Five Nodulation Mutants of White Sweetclover (Melilotus alba Desr.) Exhibit Distinct Phenotypes Blocked at Root Hair Curling, Infection Thread Development, and Nodule Organogenesis

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Abstract
In an effort to obtain a developmental sequence of mutations in the Rhizobium-legume interaction within a single legume species, we have characterized the early events of nodule development in 10 nodulation mutants of sweetclover, Melilotus alba Desr. cv U389, representing five genetic loci. Both seed and root exudates from all of the sweetclover mutants induced expression of the nod genes of Rhizobium meliloti. Mutants in three loci were blocked in the early stages of root hair curling. Of these, a mutant in the sym-3 locus exhibited root hair deformations in response to inoculation with R. meliloti but produced no nodules or emerging nodule primordia, suggesting a blockage in the signal transduction events leading to nodule organogenesis. In contrast, mutants in both the sym-1 and sym-5 loci formed ineffective nodules in response to inoculation but differed slightly in the type of root hair response observed. None of these three early mutants formed infection threads. Infection threads were observed in mutant sym-2 as well as in ineffective nodules. Mutant sym-4 also formed infection threads but lacked nodules. The phenotypes observed for mutants from these five loci suggest that a secondary receptor or signal produced by the plant is required for nodule development.