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Understanding dinitrogen-fixing symbioses in economically important, temperate, woody legumes requires evaluating their nodulation status and studying molecular mechanisms of nodulation and dinitrogen fixation. Styphnolobium japoniciim (L.) Schott and Cladrastis kentiikea (Dum.-Cours.) Rudd were examined for the capacity to form root nodules. Inoculations with various broad-range rhizobia, soil and rhizobia from closely related species of Sophora, and soil from S. japoniciim and C. kentiikea in Japan, China, and the United States did not elicited nodulation. As an unexpected consequence of these experiments, rhizobia were isolated for the first time from nodules of Maackia floribiinda Takeda, and most tested isolates also infected Maackia amiirensis Rupr. & Maxim. An early nodulation gene, ENOD2, was isolated and described in M. amiirensis, S. japoniciim, and C. kentiikea. The cDNAs had 52 to 82% identity to other ENOD2 sequences, and the cDNAs encoded proteir\s with amino acid compositions and coriserved pentapeptides (PPHEK, PPYEK, and PPEYQ) characteristic of ENOD2 proteins. Southern analyses confirmed cDNAs in each species belonged to small gene families. Transcripts were detected in nodules, roots, and flowers from M. amiirensis, and in roots, stems, and flowers from S. japoniciim and C. kentiikea. Four days after inoculation with rhizobia, ENOD2 transcripts were detected in roots of M. amiirensis, and expression was enhanced as nodules developed. In situ hybridization showed that ENOD2 transcripts are restricted to the distribution zone in indeterminant nodules of M. amurensis. Pseudonodules developed on roots of M. amiirensis after treatment with TIBA, an auxin transport inhibitor, but did not form on S. japoniciim and C. kentiikea. Zeatin and TIBA enhanced EN0D2 expression in roots of M. amurensis only during the first 10 days of a 40-day treatment. Although transcript accumulation in roots of S. japoniciim and C. kentiikea was inhibited initially after exposure to TIBA and zeatin, acciimulation was erUianced after 30 days. My results suggest ENOD2 activity is not a marker for nodulation in legumes, but instead may be a cellular response to concentrations of plant hormones in diverse tissues. In M. amurensis, ENOD2 proteins may be components of cell walls in tissues that regulate nutrient flow to sinks.

Nodulation and expression of the early nodulation gene, ENOD2, in temperate woody legumes of the Papilionoideae