生长素对水稻根系生长发育调控的研究进展

doi:10.3969/j.issn.1006-8082.2014.04.001

摘要/Abstract

摘要： 水稻根系是非常重要的吸收营养和水分的器官，其生长与发育受多个植物激素协同调控。本文综述了生长素对水稻根形态建成调控的分子遗传学研究进展，影响生长素合成的YUCCA基因是通过控制生长素在植株体内的浓度来改变根的形态。而生长素运输主要通过调控生长素输出载体PIN和生长素输入载体AUXI的极性分布，影响侧根和不定根的发育，以及根的向重性。此外，TIR1、Aux/IAA与ARF互作在生长素信号转导中起重要的调控作用。其他激素可以通过信号转导途径影响生长素的分布，调控根系统的建成。但目前水稻相关基因的克隆进展缓慢，对生长素调控网络认识还不够清晰。因此，更为深入的研究水稻生长素相关基因将对理解水稻根系发育的分子机制具有重要意义。

关键词: 水稻, 根形态建成, 生长素, 基因, 分子机制

Abstract: Rice root is the main organ for nutrient and water absorption, its growth and development are regulated by several plant hormones collaboratively. Of which，auxin is the extensively studied. In this paper, the author elucidate the development process and molecular mechanism of rice root architecture by analyzing the regulating genes that have been cloned. Comprehensive analysis reveals that the root shape is altered by auxin biosynthesis genes that regulate auxin concentration. Auxin transport regulate the development of lateral root and crown root through controlling auxin influx and efflux. Auxin signal transduction, which affects the root system, is regulated by TIR1、Aux/IAA and ARF. Besides the distrubition of auxin is influenced by acting with other plant hormones. However，these cloned root shape genes in rice is still insufficient, and the outline of molecular regulation network of auxin is not clear. Therefore，it is important to further understand molecular mechanism of auxin in rice root by screening and collecting rice auxin mutants with root mutations.

Key words: rice, root morphology, auxin, gene, molecular mechanism

中图分类号:

S511

康书静12, 钱前1, 朱丽1 *. 生长素对水稻根系生长发育调控的研究进展[J]. 中国稻米, 2014, 20(4): 1-8.

KANG Shu-Jing-12, QIAN Qian-1, ZHU Li-1 *. Advances on Molecular Mechanism of Auxin in Rice Root[J]. , 2014, 20(4): 1-8.

参考文献

[1]    Ishikawa H, Evans M L. Specialized zones of development in roots [J]. Plant Physiol, 1995, 109（3）: 725–727.

[2]   Berg V, Willemsen C, Hendriks V, et al. Short-range control of cell differentiation in the Arabidopsis root meristem [J]. Nature, 1997, 390: 287-289.

[3]    Liu H J, Wang S F, Yu X B, et al. ARL1, a LOB-domain protein required for adventitious root formation in rice [J]. Plant J, 2005, 43（1）: 47-56.

[4]    Itoh J, Nonomura K, Ikeda K, et al. Rice plant development: from zygote to spikelet [J]. Plant Cell Physiol, 2005, l46: 23-47.

[5]    Casimiro, Beeekman T, Graham N, et al. Dissecting Arabidopsis lateral root development [J]. Trends Plant Sci, 2003, 8: 165-171.

[6]    Malamy J E. Intrinsic and environmental response pathways that regulate root system architecture [J]. Plant Cell Environ, 2005, 28: 67-77.

[7]    Ljung K. Auxin metabolism and homeostasis during plant development [J]. Development, 2013, 40: 943-950.

[8]    Mashiguchi K, Tanaka K, Sakai T, et al. The main auxin biosynthesis pathway in Arabidopsis [J]. PNAS, 2011, 108（45） : 18512-18517.

[9]    Zhao Y. Auxin biosynthesis: a simple two-step pathway converts tryptophan to indole-3-acetic acid in plants [J]. Mol Plant, 2012, 5（2）: 334-348.

[10] Yamamoto Y, Kamiya N, Morinaka Y, et al. Auxin biosynthesis by the YUCCA genes in rice [J]. Plant Physiol, 2007, 143（3）: 1362-1371.

[11] Woo Y M, Park H J, Park J J, et al. Constitutively wilted 1, a member of the rice YUCCA gene family, is required for maintaining water homeostasis and an appropriate root to shoot ratio [J]. Plant Mol Biol, 2007, 65（1-2）: 125-136.

[12] Sazuka T, Kamiya N, Nishimura T, et al. A rice tryptophan deficient dwarf mutant, tdd1, contains a reduced level of indole acetic acid and develops abnormal flowers and organless embryos [J]. Plant J, 2009, 60（2）: 227-241.

