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      How Many Peas in a Pod? Legume Genes Responsible for Mutualistic Symbioses Underground

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          Abstract

          The nitrogen-fixing symbiosis between legume plants and Rhizobium bacteria is the most prominent plant–microbe endosymbiotic system and, together with mycorrhizal fungi, has critical importance in agriculture. The introduction of two model legume species, Lotus japonicus and Medicago truncatula, has enabled us to identify a number of host legume genes required for symbiosis. A total of 26 genes have so far been cloned from various symbiotic mutants of these model legumes, which are involved in recognition of rhizobial nodulation signals, early symbiotic signaling cascades, infection and nodulation processes, and regulation of nitrogen fixation. These accomplishments during the past decade provide important clues to understanding not only the molecular mechanisms underlying plant–microbe endosymbiotic associations but also the evolutionary aspects of nitrogen-fixing symbiosis between legume plants and Rhizobium bacteria. In this review we survey recent progress in molecular genetic studies using these model legumes.

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          Most cited references142

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          CERK1, a LysM receptor kinase, is essential for chitin elicitor signaling in Arabidopsis.

          Chitin is a major component of fungal cell walls and serves as a microbe-associated molecular pattern (MAMP) for the detection of various potential pathogens in innate immune systems of both plants and animals. We recently showed that chitin elicitor-binding protein (CEBiP), plasma membrane glycoprotein with LysM motifs, functions as a cell surface receptor for chitin elicitor in rice. The predicted structure of CEBiP does not contain any intracellular domains, suggesting that an additional component(s) is required for signaling through the plasma membrane into the cytoplasm. Here, we identified a receptor-like kinase, designated CERK1, which is essential for chitin elicitor signaling in Arabidopsis. The KO mutants for CERK1 completely lost the ability to respond to the chitin elicitor, including MAPK activation, reactive oxygen species generation, and gene expression. Disease resistance of the KO mutant against an incompatible fungus, Alternaria brassicicola, was partly impaired. Complementation with the WT CERK1 gene showed cerk1 mutations were responsible for the mutant phenotypes. CERK1 is a plasma membrane protein containing three LysM motifs in the extracellular domain and an intracellular Ser/Thr kinase domain with autophosphorylation/myelin basic protein kinase activity, suggesting that CERK1 plays a critical role in fungal MAMP perception in plants.
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            Plant recognition of symbiotic bacteria requires two LysM receptor-like kinases.

            Although most higher plants establish a symbiosis with arbuscular mycorrhizal fungi, symbiotic nitrogen fixation with rhizobia is a salient feature of legumes. Despite this host range difference, mycorrhizal and rhizobial invasion shares a common plant-specified genetic programme controlling the early host interaction. One feature distinguishing legumes is their ability to perceive rhizobial-specific signal molecules. We describe here two LysM-type serine/threonine receptor kinase genes, NFR1 and NFR5, enabling the model legume Lotus japonicus to recognize its bacterial microsymbiont Mesorhizobium loti. The extracellular domains of the two transmembrane kinases resemble LysM domains of peptidoglycan- and chitin-binding proteins, suggesting that they may be involved directly in perception of the rhizobial lipochitin-oligosaccharide signal. We show that NFR1 and NFR5 are required for the earliest physiological and cellular responses to this lipochitin-oligosaccharide signal, and demonstrate their role in the mechanism establishing susceptibility of the legume root for bacterial infection.
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              The CLAVATA1 gene encodes a putative receptor kinase that controls shoot and floral meristem size in Arabidopsis.

              The shoot apical meristem is responsible for above-ground organ initiation in higher plants, accomplishing continuous organogenesis by maintaining a pool of undifferentiated cells and directing descendant cells toward organ formation. Normally, proliferation and differentiation are balanced, so that the structure and size of the shoot meristem is maintained. However, Arabidopsis plants homozygous for mutations at the CLAVATA1 (CLV1) locus accumulate excess undifferentiated cells. We describe the molecular cloning and expression pattern of the CLV1 gene. It encodes a putative receptor kinase, suggesting a role in signal transduction. The extracellular domain is composed of 21 tandem leucine-rich repeats that resemble leucine-rich repeats found in animal hormone receptors. We provide evidence that CLV1 expression in the inflorescence is specifically associated with meristematic activity.
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                Author and article information

                Journal
                Plant Cell Physiol
                pcp
                pcellphys
                Plant and Cell Physiology
                Oxford University Press
                0032-0781
                1471-9053
                September 2010
                21 July 2010
                21 July 2010
                : 51
                : 9
                : 1381-1397
                Affiliations
                1Department of Plant Sciences, National Institute of Agrobiological Sciences, Tsukuba, 305-8602 Japan
                2Department of Life Science, Aichi University of Education, Kariya, 448-8542 Japan
                3Department of Evolutionary Biology and Biodiversity, National Institute for Basic Biology, Okazaki, 444-8585 Japan
                4Present address: Graduate School of Agricultural and Life Sciences, The University of Tokyo, Tokyo, 113-8657 Japan
                Author notes
                *Corresponding author: E-mail, kouchih@ 123456nias.affrc.go.jp ; Fax, +81-29-838-8347
                Article
                pcq107
                10.1093/pcp/pcq107
                2938637
                20660226
                3a19e100-278a-4a8e-a0d8-08fb2653aa10
                © The Author 2010. Published by Oxford University Press on behalf of Japanese Society of Plant Physiologists.

                This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License ( http://creativecommons.org/licenses/by-nc/2.5), which permits unrestricted non-commercial use distribution, and reproduction in any medium, provided the original work is properly cited.

                History
                : 28 May 2010
                : 17 July 2010
                Categories
                Special Issue Mini Reviews

                Plant science & Botany
                model legumes,nitrogen fixation,lotus japonicus,medicago truncatula,nodules,plant–microbe symbiosis

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