拟南芥根毛发育相关基因SRH1的功能和调控研究
摘要
通过激光辅助显微捕获(LCM)配合Affymetrix基因芯片技术,分析根毛细胞分化和发育相关的H细胞、N细胞和皮层细胞之间的差异表达基因并筛选突变体,获取了呈短根毛性状的T-DNA插入突变体srh1(Short Root Hair 1)。srh1共有4个等位突变体,具有相同的表型。SRH1蛋白定位于细胞膜,其蛋白结构分析结果显示,SRH1属于Beach蛋白类基因,此类基因在植物中的功能目前尚无报道。SRH1是Beach蛋白参与植物根毛发育调控的首例报道,也是高等植物Beach蛋白功能的首次发现。
在低磷胁迫和外源激素处理下,srh1具有与野生型拟南芥相似的表型变化,但根毛伸长能力受限。在srh1根毛中可观察到微丝骨架与顶端分离的异常情况。通过基因芯片和定量PCR研究野生型和srh1背景下部分已知根毛发育相关基因的表达规律,发现SRH1功能缺失仅影响少量基因的表达。但拟南芥NADPH氧化酶基因RHD2在srh1突变体中诱导表达,揭示SRH1功能缺失可能导致细胞氧化还原反应状态提高。为进一步了解SRH1、RHD2和另一与RHD2相关的基因SCN1之间的关系,设计了srh1与根毛突变体rhd2和scn1的杂交试验,构建双突变体与三突变体。以此为基础,本文探讨了SRH1基因影响根毛发育的作用机理,提出SRH1可能与根毛尖端生长阶段的囊泡运输或胞壁合成过程有关。
Based on candidate genes picked from LCM (Laser Capture Microdissection)-microarray(Affymetrix GENECHIP system) analysis for H cell, N cell and cortex, a T-DNA instertional Arabidopsis mutant with defect in root hair elogation was isolateded and named as srh1 (Short Root Hair 1). In response to phosphate starvation or addition of external hormones, srhl is similar with wide type(WT) except its weakness in root hair elongation. Microarray and q-RT-PCR analysis revealed that a function-loss results in a few genes to shift in expression level. Our result show that SRH1 was located in plasma membrane, and that a microfiber disorder was detected in root hair tip. Expression level of RHD2 encoding NADPH oxydase for root hair initiation was upregulated in srh1 mutant. Then the double mutants srh1/rhd2, srh1/scn1, the triple mutant srh1/rhd2/scn1 were developed for genetic and phenotyping analysis. The results suggest that SRH1 is involved vesicle trafficking or cell wall construction.
Gene cloning and bioinformatics analysis indicate that SRH1 is a member of Beach protein. It's the first report that Beach protein affected in root hair development, and also the first gene with a known function in plant Beach family.
在低磷胁迫和外源激素处理下,srh1具有与野生型拟南芥相似的表型变化,但根毛伸长能力受限。在srh1根毛中可观察到微丝骨架与顶端分离的异常情况。通过基因芯片和定量PCR研究野生型和srh1背景下部分已知根毛发育相关基因的表达规律,发现SRH1功能缺失仅影响少量基因的表达。但拟南芥NADPH氧化酶基因RHD2在srh1突变体中诱导表达,揭示SRH1功能缺失可能导致细胞氧化还原反应状态提高。为进一步了解SRH1、RHD2和另一与RHD2相关的基因SCN1之间的关系,设计了srh1与根毛突变体rhd2和scn1的杂交试验,构建双突变体与三突变体。以此为基础,本文探讨了SRH1基因影响根毛发育的作用机理,提出SRH1可能与根毛尖端生长阶段的囊泡运输或胞壁合成过程有关。
Based on candidate genes picked from LCM (Laser Capture Microdissection)-microarray(Affymetrix GENECHIP system) analysis for H cell, N cell and cortex, a T-DNA instertional Arabidopsis mutant with defect in root hair elogation was isolateded and named as srh1 (Short Root Hair 1). In response to phosphate starvation or addition of external hormones, srhl is similar with wide type(WT) except its weakness in root hair elongation. Microarray and q-RT-PCR analysis revealed that a function-loss results in a few genes to shift in expression level. Our result show that SRH1 was located in plasma membrane, and that a microfiber disorder was detected in root hair tip. Expression level of RHD2 encoding NADPH oxydase for root hair initiation was upregulated in srh1 mutant. Then the double mutants srh1/rhd2, srh1/scn1, the triple mutant srh1/rhd2/scn1 were developed for genetic and phenotyping analysis. The results suggest that SRH1 is involved vesicle trafficking or cell wall construction.