[13] Zhao Z G, Zhang Y H, Liu X, et al. A role for a dioxygenase in auxin metabolism and reproductive development in rice[J]. Dev Cell, 2013, 27（1）: 113-122.

[14] Bhalerao R P, Eklof J, Ljung K, et al. Shoot-derived auxin is essential for early lateral root emergence in Arabidopsis seedlings [J]. Plant J, 2002, 29（3）: 325-332.

[15] Yang H B, Murphy A S. Functional expression and characterization of Arabidopsis ABCB, AUX1 and PIN auxin transporters in Schizosaccharomyces pombe [J]. Plant J, 2009, 59（1）: 179-191.

[16] Elke B, Martin K, Jakub R, et al. A novel putative auxin carrier family regulates intracellular auxin homeostasis in plants [J]. Nature, 2012, 485（7396）: 119-122.

[17] 张俊红，孟成生，张彩英，等．小麦和水稻auxin基因家族的生物信息学比较分析[J]．华北农学报，2009，24（6）：15-19.

[18] Xu M, Zhu L, Shou H X, et al. A PIN1 family gene, OsPIN1, involved in auxin-dependent adventitious root emergence and tillering in rice [J]. Plant Cell Physiol, 2005, 46（10）: 1674-1681.

[19] Kitomi Y, Ogawa A, Kitano H, et al. CRL4 regulates crown root formation through auxin transport in rice [J]. Plant Root, 2008, 2: 19-28.

[20] Liu S P, Wang J R, Wang L, et al. Adventitious root formation in rice requires OsGNOM1 and is mediated by the OsPINs family [J]. Cell Res, 2009, 19（9）: 1110-1119.

[21] Geldner N, Richter S, Vieten A, et al. Partial loss-of-function alleles reveal a role for GNOM in auxin transport-related, post-embryonic development of Arabidopsis [J]. Development, 2004, 131: 389-400.

[22] Christensen S K, Dagenais N, Chory J, et al. Regulation of auxin response by the protein kinase PINOID [J]. Cell, 2000, 100（ 4） : 469-478.

[23] Zhuang X L, Jiang X F, Li J H, et al. Over-expression of OsAGAP, an ARF-GAP, interferes with auxin influx, vesicle trafficking and root development [J]. Plant J, 2006, 48（4）: 581-591.

[24] Zhao F Y, Hu F, Zhang S Y, et al. MAPKs regulate root growth by influencing auxin signaling and cell cycle-related gene expression in cadmium-stressed rice [J]. Environ Sci Pollut Res, 2013, 20: 5449-5460.

[25] Sauer M, Kleine-Vehn J. Auxin binding protein1: the outsider [J]. Plant Cell, 2011, 23（6）: 2033-2043 .

[26] 胡应红，李正国，宋红丽，等. 植物生长素受体 [J]. 植物生理学通讯，2007，43（1）：168-172.

[27] Tan X, Calderon-Villalobos L, Sharon M, et al. Mechanism of auxin perception by the TIR1 ubiquitin ligase [J]. Nature, 2007, 446: 640-645.

[28] Xia K, Wang R, Ou X, et al. OsTIR1 and OsAFB2 downregulation via OsmiR393 overexpression leads to more tillers, early flowering and less tolerance to salt and drought in rice [J]. PLoS ONE, 2012, 7（1）: e30039.

[29] Han Y, Cao H, Jiang J F, et al. Rice ROOT ARCHITECTURE ASSOCIATED1 binds the proteasome subunit RPT4 and is degraded in a D-box and proteasome-dependent manner [J]. Plant Physiol, 2008, 148（2）: 843-855.

[30] Ge L, Chen H, Jiang J F, et al. Overexpression of OsRAA1 causes pleiotropic phenotypes in transgenic rice plants, including altered leaf, flower, and root development and root response to gravity [J]. Plant Physiol, 2004, 135（3）: 1502-1513.

[31] Wang X F, He F F, Ma X X, et al. OsCAND1 is required for crown root emergence in rice [J]. Mol Plant, 2011, 4（2）: 289-299.

[32] Kang B, Zhang Z C, Wang L L, et al. OsCYP2, a chaperone involved in degradation of auxin-responsive proteins, plays crucial roles in rice lateral root initiation [J]. Plant J, 2013, 74（1）: 86-97.

[33] Ramos A, Zenser N, Leyser O, et al. Rapid degradation of auxin/indoleacetic acid proteins requires conserved amino acids of domain II and is proteasome dependent [J]. Plant Cell, 2001, 13: 2349-2360.

[34] Fu J, Yu H H, Li X H, et al. Rice GH3 gene family: regulators of growth and development [J]. Plant Signal Behav, 2011, 6（4）: 570-574.

