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作物学报 ›› 2026, Vol. 52 ›› Issue (6): 1698-1710.doi: 10.3724/SP.J.1006.2026.54133

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

甘蓝自交不亲和相关基因BoPUB3L的克隆与表达分析

左同鸿1,**(), 张贺翠1,**(), 曾静2, 朱利泉1,*()   

  1. 1 西南大学农学与生物科技学院, 重庆 400716
    2 长江师范学院现代农业与生物工程学院, 重庆 408100
  • 收稿日期:2025-10-21 接受日期:2026-02-27 出版日期:2026-06-12 网络出版日期:2026-03-09
  • 通讯作者: * 朱利泉, E-mail: zhuliquan@swu.edu.cn
  • 作者简介:左同鸿, E-mail: zuotongh@163.com;张贺翠, E-mail: zhanghecui226@163.com

    **同等贡献

  • 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(CYB22138);重庆市研究生科研创新项目(CYS18085);横向项目(M2022052)

Molecular cloning and expression analysis of BoPUB3L associated with self-incompatibility in Brasscia oleracea

Zuo Tong-Hong1,**(), Zhang He-Cui1,**(), Zeng Jing2, Zhu Li-Quan1,*()   

  1. 1 College of Agronomy and Biotechnology, Southwest University, Chongqing 400716, China
    2 College of Modern Agriculture and Bioengineering, Yangtze Normal University, Chongqing 408100, China
  • Received:2025-10-21 Accepted:2026-02-27 Published:2026-06-12 Published online:2026-03-09
  • Contact: * Zhu Li-Quan, E-mail: zhuliquan@swu.edu.cn
  • About author:

    **Contributed equally to this work

  • Supported by:
    National Natural Science Foundation of China(31572127);Chongqing Graduate Research and Innovation Project(CYB22138);Chongqing Graduate Research and Innovation Project(CYS18085);Horizontal Project(M2022052)

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摘要：

PUB蛋白(plant U-box protein)是一类E3泛素连接酶, 在植物自交不亲和、激素反应、防御和非生物胁迫反应等过程中发挥作用。本研究通过对甘蓝自花授粉(self-pollination, SP) 0~60 min的花柱转录组数据进行分析, 筛选到1个受SP诱导上调表达的PUB蛋白家族基因BoPUB3L。该基因的开放阅读框为2214 bp, 编码737个氨基酸, 理论等电点为6.04, 其相对分子质量为81.27 kD, 不含信号肽和跨膜域, 含有1个U-box结构域和5个ARM臂重复结构域, 定位于细胞核中。BoPUB3L启动子中含有光响应、脱落酸应答、生长素应答、赤霉素应答、与分生组织表达相关等多种顺式作用元件。BoPUB3L基因在甘蓝的不同组织中均有表达, 在花柱中的表达量最高, 在萼片、花瓣、花蕾中的表达量次之, 同GUS染色结果一致。该基因在自花授粉0~30 min持续上调表达, 在自花授粉15 min时差异达到最大, 是异花授粉时的15.65倍。酵母双杂交、pull down和BiFC试验结果表明, BoPUB3L蛋白与SRK胞内激酶域有相互作用。以上研究结果推测, BoPUB3L可能是一种参与甘蓝自交不亲和反应过程的新基因, 这对进一步了解甘蓝自交不亲和机制提供了新的研究视角。

关键词: 甘蓝, 基因克隆, 自交不亲和, 表达分析, 酵母双杂交

Abstract:

U-box E3 ligases (PUBs) regulate plant self-incompatibility (SI), hormone signaling, and stress responses. By mining time-course transcriptome data from Brassica oleracea following self-pollination (0-60 min), we identified BoPUB3L, a PUB gene that is rapidly and specifically upregulated by self-pollination. The 2214 bp open reading frame encodes a 737-aa protein (81.27 kD; pI 6.04) containing a single U-box and five ARM repeats, lacking a signal peptide and transmembrane domain, and localizing to the nucleus. The BoPUB3L promoter contains multiple cis-acting elements related to light responsiveness, abscisic acid, auxin, and gibberellin responses, as well as meristem-associated expression. BoPUB3L is most highly expressed in the style, followed by sepals, petals, and buds, as confirmed by GUS staining. Transcript abundance increased continuously during the first 30 min after self-pollination and peaked at 15 min, reaching a level 15.65-fold higher than that after cross-pollination. Yeast two-hybrid, pull-down, and BiFC assays further showed that BoPUB3L interacts with the kinase domain of the S-locus receptor kinase (SRK). Together, these results identify BoPUB3L as a previously uncharacterized component of the SI response in B. oleracea and provide new insight into the molecular basis of self-incompatibility in crucifers.

