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作物学报 ›› 2020, Vol. 46 ›› Issue (12): 1850-1861.doi: 10.3724/SP.J.1006.2020.04004

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

甘蓝自交不亲和性相关基因BoGSTL21的克隆与表达分析

左同鸿1(), 张贺翠1, 刘倩莹1, 廉小平2, 谢琴琴1, 胡燈科1, 张以忠1, 王玉奎1, 白晓璟1, 朱利泉1,*()   

  1. 1西南大学农学与生物科技学院, 重庆 400716
    2西南大学园艺园林学院, 重庆 400716
  • 收稿日期:2020-01-08 接受日期:2020-07-02 出版日期:2020-12-12 网络出版日期:2020-09-24
  • 通讯作者: 朱利泉
  • 基金资助:
    国家自然科学基金项目(31572127);重庆市研究生科研创新项目(CYS18085);中央高校基本科研业务费(XDJK2017C032)

Molecular cloning and expression analysis of BoGSTL21 in self-incompatibility Brasscia oleracea

Tong-Hong ZUO1(), He-Cui ZHANG1, Qian-Ying LIU1, Xiao-Ping LIAN2, Qin-Qin XIE1, Deng-Ke HU1, Yi-Zhong ZHANG1, Yu-Kui WANG1, Xiao-Jing BAI1, Li-Quan ZHU1,*()   

  1. 1College of Agronomy and Biotechnology, Southwest University, Chongqing 400716, China
    2College of Horticulture and Landscape Architecture, Southwest University, Chongqing 400716, China
  • Received:2020-01-08 Accepted:2020-07-02 Published:2020-12-12 Published online:2020-09-24
  • Contact: Li-Quan ZHU
  • Supported by:
    National Natural Science Foundation of China(31572127);Chongqing Graduate Research and Innovation Project(CYS18085);Basic Scientific Research Business Expenses Project of the Central University(XDJK2017C032)

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摘要：

谷胱甘肽-S-转移酶(glutathione S-transferases, GSTs)对植物抵御逆境胁迫、解除细胞毒素和植物生长发育起着重要作用。本研究通过甘蓝自花授粉0~60 min的柱头转录组数据分析, 筛选到1个受自花授粉诱导上调表达的谷胱甘肽-S-转移酶基因BoGSTL21。BoGSTL21基因开放阅读框长度为900 bp, 编码299个氨基酸, 理论等电点为8.49, 不包含信号肽和跨膜区, 含有GST-N和GST-C结构域。BoGSTL21基因启动子中含有光响应、生长素应答、脱落酸反应、低温和干旱响应等多种顺式作用元件。BoGSTL21基因在甘蓝不同组织中均有表达, 柱头中的表达量随发育时间而变化, 在成熟的柱头中高表达。荧光定量PCR结果证实, BoGSTL21基因在0~60 min的表达量变化趋势与转录组分析结果一致。通过酵母双杂交发现, BoGSTL21蛋白与花粉发育相关蛋白BoFAB1C、生长素相关的蛋白BoPATL2、醛缩酶型TIM桶家族蛋白BoF9N12_9存在相互作用。BoGSTL21基因在大肠杆菌中被成功诱导表达, 纯化蛋白大小为34 kD, 与预测结果一致。表明BoGSTL21可能是参与SI反应过程的新蛋白, 这为甘蓝自交不亲和的进一步研究和利用提供了新内容。

关键词: 甘蓝, 基因克隆, 自交不亲和, 表达分析, 酵母双杂交

Abstract:

Glutathione-S-transferase (GSTs) plays an important role in plant resistance to stress, cytotoxic release and plant growth and development. In this study, we identified an up-regulated gene named BoGSTL21 based on the stigma transcriptome data in 0-60 min self-pollination. BoGSTL21 had an open reading frame (ORF) with the length of 900 bp, encoded a protein of 299 amino acid residues, which contained GST-N and GST-C domains without signal peptide and transmembrane domain, the theory isoelectric point of BoGSTL21 was 8.49. The promoter of BoGSTL21 gene contained many cis-acting elements such as light response, auxin response, abscisic acid response, low temperature and drought response. BoGSTL21 expresses in different tissues of Brassica oleracea. The expression level in stigma varies with developmental time, and was mainly overexpressed in mature stigma. The results of qRT-PCR revealed that BoGSTL21 mRNA expression level after self-and cross-pollinations for 0 min to 60 min was consistent with that of RNA-seq data. It was found through yeast two-hybrid that BoGSTL21 protein interacted with pollen development-related protein BoFAB1C, auxin-related protein BoPATL2, and aldolase-type TIM barrel family protein BoF9N12_9. BoGSTL21 gene was successfully induced and expressed in E. coli BL21 (DE3) with a purified protein size of 34 kD, which was consistent with the predicted results. According to the above results, BoGSTL21 may be a novel protein involved in the SI response process, which provides a new content for further research and utilization of self-incompatibility in Brasscia oleracea.

