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作物学报 ›› 2018, Vol. 44 ›› Issue (7): 1021-1031.doi: 10.3724/SP.J.1006.2018.01021

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

马铃薯块茎花色素苷合成相关R2R3 MYB蛋白基因的克隆和功能
分析

谈欢1,刘玉汇1,*(),李丽霞1,王丽2,李元铭1,张俊莲1,*()   

  1. 1 甘肃省作物遗传改良与种质创新重点实验室 / 甘肃农业大学园艺学院 / 甘肃省干旱生境作物学重点实验室, 甘肃兰州 730070
    2 甘肃农业大学生命科学技术学院, 甘肃兰州 730070
  • 收稿日期:2017-10-27 接受日期:2018-03-26 出版日期:2018-07-10 网络出版日期:2018-04-09
  • 通讯作者: 刘玉汇,张俊莲
  • 基金资助:
    本研究由国家自然科学基金项目(31601356), 甘肃省杰出青年科学基金项目(17JR5RA138), 甘肃省农业生物技术研究与应用开发专项(GNSW-2015-15), 甘肃省高等学校科研项目(2016A-030)和甘肃农业大学“伏羲人才”计划项目(Gaufx-02Y04)资助

Cloning and Functional Analysis of R2R3 MYB Genes Involved in Anthocyanin Biosynthesis in Potato Tuber

Huan TAN1,Yu-Hui LIU1,*(),Li-Xia LI1,Li WANG2,Yuan-Ming LI1,Jun-Lian ZHANG1,*()   

  1. 1 Gansu Key Laboratory of Crop Genetic Improvement and Germplasm Enhancement / College of Horticulture / Gansu Provincial Key Laboratory of Aridland Crop Science, Gansu Agricultural University, Lanzhou 730070, Gansu, China
    2 College of Life Science and Technology, Gansu Agricultural University, Lanzhou 730070, Gansu, China
  • Received:2017-10-27 Accepted:2018-03-26 Published:2018-07-10 Published online:2018-04-09
  • Contact: Yu-Hui LIU,Jun-Lian ZHANG
  • Supported by:
    This study was supported by the National Natural Science Foundation of China (31601356), Gansu Science Foundation for Distinguished Young Scholars (17JR5RA138), the Agricultural Biotechnology Research and Application Development Foundation of Gansu Province (GNSW-2015-15), Gansu Scientific Research Foundation for the Universities (2016A-030), and Fuxi Talent Project of Gansu Agricultural University (Gaufx-02Y04).

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摘要：

植物花色素苷的合成代谢受转录因子的调控, 其中R2R3 MYB为最主要的转录调控因子。本研究以彩色四倍体马铃薯为试材, 克隆了R2R3 MYB基因家族里调控马铃薯块茎花色素苷合成R2R3 MYB-StAN1的3个同源基因, 并利用生物信息学分析、稳定烟草遗传转化、qPCR等方法对这3个同源基因的结构和功能进行分析和鉴定, 结果表明, 这3个同源基因均含有R2和R3保守结构域, 其主要差别在于C端由10个氨基酸序列组成的重复结构(R)数目不同, 根据R数目将其分别命名为StAN1-R0、StAN1-R1和StAN1-R3, 其蛋白分子量分别为28 047.91、29 458.35和31 527.60 Da, 等电点(pI)分别为6.14、6.90和8.39, 均为亲水蛋白。通过转化烟草发现, 转入StAN1-R0、StAN1-R1和StAN1-R3后, 烟草叶片叶色变化明显, 其中转StAN1-R1烟草叶色呈深红色, 其叶片花色素苷含量最高。进一步利用qPCR分析表明, 外源StAN1使烟草叶片花色素苷合成代谢途径的关键基因(NtCHS、NtCHI、NtF3H、NtF3’H、NtDFR、NtANS、NtUFGT)上调表达, 同时烟草内源NtbHLH基因的表达显著上升; StAN1-R1可以高效地调控烟草内源NtbHLH基因和结构基因NtDFR和NtANS的表达。结果表明, StAN1的3个同源蛋白均可以调控花色素苷的合成, 而只含一个重复序列R的StAN1调控花色素苷合成的能力最强。

关键词: 马铃薯, 花色素苷, R2R3 MYB, 基因克隆, 生物信息学分析, 烟草转化, 基因表达分析

Abstract:

The anthocyanins biosynthesis is regulated by transcription factors, and R2R3 MYB is the most important transcriptional regulator in plant. In this study, three homologous genes were isolated from tetraploid potato, belonging to R2R3 MYB gene family. The structure and function analysis of the three homologous genes were characterized by bioinformatics analysis, stable tobacco genetic transformation and qPCR assays. The three homologous genes contained R2 and R3 conserved domains, differed in the number of repeats (R) consisting of 10 amino acid sequences, and named as StAN1-R0, StAN1-R1, and StAN1-R3 according to the number of R. The coding proteins are hydrophilic with molecular weight of 28 047.91, 29 458.35, 31 527.60 Da, and isoelectric points (pI) of 6.14, 6.90, and 8.39, respectively. The accumulation of anthocyanin was significantly increased in StAN1-R0, StAN1-R1 and StAN1-R3 overexpressed plants. The leaf color of StAN1-R1-overexpressed plants was dark red with the highest anthocyanin content among the three transgenic events. The qPCR assays showed that exogenous StAN1 genes enhanced the expression of endogenous NtbHLH transcription factor as well as NtCHS, NtCHI, NtF3H, NtF3’H, NtDFR, NtANS, and NtUFGT, involved in anthocyanin biosynthesis in transgenic tobacco leaves. The overexpression of StAN1-R1 resulted in higher expression of NtDFR, NtANS, and endogenous NtbHLH in transgenic tobacco. The results showed that three homologous genes of StAN1 can regulate anthocyanin biosynthesis, among them StAN1-R1 containing one R has the strongest regulatory capacity.