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

表1

本研究所用引物"

引物名称 引物序列 用途
Primer name Primer sequence (5′-3′) Function
BoPUB3L-BK F: ATGGCCATGGAGGCCGAATTCATGGATCCTGTGCCGGTTC 酵母双杂交
Yeast two-hybrid
R: ATGCGGCCGCTGCAGGTCGACTGATCTACCTTTCTTCATCCTTGC
BoSRKj-AD F: CCGGAATTCCTCTGGAAAAGAAAACAAAAGC
R: CGCGGATCCTTAGGCATCGATGAGTGAGCAGGT
BoPUB3L-GUS F: CAAGCTTGGCTGCAGGTCGACAGTGTTCTCAGTCACACCACATATG 启动子活性分析Promoter activity analysis
R: GGTGGACTCCTCTTAGAATTCTCAAAACTACAAAGCTGAAGAACAC
RT-qPCR F: ATTGCAAAGTACCTCCTGATGA 荧光定量PCR引物
RT-qPCR primers
R: CATTCAATCTCCTGCATACACC
dAction F: GGCTGATGGTGAAGATATTCA 内参引物
Internal reference primers
R: CAAGCACAATACCAGTAGTAC
BoPUB3L-nYFP F: GGGGACAAGTTTGTACAAAAAAGCAGGCTTCATGGATCCTGTGCCGGTTC 双分子荧光互补Bimolecular fluorescence complementation
R: GGGGACCACTTTGTACAAGAAAGCTGGGTCTGATCTACCTTTCTTCATCCTTGC
BoSRK-cYFP F: GGGGACAAGTTTGTACAAAAAAGCAGGCTTCAAAAGAAAACAAAAGCGTGCAAAAG
R: GGGGACCACTTTGTACAAGAAAGCTGGGTCTTAGGCATCGATGAGTGAGCAGGTG
BoPUB3L-GST F: GATCTGGTTCCGCGTGGATCCATGGATCCTGTGCCGGTTC 原核表达
Prokaryotic expression
R: CTCGAGTCGACCCGGGAATTCTGATCTACCTTTCTTCATCCTTGC
BoPUB3L-GFP F: AAGTCCGGAGCTAGCTCTAGAATGGATCCTGTGCCGGTTC 亚细胞定位
Subcellular location
R: GCCCTTGCTCACCATGGATCCTGATCTACCTTTCTTCATCCTTGC

图1

自花授粉与异花授粉后差异表达基因维恩图 UP: 未授粉; SP: 自花授粉; CP: 异花授粉。UP&SP指以未授粉为对照, 自花授粉差异表达基因; UP&CP指以未授粉为对照, 异花授粉差异表达基因。15、30、60指授粉时间(min)。"

图2

转录组数据分析BoPUB3L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3

甘蓝BoPUB3L基因的cDNA和启动子序列扩增电泳图 M: DNA分子量标准; 1: BoPUB3L基因cDNA的PCR扩增; 2: BoPUB3L启动子序列的PCR扩增。"

图4

甘蓝BoPUB3L的cDNA与其氨基酸序列 双虚线框为酪蛋白激酶II磷酸化位点; 矩形为蛋白激酶C磷酸化位点; 椭圆为N-糖基化位点; 梯形虚线框为cAMP或cGMP依赖性蛋白激酶磷酸化位点; 阴影部分表示N-肉豆蔻酰化位点; 中括号表示酰胺化位点; 下画虚线表示富含丝氨酸区域; 实线框表示U-box结构域; 下画实线为ARM结构域。"

图5

BoPUB3L与其他物种PUB3蛋白的系统发育(A)、保守基序(B)、结构域(C)分析"

图6

甘蓝BoPUB3L授粉后表达分析(A)和组织表达分析(B) SP: 自花授粉; CP: 异花授粉。"

表2

BoPUB3L基因上游调控区顺式作用元件"

启动子顺式作用元件 相关功能预测
Cis-elements in the promoter region Associated putative function
AE-box, AT1-motif, ATC-motif, Box 4, Box II 光响应Light responsive
ABRE, AAGAA-motif 脱落酸(ABA)应答Abscisic acid responsiveness
TGA-element, ERE 生长素(IAA)应答Auxin-responsive
GARE-motif, F-box 赤霉素(GA)应答Gibberellin response
TCA-element 水杨酸(SA)反应Salicylic acid responsiveness
GCN4_motif 参与胚乳表达的顺式调控元件Cis-regulatory element involved in endosperm expression
CAT-box 与分生组织表达有关的顺式调控元件
Cis-acting regulatory element related to meristem expression
ARE 厌氧诱导Anaerobic induction
W box WRKY转录因子结合位点WRKY transcription factor binding site
STRE 热应激响应的顺式作用元件Cis-acting regulatory element related to heat stress response
MYB, MYB-binding site, MYC 参与干旱、低温、盐胁迫等逆境响应
Involved in drought, low temperature, salt stress and other stress responses
TC-rich repeats 压力与防御相关反应元件Stress and defense related response elements
WUN-motif 创伤响应元件Trauma response element

图7

GUS染色图 a, b: 幼苗; c: 叶片; d-g: 花; h: 果荚。"

图8

BoPUB3L蛋白的原核表达 M: 蛋白分子量标准; 1: BoPUB3L-GST未诱导蛋白; 2: BoPUB3L-GST诱导蛋白; 3: pGEX-4T-1纯化蛋白; 4: BoPUB3L-GST纯化蛋白; 箭头指纯化后蛋白。"

图9

BoPUB3L-GFP融合蛋白在拟南芥原生质体(A)和烟草叶肉细胞(B)中的亚细胞定位 红色箭头所指为细胞核; 标尺为20 μm。"

图10

酵母融合株的相互作用检测(A)、pull down试验结果(B)和烟草叶肉细胞中的荧光共聚焦检测结果(C) DDO: SD/-Leu/-Trp酵母二缺培养基; QDO: SD/-Leu/-Trp/-His/-Ade/酵母四缺培养基。标尺为20 μm。"

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