Key words: Brassica oleracea, gene cloning, self-incompatibility, expression analysis, yeast two-hybrid

表1

基因克隆及其荧光定量PCR分析所用引物"

引物名称
Primer name		 引物序列
Primer sequence (5°-3°)		 用途
Functions
BoGSTL21-PGEX-4T-1 F: GATCTGGTTCCGCGTGGATCCATGAGTGCCGGAGTGAGAGTTAG 基因的原核表达
R: CTCGAGTCGACCCGGGAATTCGGGACGTGCTTCTGCTTGG Prokaryotic expression
qRT-PCR F: TTCCTTTGCCGATTTAGTTTGG 荧光定量PCR引物
R: AGTGTTCATCTCCTTAAGCCAA qRT-PCR
dActin F: GGCTGATGGTGAAGATATTCA 内参引物
R: CAAGCACAATACCAGTAGTAC Internal reference primers
BoGSTL21-BK F: TCAGAGGAGGACCTGCATATGAGTGCCGGAGTGAGAGTTA 酵母双杂交引物
Yeast two-hybrid primers
R: TCGACGGATCCCCGGGAATTCGGGACGTGCTTCTGCTTG
BoFAB1C-AD F: GTACCAGATTACGCTCATATGGGGATGGTGAAGTTCTCTGTG
R: ATGCCCACCCGGGTGGAATTCGTTCCATGGCTCAGGAACC
BoPATL2-AD F: GTACCAGATTACGCTCATATGGCTCAAGAAGAGATACAGAAG
R: ATGCCCACCCGGGTGGAATTCTTATGCTTGCGTTTTGAACC
BoF9N12_9-AD F: GTACCAGATTACGCTCATATGGTGGTGTCGCCAAAGATAG
R: ATGCCCACCCGGGTGGAATTCTCAGGTGATGGGTTGGGC
1391 F: GAACTGATCGTTAAAACTGC 通用引物
R: TGGTCTTCTGAGACTGTATC Universal primer
BoGSTL21-GUS F: CAAGCTTGGCTGCAGGTCGACATGTTATACGTTGCGAACGC 基因的启动子活性分析
R: GGTGGACTCCTCTTAGAATTCACTCAATCGTTCTTCTTCCGT Promoter activity analysis

图1

用柱头转录组数据分析BoGSTL21自花授粉和异花授粉后的表达模式 SP: 自花授粉; CP: 异花授粉。"

图2

甘蓝BoGSTL21的cDNA和gDNA序列扩增产物电泳图"

图3

甘蓝BoGSTL21 cDNA结构图及其推导的氨基酸序列 虚线框为N-糖基化位点, 实线框为GST-N结构域, 下画虚线为GST-C结构域。"

图4

BoGSTL21与其他物种GSTL2氨基酸序列的系统进化树"

图5

甘蓝BoGSTL21与其他物种的同源蛋白氨基酸序列比对 黑色: 氨基酸一致性为100%; 深灰色: 氨基酸一致性为75%; 白色: 氨基酸一致性为50%。三角形标记BoGSTL21和BoGSTL2的不同之处; 下画直线为GST-N端结构域, 下画虚线为GST-C端结构域, 同时也代表该蛋白在这一区段保守性高。"

表2

BoGSTL21基因上游调控区顺式作用元件"

相关功能预测
Associated putative function		 启动子顺式作用元件
Cis-elements in the promoter region
脱落酸(ABA)应答 Abscisic acid response ABRE
光响应 Light response G-Box, GT1-motif, TCT-motif
低氧诱导 Anaerobic induction response ARE
促进和增强基因转录 Promote and enhance gene transcription CAAT-box
赤霉素应答 Gibberellin-response GARE-motif, P-box
MYBHv1结合位点 MYBHv1 binding site CCAAT-box
涉及光响应的MYBHv1结合位点 MYB binding site involved in light response MRE
转录启动约-30的核心启动子 Core promoter element around -30 of transcription start TATA-box
低温响应 Low-temperature response LTR
干旱诱导 Drought induced response MBS
水杨酸(SA)反应 Salicylic acid response TCA-element
MeJA茉莉酸响应 MeJA-response CGTCA-motif, TGACG-motif
生长素(IAA)应答 Auxin-response TGA-element

