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Published on March 16, 2007; 10.1104/pp.106.093021

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Received November 14, 2006
Accepted March 7, 2007

Medicago truncatula NIN Is Essential for Rhizobium-Independent Nodule Organogenesis Induced by Autoactive CCaMK

John F. Marsh *, Alexandra Rakocevic , Raka M. Mitra , Lysiane Brocard , Jongho Sun , Alexis Eschstruth , Sharon R. Long , Michael Schultze , Pascal Ratet , and Giles E.D. Oldroyd

John Innes Centre, Norwich Research Park, Colney, Norwich, NR4 7UH, UK; Institut des Sciences du Vegetale, CNRS, Avenue de la Terrasse, 91198, Gif sur Yvette, France; Department of Biological Sciences, 371 Serra Mall, Stanford University, Stanford, CA 94305-5020; Department of Biology, University of York, YO10 5YW, UK

* Corresponding author; email: john.marsh{at}bbsrc.ac.uk.

The symbiotic association between legumes and nitrogen fixing bacteria collectively known as rhizobia results in the formation of a unique plant root organ called the nodule. This process is initiated following the perception of rhizobial nodulation factors by the host plant. Nod factor (NF) stimulated plant responses, including nodulation specific gene expression, is mediated by the early NF signaling pathway. Plant mutants in this pathway are unable to nodulate. We describe here the cloning and characterization of two mutant alleles of the Medicago truncatula ortholog of the Lotus japonicus and pea NIN gene. The Mtnin mutants undergo excessive root hair curling, but are impaired in infection and fail to form nodules following inoculation with Sinorhizobium meliloti. Our investigation of early NF-induced gene expression using the reporter fusion ENOD11::GUS in the Mtnin-1 mutant, demonstrates that MtNIN is not essential for early NF signaling, but may negatively regulate the spatial pattern of ENOD11 expression. It was recently shown that an autoactive form of a nodulation-specific calcium/calmodulin-dependent protein kinase (CCaMK) is sufficient to induce nodule organogenesis in the absence of rhizobia. We show here that MtNIN is essential for autoactive CCaMK-induced nodule organogenesis. The non-nodulating hcl mutant has a similar root hair-curling phenotype to Mtnin, but we demonstrate that HCL is not required in this process. Based on our data, we suggest that MtNIN functions downstream of the early NF signaling pathway to coordinate and regulate the correct temporal and spatial formation of root nodules.




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