on one water-soluble UV-absorbing pigment (released from the glycan by acid hydrolysis) indicated the presence of an oligosaccharide (4), raising the possibility that the pigment may be covalently linked to the glycan in the desiccated state.An understanding of the biochemical and biophysical properties of such biopolymers and the isolation of genes and enzymes required for their synthesis and modification can lead to an understanding of the underlying principles of extremophile stability. Furthermore, one can envision the utilization of such materials f...