[35] 鄂志国，王磊. 水稻中生长素作用的分子机理研究进展[J]. 核农学报，2011，25（4）：730 -735.

[36] Zhang S W, Li C H, Cao J, et al. Altered architecture and enhanced drought tolerance in rice via the down-regulation of indole-3-acetic acid by TLD1/OsGH3.13 activation [J]. Plant Physiol, 2009, 151（4）: 1889-1901.

[37] Kitomi Y, Inahashi H, Takehisa H, et al. OsIAA13-mediated auxin signaling is involved in lateral root initiation in rice [J]. Plant Sci, 2012, 190: 11.

[38] Zhu Z X, Liu Y, Liu S J, et al. A gain-of-function mutation in OsIAA11 affects lateral root development in rice [J]. Mol Plant, 2012, 5（1）: 154-161.

[39] Ni J, Wang G H, Zhu Z X, et al. OsIAA23-mediated auxin signaling defines postembryonic maintenance of QC in rice [J]. Plant J, 2011, 68（3）: 433-442.

[40] 吴蓓，吴建勇，蔡刘体，等. 生长素反应因子[J]. 植物生理学通讯，2005，41（3）：273-278.

[41] Inukai Y, Sakamoto T, Ueguchi-Tanaka M, et al. Crown rootless1, which is essential for crown root formation in rice, is a target of an AUXIN RESPONSE FACTOR in auxin signaling [J]. Plant Cell, 2005, 17（5）: 1387-1396.

[42] Liu H J, Wang S F, Yu X B, et al. ARL1, a LOB-domain protein required for adventitious root formation in rice [J]. Plant J, 2005, 43（1）: 47-56.

[43] Qi Y H, Wang S K, Shen C J, et al. OsARF12, a transcription activator on auxin response gene, regulates root elongation and affects iron accumulation in rice [J]. New Phytol, 2012, 193（1）: 109-120.

[44] Shen C J, Wang S K, Zhang S N, et al. OsARF16, a transcription factor, is required for auxin and phosphate starvation response in rice （Oryza sativa L.） [J]. Plant Cell Environ, 2013, 36: 607-620.

[45] Kitomi Y, Hidemi K, Inukai Y. Molecular mechanism of crown root initiation and the different mechanisms between crown root and radicle in rice [J]. Plant Signal Behav, 2011, 6（9）: 1276-1278.

[46] Zhao Y, Hu Y F, Dai M Q, et al. The WUSCHEL-related homeobox gene WOX11 is required to activate shoot-borne crown root development in rice [J]. Plant Cell, 2009, 21（3）: 736-748.

[47] Liu W, Xu Z H, Luo D, et al. Roles of OsCKI1, a rice casein kinase I, in root development and plant hormone sensitivity [J]. Plant J, 2003, 36（2）: 189-202.

[48] Nakamura A, Fujioka S, Sunohara H, et al. The role of OsBRI1 andits homologous genes, OsBRL1 and OsBRL3 in rice [J]. Plant Physiol, 2006, 140: 580-590.

[49] Nakamura A, Umemura I, Gomi K, et al. Production and characterization of auxininsensitive rice by overexpression of a mutagenized rice IAA protein [J]. Plant J, 2006, 46: 297-306.

[50] Zhao Y, Hu Y F, Dai M Q, et al. Modulation of ethylene responses by OsRTH1 overexpression reveals the biological significance of ethylene in rice seedling growth and development [J]. J Exp Bot, 2012, 63（11）: 4151-4164.

[51] Koltai H. Strigolactones are regulators of root development [J]. New Phytol, 2011, 190（3）: 545-549.

[52] Lofke C, Zwiewka M, Heilmannd I, et al. Asymmetric gibberellin signaling regulates vacuolar trafficking of PIN auxin transporters during root gravitropism [J]. PNS, 2013, 110（9）: 3627-3632 .

[53] Cho S H, Yoo S C, Zhang H, et al. The rice narrow leaf2 and narrow leaf3 loci encode WUSCHEL-related homeobox 3A （OsWOX3A） and function in leaf, spikelet, tiller and lateral root development [J]. New Phytol, 2013, 198（4）: 1071-1084.

[54] Song Y L, You J, Xiong L Z. Characterization of OsIAA1 gene, a member of rice Aux/IAA family involved in auxin and brassinosteroid hormone responses and plant morphogenesis [J]. Plant Mol Biol, 2009, 70（3）: 297-309.

[55] Zhang J, Peng Y L, Guo Z J. Constitutive expression of pathogen-inducible OsWRKY31 enhances disease resistance and affects root growth and auxin response in transgenic rice plants [J]. Cell Res, 2008, 18（4）: 508-521.