1State Key Laboratory of Rice Biology, China National Rice Research Institute, Hangzhou 310006, China; 2College of Life and Environment Science, Hangzhou Normal University, Hangzhou 310016, China; 3Key Laboratory of Plant Functional Genomics, Ministry of Education, Yangzhou University, Yangzhou 225009, China; *Corresponding author, E-mail: firstname.lastname@example.org
Abstract A rumpled and twisted leaf 1 (rtl1) mutant was derived from japonica cultivar Nipponbare by ethyl methane sulfonate (EMS) treatment, which was characterized by rumpled and twisted leaf at the seedling stage. The F2 population was constructed by crossing with indica cultivars TN1 and Zhefu 802, respectively. Genetic analysis confirmed that it was controlled by one recessive nuclear gene. The closely linked SSR marker RM1155 was obtained through bulked segregant analysis．Subsequently, new STS markers were developed using published rice genome sequence, and the gene was finally located between the STS marker T1591 and SSR marker RM1359 with the distances of 0.48 and 0.96 cM, respectively. This will contribute to cloning of the target gene in further studies.
Scanlon M J. Developmental complexities of simple leaves. Curr Opin Plant Biol, 2000, 3(1): 31-36.
Bowman J, Eshed Y, Baum S F. Establishment of polarity in angiosperm lateral organs. Trends Genet, 2002, 18(3): 134-141.
Laux T, Mayer K F, Berger J, et al. The WUSCHEL gene is required for shoot and floral meristem integrity in Arabidopsis. Development, 1996, 122(1): 87-96.
Schoof H, Lenhard M, Haecker A, et al. The stem cell population of Arabidopsis shoot meristems in maintained by a regulatory loop between the CLAVATA and WUSCHEL genes. Cell, 2000, 100(6): 635-644.
Fletcher J C, Brand U, Running M P, et al. Signaling of cell fate decisions by CLAVATA3 in Arabidopsis shoot meristems. Science, 1999, 283(5409): 1911-1914.
Lincoln C, Long J, Yamaguchi J, et al. A knotted1-like homeobox gene in Arabidopsis is expressed in the vegetative meristem and dramatically alters leaf morphology when overexpressed in transgenic plants. Plant Cell, 1994, 6(12): 1859-1876.
Ito Y, Eiguchi M, Kurata N. KNOX homeobox genes are sufficient in maintaining cultured cells in an undifferentiated state in rice. Genesis, 2001, 30(4): 231-238.
Sato Y, Hong S K, Tagiri A, et al. A rice homeobox gene, OSH1, is expressed before organ differentiation in a specific region during early embryogenesis. Proc Natl Acad Sci USA, 1996, 93(15): 8117-8122.
Scofield S, Murray J A. KNOX gene function in plant stem cell niches. Plant Mol Biol, 2006, 60(6): 929-946.
Byrne M E, Barley R, Curtis M, et al. Asymmetric leaves 1 mediates leaf patterning and stem cell function in Arabidopsis. Nature, 2000, 408(6815): 967-971.
Ori N, Eshed Y, Chuck C R, et al. Mechanisms that control knox gene expression in the Arabidopsis shoot. Development, 2000, 127(24): 5523-5532.
Semiarti E, Ueno Y, Tsukaya H, et al. The ASYMMETRIC LEAVES2 gene of Arabidopsis thaliana regulates formation of a symmetric lamina, establishment of venation and repression of meristem related homeobox genes in leaves. Development, 2001, 128(10): 1771-1783.
Endrizzi K, Moussian B, Haecker A, et al. The SHOOT MERISTEMLESS gene is required for maintenance of undifferentiated cells in Arabidopsis shoot and floral meristems and acts at a different regulatory level than the meristem genes WUSCHEL and ZWILLE. Plant J, 1996, 10(6): 967-979.
Zhong R, Ye Z H. IFL1, a gene regulating interfascicular fiber differentiation in Arabidopsis, encodes a homeodomain-leucine zipper protein. Plant Cell, 1999, 11(11): 2139-2152.
Ratcliffe O J, Riechmann J L, Zhang J Z. INTERFASCICULAR FIBERLESSI is the same gene as REVOLUTA. Plant Cell, 2000, 129(3): 315-317.
McConnell J R, Emery J, Eshed Y, et al. Role of PHABULOSA and PHAVOLUTA in determining radial patterning in shoots. Nature, 2001, 411(6838): 709-713.
Siegfried K R, Eshed Y, Baum S F, et al. Members of the YABBY gene family specify abaxial cell fate in Arabidopsis. Development, 1999, 126(18): 4117-4128.
Kumaran M K, Bowman J L, Sundaresan V. YABBY polarity genes mediate the repression of KNOX homeobox genes in Arabidopsis. Plant Cell, 2002, 14(11): 2761-2770.
Yamaguchi T, Nagasawa N, Kawasaki S, et al. The YABBY gene DROOPING LEAF regulates carpel specification and midrib development in Oryza sativa. Plant Cell, 2004, 16(2): 500-509.
Eshed Y, Baum S F, Perea J V, et al. Establishment of polarity in lateral organs of plants. Curr Biol, 2001, 11(16): 1251-1260.
