水稻OsRAA1基因的克隆与功能分析
摘要
FPF1(flowering promoting factor1)蛋白最早在白芥中研究发现是开花促进因子，可能参与了赤霉素的信号传导途径。它可以和AP1和LFY蛋白协同作用，促进茎顶端分生组织向花分生组织转化。本论文将AtFPF1基因转化水稻，转基因水稻抽穗时间只有微弱的提前。然而异源表达AtFPF1却抑制了转基因水稻根的生长，促进了成苗根系的发达。这些表型类似于我们克隆的水稻OsRAA1 (Oryza sativa Root Architecture Associated 1)的功能。这是首次报道AtFPF1／OsRAA1在水稻中具有控制根系发育的功能
生物信息学分析表明水稻OsRAA1基因定位于水稻1号染色体，它编码的蛋白推测分子量为12kD和AtFPF1有58％的同源。通过RNA原位杂交和OsRAA1基因启动子调控GUS基因表达的模式，证实OsRAA1基因主要在根尖的顶端分生区和伸长区，根尖分支区和幼侧根的中柱，侧根的原基表达。同时在幼穗分支顶端，根茎结合区的边周维管束，稃片，花药与花丝的结合区也有表达。OsRAA1在玉米泛素启动子驱动下组成型表达，可以抑制水稻初生根的生长，促进不定根的形成，部分植物形成不同程度螺旋状的初生根。这些表型和野生型水稻用生长素处理的表型类似。OsRAA1组成型表达，在成苗阶段，特别是孕穗前，大大促进叶片伸长，并导致部分小花败育。光学镜检表明OsRAA1组成型表达的水稻的花丝伸长过快，部分小花花药萎缩败育。剑叶表面细胞电镜扫描表明，OsRAA1组成型表达的水稻剑叶的硅化细胞比对照植株要长。野生型水稻根系生长素处理的Northern杂交和OsRAA1基因启动子调控GUS基因表达的水稻生长素处理后GUS活性染色表明，OsRAA1基因的转录受生长素诱导。而且OsRAA1组成型表达的水稻根的向地性反应减缓。这些结果表明，OsRAA1可能参与了生长素的信号转导途径。
与此同时，从基因序列数据库中，在很多植物中寻找到很多表达片段和FPF1／OsRAA1基因同源。从已有报道和我们的结果表明，在植物中可能普遍存在一个受赤霉素和生长素调控FPF1／OsRAA1基因家族，调控着植物各个器官的发育。
FPF1 protein was firstly studied as a flowering promoting factor in mustard. It was indicated that FPF1 is involved in GA-dependent signaling pathway. And it may work synergistically with AP1 and LFY to modulate the competence to flowering in the shoot apical meristem. In the dissertation,AtFPF1 gene was transformed into rice. It is unfortunately that it just weakly promoted the heading of rice. On the contrary,it inhibited the root growth and promoted mature adventitious roots formation. These functions were just like those of the gene Oryza sativa Root Architecture Associated 1 (OsRAA1) which was cloned in rice. The protein shared 58% homology to the AtFPFI in Arabidopsis. It is the first report that the AtFPF1/OsRAA1 gene has a function to control root development in rice.
The gene OsRAAl was located in chromosome 1 of rice based on the data of bioinformatics. Now this gene has been characterized molecularly. The OsRAAl gene encodes a 12.0 kDa protein. The results of in situ hybridization showed that its expression was detectable clearly in the apical meristem,the elongation zone of root tip,steles of the branch zone and the young lateral root. At the same time,its transcript also appeared in the young spike and lateral root primordia. Analysis of GUS activity,which was controlled by the promoter of OsRAAl gene,suggested that OsRAAl may be also expressed in the anther and palea/low palea. Constitutive expression of the gene under control of maize ubiquitin promoter resulted in inhibiting the growth of primary root,promoting the formation of adventitious roots,forming helix primary root in rice,which is similar to the phenotype of the plant treated with auxin. Its constitutive expression also caused the longer leaves and sterile of part florets at the late stage of plant develo
pment. The data of scan electronic microscope showed that flag leafs cells of the transgenic plant has been elongated more than those of the control. The analyses of Northen blot and GUS activity,which was controlled under OsRAAl promoter,demonstrated that the transcription of OsRAAl was induced by auxin. The gravitropic response was also delayed in the transgenic
rice,which constitutively expressed OsRAAl. These data suggested that OsRAAl might be involved in the pathway of Auxin/IAA signal transduction in establishing the root architecture in rice.
At the same time,many genes or ESTs were mined from the database showed that they have high homology to FPF1/OsRAA1 gene. From others and our data,it was suggested that there may be a FPF1/OsRAAl gene family in the plant which may be regulated by GA and auxin to control many aspects of plant development.
