Rhizobium nodulation (Nod) factors are lipo-chitooligosaccharides that act as symbiotic signals, eliciting several key developmental responses in the roots of legume hosts. Using nodulation-defective mutants of Medicago truncatula , we have started to dissect the genetic control of Nod factor transduction. Mutants in four genes ( DMI1 , DMI2 , DMI3 , and NSP ) were pleiotropically affected in Nod factor responses, indicating that these genes are required for a Nod factoractivated signal transduction pathway that leads to symbiotic responses such as root hair deformations, expressions of nodulin genes, and cortical cell divisions. Mutant analysis also provides evidence that Nod factors have a dual effect on the growth of root hair: inhibition of endogenous (plant) tip growth, and elicitation of a novel tip growth dependent on (bacterial) Nod factors. dmi1 , dmi2 , and dmi3 mutants are also unable to establish a symbiotic association with endomycorrhizal fungi, indicating that there are at least three common steps to nodulation and endomycorrhization in M. truncatula and providing further evidence for a common signaling pathway between nodulation and mycorrhization.
INTRODUCTIONSymbiotic bacteria of the genera Rhizobium , Bradyrhizobium , Azorhizobium , and Sinorhizobium , collectively referred to as rhizobia, are able to elicit on their leguminous hosts the formation of specialized organs, called nodules, capable of fixation of atmospheric nitrogen. Among the earliest visible manifestations of symbiotic development are the infection of root hair cells by bacteria, contained in plant-derived infection structures called infection threads, and the induction of cortical divisions to form the nascent nodule primordium (Mylona et al., 1995;Schultze and Kondorosi, 1998).Genetic analysis of rhizobia has led to identification of nod genes, which are involved in the control of host specificity, infection, and nodulation. nod gene expression is under the control of plant signals, essentially flavonoids, excreted in the rhizosphere. Once activated, nod genes specify the synthesis of nodulation (Nod) factors, which are lipo-chitooligosaccharides (Dénarié et al., 1996). Purified Nod factors are capable of eliciting in the roots of the legume hosts many of the plant responses characteristic of the bacteria themselves (Dénarié and Cullimore, 1993): root hair deformations, activation of the plant genes that are specifically induced during early stages of nodulation (early nodulin genes), initiation of cortical cell division, and triggering of a plant organogenic program that leads to nodule formation. Nod factors thus act as symbiotic signaling molecules for initiating nodule development. They also play a key role in the control of specificity of infection and nodulation, the result of particular substitutions present on the basic backbone of the Nod factor molecule (reviewed in Long, 1996; Cohn et al., 1998;Schultze and Kondorosi, 1998).One of the most challenging areas of research in the study of the Rhizobium-legume symbiosis...