Emery J F, Floyd S K, Alvarez J, et al. Radial patterning of Arabidopsis shoots by class ⅢHD-ZIP and KANADⅠgenes. Curr Biol, 2003, 13(20): 1768-1774.
Kerstetter R A, Bollman K, Taylor R A, et al. KANADⅠ regulates organ polarity in Arabidopsis. Nature, 2001, 411(6838): 706-709.
Vogler H, Kuhlemeier C. Simple hormones but complex signaling. Curr Opin Plant Biol, 2003, 6(1): 51-56.
Okadala K, Uedal J, Komaki M K, et al. Requirement of the auxin polar transport system in early stages of Arabidopsis floral bud formation. Plant Cell, 1991, 3(7): 677-684.
Reinhardt D, Pesce E R, Stieger P, et al. Regulation of phyllotaxis by polar auxin transport. Nature, 2003, 426(6964): 255-260.
Rhoades M W, Reinhart B J, Lim L P, et al. Prediction of plant microRNA targets. Cell, 2002, 110(4): 513-520.
Juarez M T, Kui J S, Thomas J, et al. MicroRNA-mediated repression of rolled leaf1 specifies maize polarity. Nature, 2004, 428(6978): 84-88.
Murray M G, Thompson W F. Rapid isolation of high molecular weight plant DNA. Nucl Acids Res, 1980, 8(19): 4321-4325.
Vollbrecht E, Veit B, Sinha N, et al. The developmental gene Knotted-1 is a member of a maize homeobox gene family. Nature, 1991, 350(6315): 241-243.
Smith L G, Greene B, Veit B, et al. A dominant mutation in the maize homeobox gene, Knotted-1, causes its ectopic expression in leaf cells with altered fates. Development, 1992, 116(1): 21-30.
Kano-Murakami Y, Yanai T, Tagiri A, et al. A rice homeotic gene, OSH1, causes unusual phenotypes in transgenic tobacco. FEBS Lett, 1993, 334(3): 365-368.
Sinha N R, Williams R E, Hake S. Overexpression of the maize homeo box gene, KNOTTED-1, causes a switch from determinate to indeterminate cell fates. Genes Dev, 1993, 7(5): 787-795.
Postma-Haarsma A D, Verwoert I I, Stronk O P, et al. Characterization of the KNOX class homeobox genes Oskn2 and Oskn3 identified in a collection of cDNA libraries covering the early stages of rice embryogenesis. Plant Mol Biol, 1999, 39(2): 257-271.
Hareven D, Gutfinger T, Parnis A, et al. The making of a compound leaf: Genetic manipulation of leaf architecture in tomato. Cell, 1996, 84(5): 735-744.
Ma Y, Wang F, Guo J, et al. Rice OsAS2 gene, a member of LOB domain family, functions in the regulation of shoot differentiation and leaf development. J Plant Biol, 2009, 52(5): 374-381.
Dai M, Hu Y, Zhao Y, et al. A WUSCHEL-LIKE HOMEOBOX gene represses a YABBY gene expression required for rice leaf development. Plant Physiol, 2007, 144(1): 380-390.
Hu J, Zhu L, Zeng D, et al. Identification and characterization of NARROW AND ROLLED LEAF 1, a novel gene regulating leaf morphology and plant architecture in rice. Plant Mol Biol, 2010, 73(3): 283-292.
Fujino K, Matsuda Y, Ozawa K, et al. NARROW LEAF 7 controls leaf shape mediated by auxin in rice. Mol Genet Genomics, 2008, 279(5): 499-507.
Qi J, Qian Q, Bu Q, et al. Mutation of the rice Narrow leaf1 gene, which encodes a novel protein, affects vein patterning and polar auxin transport. Plant Physiol, 2008, 147(4): 1947-1459.
Ueguchi-Tanaka M, Ashikari M, Nakajima M, et al. GIBBERELLIN INSENSITIVE DWARF1 encodes a soluble receptor for gibberellin. Nature, 2005, 437(7059): 693-698.
Zhang G H, Xu Q, Zhu X D, et al. SHALLOT-LIKE1 is a KANADⅠ transcription factor that modulates rice leaf rolling by regulating leaf abaxial cell development. Plant Cell, 2009, 21(3): 719-735.
Zhao S Q, Hu J, Guo L B, et al. Rice leaf inclination2, a VIN3-like protein, regulates leaf angle through modulating cell division of the collar. Cell Res, 2010, 20(8): 935-947.
Yamamuro C, Ihara Y, Wu X, et al. Loss of function of a rice brassinosteroid insensitive1 homolog prevents internode elongation and bending of the lamina joint. Plant Cell, 2000, 12(9): 1591-1606.
Tanabe S, Ashikari M, Fujioka S, et al. A novel cytochrome P450 is implicated in brassinosteroid biosynthesis via the characterization of a rice dwarf mutant, dwarf11, with reduced seed length. Plant Cell, 2005, 17(3): 776-790.
Hong Z, Ueguchi-Tanaka M, Umemura K, et al. A rice brassinosteroid-deficient mutant, ebisu dwarf (d2), is caused by a loss of function of a new member of cytochrome P450. Plant Cell, 2003, 15(12): 2900-2910.
Nagasawa N, Miyoshi M, Sano Y, et al. SUPERWOMAN1 and DROOPING LEAF genes control floral organ identity in rice. Development, 2003, 130(4): 705-718.