生物信息学分析表明水稻OsRAA1基因定位于水稻1号染色体，它编码的蛋白推测分子量为12kD和AtFPF1有58％的同源。通过RNA原位杂交和OsRAA1基因启动子调控GUS基因表达的模式，证实OsRAA1基因主要在根尖的顶端分生区和伸长区，根尖分支区和幼侧根的中柱，侧根的原基表达。同时在幼穗分支顶端，根茎结合区的边周维管束，稃片，花药与花丝的结合区也有表达。OsRAA1在玉米泛素启动子驱动下组成型表达，可以抑制水稻初生根的生长，促进不定根的形成，部分植物形成不同程度螺旋状的初生根。这些表型和野生型水稻用生长素处理的表型类似。OsRAA1组成型表达，在成苗阶段，特别是孕穗前，大大促进叶片伸长，并导致部分小花败育。光学镜检表明OsRAA1组成型表达的水稻的花丝伸长过快，部分小花花药萎缩败育。剑叶表面细胞电镜扫描表明，OsRAA1组成型表达的水稻剑叶的硅化细胞比对照植株要长。野生型水稻根系生长素处理的Northern杂交和OsRAA1基因启动子调控GUS基因表达的水稻生长素处理后GUS活性染色表明，OsRAA1基因的转录受生长素诱导。而且OsRAA1组成型表达的水稻根的向地性反应减缓。这些结果表明，OsRAA1可能参与了生长素的信号转导途径。
与此同时，从基因序列数据库中，在很多植物中寻找到很多表达片段和FPF1／OsRAA1基因同源。从已有报道和我们的结果表明，在植物中可能普遍存在一个受赤霉素和生长素调控FPF1／OsRAA1基因家族，调控着植物各个器官的发育。
FPF1 protein was firstly studied as a flowering promoting factor in mustard. It was indicated that FPF1 is involved in GA-dependent signaling pathway. And it may work synergistically with AP1 and LFY to modulate the competence to flowering in the shoot apical meristem. In the dissertation,AtFPF1 gene was transformed into rice. It is unfortunately that it just weakly promoted the heading of rice. On the contrary,it inhibited the root growth and promoted mature adventitious roots formation. These functions were just like those of the gene Oryza sativa Root Architecture Associated 1 (OsRAA1) which was cloned in rice. The protein shared 58% homology to the AtFPFI in Arabidopsis. It is the first report that the AtFPF1/OsRAA1 gene has a function to control root development in rice.
The gene OsRAAl was located in chromosome 1 of rice based on the data of bioinformatics. Now this gene has been characterized molecularly. The OsRAAl gene encodes a 12.0 kDa protein. The results of in situ hybridization showed that its expression was detectable clearly in the apical meristem,the elongation zone of root tip,steles of the branch zone and the young lateral root. At the same time,its transcript also appeared in the young spike and lateral root primordia. Analysis of GUS activity,which was controlled by the promoter of OsRAAl gene,suggested that OsRAAl may be also expressed in the anther and palea/low palea. Constitutive expression of the gene under control of maize ubiquitin promoter resulted in inhibiting the growth of primary root,promoting the formation of adventitious roots,forming helix primary root in rice,which is similar to the phenotype of the plant treated with auxin. Its constitutive expression also caused the longer leaves and sterile of part florets at the late stage of plant develo
pment. The data of scan electronic microscope showed that flag leafs cells of the transgenic plant has been elongated more than those of the control. The analyses of Northen blot and GUS activity,which was controlled under OsRAAl promoter,demonstrated that the transcription of OsRAAl was induced by auxin. The gravitropic response was also delayed in the transgenic
rice,which constitutively expressed OsRAAl. These data suggested that OsRAAl might be involved in the pathway of Auxin/IAA signal transduction in establishing the root architecture in rice.
At the same time,many genes or ESTs were mined from the database showed that they have high homology to FPF1/OsRAA1 gene. From others and our data,it was suggested that there may be a FPF1/OsRAAl gene family in the plant which may be regulated by GA and auxin to control many aspects of plant development.
引文
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3．王斌，李传友，郑洪刚，伏健民，杨仁崔，Henry T．Nauyen(1999)水稻的AFLP研究初报--反应条件的优化及对温敏核不育等位突变系的分析。植物学报 41(5)：502-507
4．王风平，梅明华，徐才国，张启发．(1997)光敏核不育水稻农垦58S与正常品种“58S”在pms1区段无育性基因分离。植物学报 39(10)：922-925
5．张端品，邓训安，余功新．(1990)农垦58S光敏雄性不育基因的染色体定位。华中农业大学学报 9：407-419
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2．石明松(1985)对光照长度敏感的隐性雄性不育水稻的发现和初步分析。户国农业科学(北京)2：44-48 谭克辉(1992)春化过程中特异蛋白质的合成。植物学集刊，6：13－16
3．王斌，李传友，郑洪刚，伏健民，杨仁崔，Henry T．Nauyen(1999)水稻的AFLP研究初报--反应条件的优化及对温敏核不育等位突变系的分析。植物学报 41(5)：502-507
4．王风平，梅明华，徐才国，张启发．(1997)光敏核不育水稻农垦58S与正常品种“58S”在pms1区段无育性基因分离。植物学报 39(10)：922-925
5．张端品，邓训安，余功新．(1990)农垦58S光敏雄性不育基因的染色体定位。华中农业大学学报 9：407-419
6. Aubert D, Chevillard M, Dorne AM, Arlaud G, Herzog M. (1998) Expression patterns of GASA genes in Arabidopsis thaliana: the GASA4 gene is up-regulated by gibberellins in meristematic regions. Plant Mol Biol. 36:871-83
7. Baluska F, Parker JS, Barlow PW (1993) A role for gibberellic acid in orienting microtubules and regulating cell growth polarity in the maize root cortex. Planta 191:149-157
8. Bernier, G. (1988). The control of floral evocation and morphogenesis. Annu. Rev. Plant. Physiol. Plant Mol. Biol. 39: 175-219.
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