A Model for the Development of the Rhizobial and Arbuscular Mycorrhizal Symbioses in Legumes and Its Use to Understand the Roles of Ethylene in the Establishment of these two Symbioses

We propose a model depicting the development of nodulation and arbuscular mycorrhizae. Both processes are dissected into many steps, using Pisum sativum L. nodulation mutants as a guideline. For nodulation, we distinguish two main developmental programs, one epidermal and one cortical. Whereas Nod factors alone affect the cortical program, bacteria are required to trigger the epidermal events. We propose that the two programs of the rhizobial symbiosis evolved separately and that, over time, they came to function together. The distinction between these two programs does not exist for arbuscular mycorrhizae development despite events occurring in both root tissues. Mutations that affect both symbioses are restricted to the epidermal program. We propose here sites of action and potential roles for ethylene during the formation of the two symbioses with a specific hypothesis for nodule organogenesis. Assuming the epidermis does not make ethylene, the microsymbionts probably first encounter a regulatory level of ethylene at the epidermis–outermost cortical cell layer interface. Depending on the hormone concentrations there, infection will either progress or be blocked. In the former case, ethylene affects the cortex cytoskeleton, allowing reorganization that facilitates infection; in the latter case, ethylene acts on several enzymes that interfere with infection thread growth, causing it to abort. Throughout this review, the difficulty of generalizing the roles of ethylene is emphasized and numerous examples are given to demonstrate the diversity that exists in plants

Cytokinin accumulation and an altered ethylene response mediate the pleiotropic phenotype of the pea nodulation mutant R50 (sym16)

R50 (sym16), a pleiotropic mutant of Pisum sativum L., is short, has thickened internodes and roots, and has a reduced number of lateral roots and nodules. Its low nodule phenotype can be restored with the application of ethylene inhibitors; furthermore, it can be mimicked by applying cytokinins (CKs) to the roots of the parent line 'Sparkle'. Here, we report on the etiolation phenotypes of R50 and 'Sparkle', and on the interactive roles of ethylene and CKs in these lines. R50 displayed an altered etiolation phenotype, as it was shorter and thicker, and had more developed leaves than dark-grown 'Sparkle'. Shoot morphological differences induced by exogenous ethylene or CKs were found to be less severe for R50. Ethylene inhibitor application induced root and shoot elongation and encouraged apical hook opening in both etiolated lines. Liquid chromatography - tandem mass spectrometry analysis indicated that CK concentrations in R50 were higher than in 'Sparkle', particularly in mature shoots where the levels were maintained at elevated concentrations. These differences indicate a reduction in the CK catabolism of R50. The accumulation of CKs can be directly related to several traits of R50, with the reduced number of nodules and altered shoot ethylene response being likely indirect effects