苹果BR信号转录因子基因MdBZR1的克隆及表达分析
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摘要
从矮化苹果砧木Malling 9(M9)中分离1个BR信号转录因子BRASSINAZOLE-RESISTANT1(BZR1)基因,命名为MdBZR1,其开放阅读框为888 bp,编码295个氨基酸,有5个保守结构域:核定位信号结构域NLS(MARRKPSWRERENNRRTERRR)、氮(N)端结构域(NLPKHCDNNEVLKALCLQ AGWTVEDDGT)、BRASSINOSTEROID-INSENSITIVE 2(BIN2)磷酸化结构域(STKISPYSSLNPSPIPS YOVSPSSSSYPSPTR)、PEST结构域(PTAATIPECDESDASTVDSGQ)和碳(C)端保守结构域(VKPWIGEK IHEVGLDDLELTLGNGKA)。系统进化树分析显示,MdBZR1与美国National Center for Biotechnology Information(NCBI)数据库中注释的苹果BZR1基因亲缘关系最近。MdBZR1在M9不同组织均有表达,在顶梢中的表达水平最高;外源BR和生长素处理促进苹果幼苗(M9和平邑甜茶)株高增长,MdBZR1表达量增加;GA处理虽也促进株高增长,但对MdBZR1的表达影响不显著。此外,半矮化砧穗组合长富2号/MM106和乔化砧穗组合长富2号/长富2号接穗中MdBZR1的表达量明显高于矮化砧穗组合长富2号/M9。因此,MdBZR1很可能对苹果树株高具有重要调控作用。
In order to study the regulation function of MdBZR1 on apple plant height,a BR signal transcription factor gene BRASSINAZOLE-RESISTANT 1(BZR1)was cloned from apple dwarf rootstock Malling 9(M9),which was named as MdBZR1. MdBZR1 contains an open reading frame(888 bp)and encodes a protein of 295 amino acids. Bioinformatics analysis showed that MdBZR1 shared five conserved domains:nuclear localization signal(NLS)(MARRKPSWRERENNRRTERRR),nitrogen terminal domain(N)(NLPKHCDNNEVLKALCLQAGWTVEDDGT),BIN2 phosphorylation domain(STKISPYSSLNPSPIPSYOVSPSSSSYPSPTR),PEST motif(PTAATIPECDESDASTVDSGQ)and a carboxylterminal(C)(VKPWIGEKIHEVGLDDLELTLGNGKA). A phylogenetic analysis of BZR1 genes indicated that MdBZR1 shared high homologous amino acid sequences with apple BZR1 gene annotated in National Center for Biotechnology Information(NCBI)database. The expression pattern of MdBZR1 gene in different tissues of M9 was characterized. Results indicated that MdBZR1 transcript was detected in shoot tip,leaf,root,xylem and phloem of main stem,and the highest level appeared in shoot tip. Exogenous brassinosteroid(BR)and auxin increased primary shoot length of M9 as well as Pingyitiancha(Malus hupehensis)and significantly induced the expression of MdBZR1. Primary shoot length was also increased by GA treatment,while MdBZR1 was neither induced nor repressed by GA obviously. In addition, MdBZR1 exhibited higher expression level in Nagafu/MM106 and Nagafu/Nagafu than Nagafu/M9. These results suggest that MdBZR1 may be involved in the regulation of apple tree size.
In order to study the regulation function of MdBZR1 on apple plant height,a BR signal transcription factor gene BRASSINAZOLE-RESISTANT 1(BZR1)was cloned from apple dwarf rootstock Malling 9(M9),which was named as MdBZR1. MdBZR1 contains an open reading frame(888 bp)and encodes a protein of 295 amino acids. Bioinformatics analysis showed that MdBZR1 shared five conserved domains:nuclear localization signal(NLS)(MARRKPSWRERENNRRTERRR),nitrogen terminal domain(N)(NLPKHCDNNEVLKALCLQAGWTVEDDGT),BIN2 phosphorylation domain(STKISPYSSLNPSPIPSYOVSPSSSSYPSPTR),PEST motif(PTAATIPECDESDASTVDSGQ)and a carboxylterminal(C)(VKPWIGEKIHEVGLDDLELTLGNGKA). A phylogenetic analysis of BZR1 genes indicated that MdBZR1 shared high homologous amino acid sequences with apple BZR1 gene annotated in National Center for Biotechnology Information(NCBI)database. The expression pattern of MdBZR1 gene in different tissues of M9 was characterized. Results indicated that MdBZR1 transcript was detected in shoot tip,leaf,root,xylem and phloem of main stem,and the highest level appeared in shoot tip. Exogenous brassinosteroid(BR)and auxin increased primary shoot length of M9 as well as Pingyitiancha(Malus hupehensis)and significantly induced the expression of MdBZR1. Primary shoot length was also increased by GA treatment,while MdBZR1 was neither induced nor repressed by GA obviously. In addition, MdBZR1 exhibited higher expression level in Nagafu/MM106 and Nagafu/Nagafu than Nagafu/M9. These results suggest that MdBZR1 may be involved in the regulation of apple tree size.
引文
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Du Li-sha,Qi Si-yan,Ma Juan-juan,Xing Li-bo,Fan Sheng,Li You-mei,Zhang Dong,Zhao Cai-ping,Han Ming-yu.2017.Effects of de-fruit treatment on soluble sugar accumulation and expression of flowering-related genes in terminal spur buds and leaves of‘Fuji’apple.Acta Horticulturae Sinica,44(4):622-632.(in Chinese)杜利莎,齐思言,马娟娟,邢利博,樊胜,李有梅,张东,赵彩平,韩明玉.2017.‘长富2号’苹果短枝幼果摘除对芽叶可溶性糖积累和成花相关基因表达的影响.园艺学报,44(4):622-632.
