3One of the most severe and widespread problems facing the agricultural industry is the degradation of soil quality due to desiccation and salinity. In fact, almost 40% of the world's land surface is affected by salinity-related problems (131). These two harsh environmental conditions can have a dramatic impact on the endogenous soil bacteria (38, 48). Of particular importance to the agricultural industry is the impact of these harsh environmental conditions on the endogenous group of proteobacteria, the rhizobia. These bacteria induce formation of nodules on legumes, in which atmospheric nitrogen is fixed and supplied to the plant, enhancing growth under nitrogenlimiting conditions. Desiccation and salinity negatively affect such interactions by limiting nitrogen fixation (131).The importance of nitrogen fixation for agriculture cannot be understated and is illustrated by the numerous studies of the impact of soil management on rhizobial populations in arid regions (52), as well as the isolation and characterization of desiccation-and salt-resistant strains (28, 56,128). Furthermore, to enhance nodulation and nitrogen fixation efficiency, techniques that allow close contact between the bacteria and the host seed have been developed. Despite such methods, there has been a decline in the number of viable bacteria on plant seeds, soil, and carrier material, in part because of the stresses caused by fertilizer and chemical applications (110), desiccation (100, 101), temperature changes (63, 74, 116), salinity changes (63), light (68), and growth media employed (23,35,63). These factors are encountered during the manufacture, storage, and use of the coated seeds, with desiccation as the principal cause of reduced bacterial survival on the seed (33, 119).Although improvement of long-term survival and seed inoculum storage time has been the focus of desiccation research (25, 33), relatively little work has focused primarily on the bacterial cell. Many questions remain regarding the physiological response of rhizobia to desiccation. In this review, we evaluate studies of the physiological responses of rhizobia to environmental stresses (osmotic, salt, temperature, and oxygen) that affect desiccation survival. Our discussion will focus on one species of rhizobia in particular, Sinorhizobium meliloti.
DESICCATION RESPONSES OF RHIZOBIAEarly rhizobial desiccation research. Desiccation produces many stress responses in the bacterial cell. In 1932, Fred and coworkers reported loss of viability in rhizobia used as seed inocula (41) and suggested that the nature of the suspending medium, pH, and temperature are important factors in the survival of the inoculum in the dry state. This led to the recommendation for farmers to refrain from using rhizobia in dry form. Vincent et al. (119) showed that the decline in abundance of Rhizobium trifolii during drying on glass beads correlates with the extraction of water, which suggests that part of the decline in viable counts is caused by both "seed factors" and the drying itself. The ...