Gene cloning and bioinformatics analysis indicate that SRH1 is a member of Beach protein. It's the first report that Beach protein affected in root hair development, and also the first gene with a known function in plant Beach family.
引文
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Bates, T.R. and Lynch, J.P. (1996) Stimulation of root hair elongation in Arabidopsis thaliana by low phosphorus availability. Plant Cell Environ, 19,529-538.
Baumberger, N., Ringli, C. and Keller, B. (2001) The chimeric leucine-rich repeat/extensin cell wall protein LRX1 is required for root hair morphogenesis in Arabidopsis thaliana. Genes Dev, 15, 1128-1139.
Berger, E, Haseloff, J., Schiefelbein, J. and Dolan, L. (1998) Positional information in root epidermis is defined during embryogenesis and acts in domains with strict boundaries. Curr Biol, 8, 421-430,
Bernhardt, C., Lee, M.M., Gonzalez, A., Zhang, E, Lloyd, A. and Schiefelbein, J. (2003) The bHLH genes GLABRA3 (GL3) and ENHANCER OF GLABRA3 (EGL3) specify epidermal cell fate in the Arabidopsis root. Development, 130, 6431-6439.
Bernhardt, C., Zhao, M., Gonzalez, A., Lloyd, A. and Schiefelbein, J. (2005) The bHLH genes GL3 and EGL3 participate in an intercellular regulatory circuit that controls cell patterning in the Arabidopsis root epidermis. Development, 132, 291-298.
Bibikova, T.N., Zhigilei, A. and Gilroy, S. (1997) Root hair growth in Arabidopsis thaliana is directed by calcium and an endogenous polarity. Planta, 203, 495-505.
Bibikova, T.N., Jacob, T., Dahse, I. and Gilroy, S. (1998) Localized changes in apoplastic and cytoplasmic pH are associated with root hair development in Arabidopsis thaliana. Development, 125, 2925-2934.
Bibikova, T.N., Blancaflor, E.B. and Gilroy, S. (1999) Microtubules regulate tip growth and orientation in root hairs of Arabidopsis thaliana. Plant J, 17, 657-665.
Carol, R.J., Takeda, S., Linstead, P., Durrant, M.C., Kakesova, H., Derbyshire, P., Drea, S., Zarsky, V. and Dolan, L. (2005) A RhoGDP dissociation inhibitor spatially regulates growth in root hair cells. Nature, 438, 1013-1016.
Chant, J. (1999) Cell polarity in yeast. Annu Rev Cell Dev Biol, 15, 365-391.
Daniele Werck-Reichhart, S.B.a.S.P. (2002) The Arabidopsis Book, Cytochromes P450. American Society of Plant Biologists.
De Lozanne, A. (2003) The role of BEACH proteins in Dictyostelium. Traffic, 4, 6-12.
Dolan, L., Duckett, C., Grierson, C., Linstead, P., Schneider, K., Lawson, E., Dean, C., Poethig, S. and Roberts, K. (1994) Clonal relationships and cell patterning in the root epidermis of Arabidopsis. Development, 120, 2465-2474.
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Favery, B., Ryan, E., Foreman, J., Linstead, P., Boudonck, K., Steer, M., Shaw, P. and Dolan, L. (2001) KOJAK encodes a cellulose synthase-like protein required for root hair cell morphogenesis in Arabidopsis. Genes Dev, 15, 79-89.