Gong Ming-xia,Luo Hai-ling,Yuan Hong-juan,Wei Shan-qing,Yang Xu-dong,He Long-fei.2015.Effects of exogenous gibberellin and paclobutrazol on tuber expansion and bulbil formation of Chinese yam.Acta Horticulturae Sinica,42(6):1175-1184.(in Chinese)龚明霞,罗海玲,袁红娟,韦善清,杨旭东,何龙飞.2015.外源赤霉素和多效唑对山药块茎膨大和零余子形成的影响.园艺学报,42(6):1175-1184.
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Han Ming-yu.2013.Apple intensive pattern with dwarf rootstock research progress and the next task.Northwest Horticulture(fruit tree),(3):4-6.(in Chinese)韩明玉.2013.苹果矮砧集约栽培模式研究进展与下一步任务.西北园艺(果树),(3):4-6.
Jaakola L,Pirttila A M,Halonen M,Hohtola A.2001.Isolation of high quality RNA from bilberry(Vaccinium myrtillus L.)fruit.Molecular Biotechnology,19:201-203.
Kir G,Ye H,Nelissen H,Neelakandan A K,Kusnandar A S,Luo A,Inze D,Sylvester A W,Yin Y,Becraft P W.2015.RNA interference knockdown of brassinosteroid insensitive1 in maize reveals novel functions for brassinosteroid signaling in controlling plant architecture.Plant Physiol,169(1):826-829.
Li J,Hou H,Li X,Xiang J,Yin X,Gao H,Zheng Y,Bassett C L,Wang X.2013.Genome-wide identification and analysis of the SBP-box family genes in apple(Malus×domestica Borkh.).Plant Physiol Bioch,70:100-114.
Li J M,Chory J.1999.Brassinosteroid actions in plants.J Exp Bot,50(332):275-282.
Li L,Deng X W.2005.It runs in the family:regulation of brassinosteroid signaling by the BZR1-BES1 class of transcription factors.Trends Plant Sci,10(6):266-268.
Li Ying-hui,Li Yan,Hang Xiao-ming,Wang Peng.2001.Effects of cytokinins on shoot organogenesis from cotyledon of pepper.Acta Horticulturae Sinica,28(3):270-272.(in Chinese)李英慧,李艳,杭晓明,王朋.2001.细胞分裂素对辣椒子叶再生的影响.园艺学报,28(3):270-272.
Li Zhen,Wei Jia,Jia Cheng-guo,Wang Qiao-mei.2010.Effect of Bzr1 gene transformation on characters of fruit of tomato.Scientia Agricultura Sinica,(9):1868-1876.(in Chinese)李振,魏佳,贾承国,汪俏梅.2010.Bzr1基因的转化对番茄果实性状的影响.中国农业科学,(9):1868-1876.
Liu Feng-luan,Kou Ya-ping,Chen Xiao-li,Gao Bin,Wang Ling,Zhao Liang-jun.2014.Cloning and expression analysis of PIN1 and PIN2 encoding auxin efflux carriers in Rosa canina.Acta Horticulturae Sinica,41(5):925-934.(in Chinese)刘凤栾,寇亚平,陈晓丽,高彬,王玲,赵梁军.2014.狗蔷薇生长素输出载体蛋白基因PIN1和PIN2的分离与表达分析.园艺学报,41(5):925-934.
Liu L,Liu H,Li S,Zhang X,Zhang M,Zhu N,Dufresne C P,Chen S,Wang Q.2016.Regulation of BZR1 in fruit ripening revealed by i TRAQproteomics analysis.Sci Rep,6:33-36.
Liu Xiao-jie,Fan Sheng,Li Guo-fang,Tan Ming,Mo Ning,Ma Juan-juan,Zhang Dong,Han Ming-yu.2017.Genome-wide identification of PINgene family,cloning and expression analysis of Md PIN15 during axillary bud burst in Malus.Acta Horticulturae Sinica,44(11):2041-2054.(in Chinese)刘小杰,樊胜,李国防,檀鸣,默宁,马娟娟,张东,韩明玉.2017.苹果全基因组PIN成员鉴定及Md PIN15的克隆和在腋芽萌发中的表达分析.园艺学报,44(11):2041-2054.
Livak K J,Schmittgen T D.2001.Analysis of relative gene expression data using real-time quantitative PCR and the 2-ΔΔCt.Method,25(4):402-408.
Ma J J,Han M.2016.Genomewide analysis of ABCBs with a focus on ABCB1 and ABCB19 in Malus domestica.J Genet,95(1):141-149.
Ma Y,Xue H,Zhang L,Zhang F,Ou C,Wang F,Zhang Z.2016.Involvement of auxin and Brassinosteroid in Dwarfism of autotetraploid apple(Malus×domestica).Sci Rep,6:26-39.
Mao Jiangping,Zhang Dong,Li Ke,Liu Zhen,Liu Xiaojie,Song Chunhui,Li Guofang,Zhao Caiping,Ma Juanjuan,Han Mingyu.2017.Effect of exogenous Brassinolide(BR)application on the morphology,hormone status,and gene expression of developing lateral roots in Malus hupehensis.Plant Growth Regulation,26:4-9.
Miyaji T,Yamagami A,Kume N,Sakuta M,Osada H,Asami T,Arimoto Y,Nakano T.2014.Brassinosteroid-related transcription factor BIL1/BZR1increases plant resistance to insect feeding.Biosci Biotech Bioch,78(6):960-968.
Mouchel C F,Osmont K S,Hardtke C S.2006.BRX mediates feedback between brassinosteroid levels and auxin signalling in root growth.Nature,443(7110):458-461.
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