Key words: potato, anthocyanin, R2R3 MYB, gene cloning, bioinformatics analysis, stable tobacco transformation, gene expression analysis

表1

定量PCR分析的引物序列"

基因
Gene		 登录号
Accession number		 上游引物
Forward primer (5'-3')		 下游引物
Reverse primer (5'-3')
StAN1 AY841129.1 GGCCACATATCAAGAGAGGTGACTTTG TCACATCGTTAGCTGTCCTTCCTGG
NtCHS AF311783.1 TTGTTCGAGCTTGTCTCTGC AGCCCAGGAACATCTTTGAG
NtCHI AB213651.1 GTCAGGCCATTGAAAAGCTC CTAATCGTCAATGCCCCAAC
NtF3H AB289450.1 CAAGGCATGTGTGGATATGG TGTGTCGTTTCAGTCCAAGG
NtF3'H AB289449.1 AGGCTCAACACTTCTCGT CATCAACTTTGGGCTTCT
NtDFR EF421429.1 AACCAACAGTCAGGGGAATG TTGGACATCGACAGTTCCAG
NtANS AB289447.1 TGGCGTTGAAGCTCATACTG GGAATTAGGCACACACTTTGC
NtUFGT FG627024.1 GAGTGCATTGGATGCCTTTT CCAGCTCCATTAGGTCCTTG
NtbHLH HQ589208.1/HQ589209.1 ATGKGCGCAAACGAGGTTGATAGC TRGCTGAGGTTGTTGTTGCTCA
NtEF-1α D63396.1 TGAACCATCCAKGACAGATTGG TGGGCTCCTTCTCAATCTCCTT

图1

总RNA的电泳图RS1:“甘农薯5号”红色薯皮; WS:“新大坪”白色薯皮; PS:“黑美人”紫色薯皮; RS2:“青薯9号”红色薯皮; WF1:“甘农薯5号”白色薯肉; WF:“新大坪”白色薯肉; PF:“黑美人”紫色薯肉; WF2:“青薯9号”白色薯肉; RFV:“青薯9号”红色维管束; M: DNA marker。"

图2

基因克隆过程的PCR产物电泳图RS1:“甘农薯5号”红色薯皮; WS:“新大坪”白色薯皮; PS:“黑美人”紫色薯皮; RS2:“青薯9号”红色薯皮; WF1:“甘农薯5号”白色薯肉; WF:“新大坪”白色薯肉; PF:“黑美人”紫色薯肉; WF2:“青薯9号”白色薯肉; RFV:“青薯9号”红色维管束; M: DNA marker。"

图3

序列比对图 R表示重复结构TIAPQPQEGI。"

表2

马铃薯StAN1基因编码蛋白质的一级结构预测"

基因
Gene		 蛋白质分子式
Formula		 分子量
Molecular weight (Da)		 等电点
pI		 不稳定系数
Instability index		 平均疏水性
Average of hydropathicity		 脂肪系数
Aliphatic index
StAN1-R0 C1227H1936N362O365S14 28047.91 8.39 46.97 -0.693 79.14
StAN1-R1 C1284H2019N377O390S15 29458.35 6.90 48.64 -0.720 74.88
StAN1-R3 C1371H2164N402O423S15 31527.60 6.14 50.76 -0.712 76.15

表3

StAN1基因编码蛋白质的二级结构分析"

基因
Gene		 α-螺旋
Alpha helix		 β-螺旋
Beta turn		 无规则卷曲
Random coil		 延伸链
Extended strand
StAN1-R0 34.84 9.84 40.57 14.75
StAN1-R1 35.27 8.91 42.25 13.57
StAN1-R3 28.06 7.55 47.84 16.55

图4

一些茄科植物与其他科属植物的R2R3 MYB氨基酸序列多重比对StAN1-R0_PSAK277, StAN1-R1_PSAK277, StAN1-R3_PSAK277: 马铃薯; AAF66727: 矮牵牛; AAA33482.1: 玉米; AAG42001.1-AAG42002.1: 拟南芥; ABB84753.1: 苹果; ABD72956.1: 葡萄; ABO26065.1: 智利番茄; ABX71488.1: 沙梨; ACI195261.1: 多毛番茄; ACN79541.1: 蒺藜苜蓿; ACO52470.1: 烟草; ADW94950.1: 矮牵牛; AFD31843.1: 马铃薯; AIP93873.1: 茄子; AMK01804.1: 茄子; BAF45114.1: 甘薯; NP_001311547.1: 辣椒; ABX71493.1: 甜樱桃; sp_O23892.1: 水稻。"

图5

StAN1基因与一些茄科和其他植物R2R3 MYB转录因子家族氨基酸序列的系统进化树分析"

图6

转StAN1基因烟草的表型分析a: 转StAN1基因烟草的整株; b: 转StAN1基因烟草的叶片。"

图7

转StAN1基因烟草的PCR鉴定M: DNA marker; CK: 未转基因烟草; R0、R1、R3分别为转StAN1-R0、StAN1-R1、StAN1-R3基因烟草。"

图8

转StAN1基因烟草与对照的花色素苷平均含量采用Duncan’s多重比较方法分析(P < 0.05, n = 3)。"

图9

转基因烟草中与花色素苷合成相关基因相对表达量采用Duncan’s多重比较方法分析(P < 0.05, n = 3)。"

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