图6

BoGSTL21基因在不同组织中的表达分析"

图7

GUS染色分析 a~c: 种子不同发育时期; d, e: 幼苗不同发育时期; f: 叶的不同发育时期; g~i: 花的不同发育时期; j, k: 果荚的不同发育时期。"

图8

BoGSTL21基因在不同授粉处理后柱头组织中的表达分析 SP: 自花授粉; CP: 异花授粉。"

表3

候选蛋白的功能注释分析"

候选蛋白Candidate protein 蛋白名称
Protein name		 功能注释
Functional annotations
BoFAB1C 1-磷脂酰肌醇-3-磷酸5-激酶
1-phosphatidylinositol-3-
phosphate 5-kinase		 具有1-磷脂酰肌醇-3-磷酸5-激酶活性, ATP结合,参与磷脂酰肌醇磷酸化, 花粉发育, 气孔关闭的生物过程。
It has 1-phosphatidylinositol-3-phosphate 5-kinase activity, ATP binding, participates in the biological processes of phosphatidylinositol phosphorylation, pollen development, and stomatal closure.
BoPATL2 patellin-2 PATL属于具有高尔基动力学(GOLD)结构域和Sec14p-like结构域串联的蛋白质家族。PATL受生长素调节。
PATL belongs to a family of proteins with a Golgi dynamics (GOLD) domain and a Sec14p-like domain tandem. PATL is regulated by auxin.
BoF9N12_9 磷酸核糖3-差向异构酶
Ribulose-phosphate 3-epimerase		 属于醛缩酶型TIM桶家族蛋白, 参与碳水化合物代谢过程, 磷酸戊糖途径, 氨基酸的生物合成。
Belongs to the aldolase type TIM barrel family protein, involved in carbohydrate metabolism, pentose phosphate pathway, amino acid biosynthesis.

表4

质粒共转化酵母的相互作用分析"

编号
No.		 酵母菌种(质粒)
Yeast strain (plasmid)		 培养基
Yeast medium		 菌斑
Colony		 颜色
Color
1 Y2HGold (pGADT7-T×pGBKT7-53) SD/-Leu/-Trp 是Yes 白色White
2 Y2HGold (pGADT7-T×pGBKT7-Lam) SD/-Leu/-Trp 是Yes 红色Red
3 Y2HGold (pGADT7-BoFAB1C×pGBKT7-BoGSTL21) SD/-Leu/-Trp 是Yes 白色White
4 Y2HGold (pGADT7-BoPATL2×pGBKT7-BoGSTL21) SD/-Leu/-Trp 是Yes 白色White
5 Y2HGold (pGADT7-BoF9N12×pGBKT7-BoGSTL21) SD/-Leu/-Trp 是Yes 白色White
6 Y2HGold (pGADT7-T×pGBKT7-53) SD/-Ade/-His/-Leu/-Trp 是Yes 白色White
7 Y2HGold (pGADT7-T×pGBKT7-Lam) SD/-Ade/-His/-Leu/-Trp 是Yes 无色No
8 Y2HGold (pGADT7-BoFAB1C×pGBKT7-BoGSTL21) SD/-Ade/-His/-Leu/-Trp 是Yes 白色White
9 Y2HGold (pGADT7-BoPATL2×pGBKT7-BoGSTL21) SD/-Ade/-His/-Leu/-Trp 是Yes 白色White
10 Y2HGold (pGADT7-BoF9N12×pGBKT7-BoGSTL21) SD/-Ade/-His/-Leu/-Trp 是Yes 白色White

图9

酵母双杂交验证BoFAB1C/BoGSTL21、BoPATL2/ BoGSTL21、BoF9N12_9/BoGSTL21之间的相互作用 第1~3排表示pGADT7-BoFAB1C/BoPATL2/BoF9N12_9× pGBDT7- BoGSTL21; 第4排表示阳性对照(pGADT7-T× pGBDT7-p53); 第5排表示阴性对照(pGADT7-T×pGBDT7-Lam)。"

图10

BoGSTL21蛋白的原核表达 M: 蛋白分子量标准; 1: BoGSTL21-pGEX-4T-1未诱导蛋白; 2: BoGSTL21-pGEX-4T-1诱导蛋白; 3: pGEX-4T-1蛋白; 4: BoGSTL21-pGEX-4T-1纯化蛋白。"

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