Foreman, J., Demidchik, V., Bothwell, J.H., Mylona, P., Miedema, H., Torres, M.A., Linstead, P., Costa, S., Brownlee, C., Jones, J.D., Davies, J.M. and Dolan, L. (2003) Reactive oxygen species produced by NADPH oxidase regulate plant cell growth. Nature, 422, 442-446.
Forman, H.J. and Torres, M. (2002) Reactive oxygen species and cell signaling: respiratory burst in macrophage signaling. Am J Respir Crit Care Med, 166, S4-8.
Galway, M.E., Heckman, J.W., Jr. and Schiefelbein, J.W. (1997) Growth and ultrastructure of Arabidopsis root hairs: the rhd3 mutation alters vacuole enlargement and tip growth. Planta, 201, 209-218.
Grierson, C.S., Roberts, K., Feldmann, K.A. and Dolan, L. (1997) The C0W1 locus of arabidopsis acts after RHD2, and in parallel with RHD3 and TIP1, to determine the shape, rate of elongation, and number of root hairs produced from each site of hair formation. Plant Physiol, 115, 981-990.
Jogl, G., Shen, Y., Gebauer, D., Li, J., Wiegmann, K., Kashkar, H., Kronke, M. and Tong, L. (2002) Crystal structure of the BEACH domain reveals an unusual fold and extensive association with a novel PH domain. Embo J, 21, 4785-4795.
Jones, M.A., Raymond, M.J. and Smirnoff, N. (2006) Analysis of the root-hair morphogenesis transcriptome reveals the molecular identity of six genes with roles in root-hair development in Arabidopsis. Plant J, 45, 83-100.
Jones, M.A., Raymond, M.J., Yang, Z. and Smirnoff, N. (2007) NADPH oxidase-dependent reactive oxygen species formation required for root hair growth depends on ROP GTPase. J Exp Bot.
Kirik, V., Simon, M., Wester, K., Schiefelbein, J. and Hulskamp, M. (2004) ENHANCER of TRY and CPC 2 (ETC2) reveals redundancy in the region-specific control of trichome development of Arabidopsis. Plant Mol Biol, 55, 389-398.
Knox, K., Grierson, C.S. and Leyser, O. (2003) AXR3 and SHY2 interact to regulate root hair development. Development, 130, 5769-5777.
Kwak, J.M., Mori, I.C, Pei, Z.M., Leonhardt, N., Torres, M.A., Dangl, J.L., Bloom, R.E., Bodde, S., Jones, J.D. and Schroeder, J.I. (2003) NADPH oxidase AtrbohD and AtrbohF genes function in ROS-dependent ABA signaling in Arabidopsis. Embo J, 22, 2623-2633.
Kwak, S.H., Shen, R. and Schiefelbein, J. (2005) Positional signaling mediated by a receptor-like kinase in Arabidopsis. Science, 307,1111-1113.
Lee, M.M. and Schiefelbein, J. (1999) WEREWOLF, a MYB-related protein in Arabidopsis, is a position-dependent regulator of epidermal cell patterning. Cell, 99, 473-483.
Lew, R.R. (1996) Pressure regulation of the electrical properties of growing Arabidopsis thaliana L. root hairs. Plant Physiol, 112, 1089-1100.
Lew, R.R. (2004) Osmotic effects on the electrical properties of Arabidopsis root hair vacuoles in situ. Plant Physiol, 134, 352-360.
Ma, Z., Bielenberg, D.G., Brown, K.M. and Lynch, J.P. (2001) Regulation of root hair density by phosphorus availability in Arabidopsis thaliana. Plant Cell Environ, 24, 459-467.
Masucci, J.D. and Schiefelbein, J.W. (1994) The rhd6 Mutation of Arabidopsis thaliana Alters Root-Hair Initiation through an Auxin- and Ethylene-Associated Process. Plant Physiol, 106, 1335-1346.
Masucci, J.D., Rerie, W.G., Foreman, D.R., Zhang, M., Galway, M.E., Marks, M.D. and Schiefelbein, J.W. (1996) The homeobox gene GLABRA2 is required for position-dependent cell differentiation in the root epidermis of Arabidopsis thaliana. Development, 122,1253-1260.
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