Spores of Bacillus megaterium QM B1551 germinate in response to a number of trigger compounds, including glucose, proline, leucine, and inorganic salts. An approximate 6-kb region of the 165-kb plasmid was found to harbor a tricistronic receptor operon, gerU, and a monocistronic receptor component, gerVB. The gerU operon was observed to complement the germination response in plasmidless strain PV361 to glucose and leucine, with KBr acting as a cogerminant. Proline recognition is conferred by the monocistronic gerVB gene, the presence of which also improves the germination response to other single-trigger compounds. A chimeric receptor, GerU*, demonstrates interchangeability between receptor components and provides evidence that it is the B protein of the receptor that determines germinant specificity. Introduction of the gerU/gerVB gene cluster to B. megaterium KM extends the range of germinants recognized by this strain to include glucose, proline, and KBr in addition to alanine and leucine. A chromosomally encoded receptor, GerA, the B component of which is predicted to be truncated, was found to be functionally redundant. Similarly, the plasmid-borne antiporter gene, grmA, identified previously as being essential for germination in QM B1551, did not complement the germination defect in the plasmidless variant PV361. Wild-type spores carrying an insertion-deletion mutation in this cistron germinated normally; thus, the role of GrmA in spore germination needs to be reevaluated in this species.Bacterial spores rely upon a number of small molecules, including amino acids, purine ribosides, sugars, and ions, to indicate that environmental conditions are favorable for the resumption of growth. The rapid transition from extreme dormancy to loss of resistance properties and resumption of metabolism displayed by spores upon exposure to germinant molecules is regulated by a process known as spore germination (22,23,32). Germinant interaction with cognate receptors located within the inner spore membrane (11, 26) stimulates changes in the permeability of this membrane to various metal ions and Ca-dipicolinic acid (DPA), which are effluxed from the spore core, while permitting partial hydration of the spore core. Enzymatic hydrolysis of the spore's thick layer of cortical peptidoglycan and protective outer coat layers is then initiated, permitting complete hydration and the resumption of vegetative metabolism. Since strict germination events precede macromolecular synthesis, the germination apparatus must be present in the dormant spore (22,32).A substantial amount of genetic evidence has been presented that strongly suggests that orthologous proteins belonging to the GerA family form the receptors through which the spore senses its environment (21,23,25). GerA structural genes are arranged in tricistronic operons, which are expressed during sporulation in the developing forespore (7). Multiple copies of these receptor operons reside within the genomes of all Bacillus species examined to date. Hydropathy profiling indica...