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作物学报 ›› 2021, Vol. 47 ›› Issue (1): 30-41.doi: 10.3724/SP.J.1006.2021.04090

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

棉花GbSTK基因调控开花和黄萎病抗性的功能研究

崔静(), 王志城, 张新雨, 柯会锋, 吴立强, 王省芬, 张桂寅, 马峙英, 张艳*()   

  1. 华北作物改良与调控国家重点实验室 / 河北省棉花产业协同创新中心 / 河北农业大学, 河北保定 071001
  • 收稿日期:2020-04-11 接受日期:2020-08-19 出版日期:2021-01-12 网络出版日期:2020-09-21
  • 通讯作者: 张艳
  • 作者简介:E-mail: cuijing1240861235@qq.com
  • 基金资助:
    河北省杰出青年科学基金项目(C2019204365);国家现代农业产业技术体系建设专项(CARS-15-03);河北省青年拔尖人才支持计划资助

Function study of cotton GbSTK gene in regulating flowering and Verticillium wilt resistance

CUI Jing(), WANG Zhi-Cheng, ZHANG Xin-Yu, KE Hui-Feng, WU Li-Qiang, WANG Xing-Fen, ZHANG Gui-Yin, MA Zhi-Ying, ZHANG Yan*()   

  1. State Key Laboratory of North China Crop Improvement and Regulation / Co-Innovation Center for Cotton Industry of Hebei Province / Hebei Agricultural University, Baoding 071001, Hebei, China
  • Received:2020-04-11 Accepted:2020-08-19 Published:2021-01-12 Published online:2020-09-21
  • Contact: ZHANG Yan
  • Supported by:
    Natural Science Foundation of Hebei Province(C2019204365);China Agriculture Research System(CARS-15-03);Talents Support Program of Hebei Province

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摘要：

棉花是重要的经济作物, 实现既抗病又早熟是棉花重要的育种目标, 但棉花抗病与早熟基因之间的关联性研究报道甚少。本课题组前期鉴定到一个响应黄萎病菌诱导的丝氨酸/苏氨酸蛋白激酶基因GbSTK, 可以提高转基因拟南芥的黄萎病抗性。本研究发现多个抗病棉花品种中STK基因的表达量显著高于感病对照品种H208。在抗病品种农大601中沉默STK基因, 沉默植株在接菌后20 d, 其黄萎病抗性显著降低, 病情指数由27.5 (耐病)升高到63.2 (感病), 表明该基因正向调控棉花黄萎病抗性。对转GbSTK基因拟南芥表型鉴定发现, 其开花时莲座叶片数平均为7.5个, 显著低于空载体对照植株(11.9个), 转基因植株开花时间较对照平均提早5~7 d。进一步对拟南芥开花途径标志基因FT、SOC1和LFY的转录水平检测发现, 转基因不同株系中FT和SOC1的表达水平显著高于对照植株, 而LFY基因的表达受影响较小, 表明STK基因可以影响开花关键基因FT和SOC1的表达, 进而调控植株提早开花。基于酵母双杂交互作蛋白筛选试验, 初步获得30个与GbSTK具有互作关系的候选蛋白, 为进一步揭示GbSTK的分子调控网络奠定了基础。本研究首次鉴定出可同时提高黄萎病抗性和提早开花的一个功能基因, 为棉花抗病早熟遗传改良提供了参考依据。

关键词: 棉花, 丝氨酸/苏氨酸蛋白激酶(GbSTK), 黄萎病抗性, 早熟性

Abstract:

Cotton is an important cash crop, and achieving disease resistance as well as early maturity is an important breeding goal. However, the molecular relationship between disease resistance and early maturity of cotton is still a research gap. In previous study, we identified a GbSTK gene response to V. dahliae that could enhance Verticillium wilt resistance in transgenic Arabidopsis. In this study, we found that GbSTK was more highly expressed in resistance and/or tolerant varieties than in susceptible varieties. Knocking down of GbSTK expression in Nongda 601 showed that the silenced plants displayed serious disease symptoms, with disease index from 27.5 (tolerant) to 63.2 (susceptible), suggesting that GbSTK positively regulates cotton Verticillium wilt resistance. More interesting, we found that GbSTK could promote the development of Arabidopsis. At the flowering time point, the average number of rosette leaves in transgenic lines was 7.5, significantly less than the mock plants, and the transgenic plants bloomed five to seven days earlier than the control. The FT and SOC1 gene, regarded as marker genes in Arabidopsis flowering signaling pathway, maintained higher expression level in transgenic lines than in the mock, indicating that GbSTK could accelerate flowering via regulating FT and SOC1 gene expression. We also obtained 30 candidate proteins that could interacted with GbSTK via yeast double hybrid test, which helped to further revealing the molecular regulatory network of GbSTK. Taken together, we first identified a functional gene that could improve disease resistance and early flowering, which would provide a reference for genetic improvement of cotton disease resistant and early maturity.

Key words: cotton, GbSTK, Verticillium wilt resistance, early maturity

附图1

苗期蘸根法接种黄萎病菌孢子悬浮液 a: 棉苗水培培养。b: 蘸根法接黄萎病菌。"

表1

试验中所用引物"

引物名称
Primer name		 序列
Primer sequence (5°-3°)		 用途
Purpose
STK-F1 CCATATGAAGAATTCAAGCAAATCCTC 基因扩增Gene amplification
STK-R1 GCTGCAGTCAAGGGGCCGTAGGTTGT 基因扩增Gene amplification
VF ACTAGTGCTGCTCGAACTTCTATTGGGAAG 基因扩增Gene amplification
VR GGCGCGCCAGGGGCCGTAGGTTGTGTAACTCT 基因扩增Gene amplification
STK-F2
STK-R2		 ATGCGGCAACGAACAAGAAT
TAGATTGGCACTGAGCTGGT		 实时定量qPCR
实时定量qPCR
GhUBQ14-F CAACGCTCCATCTTGTCCTT 实时定量qPCR
GhUBQ14-R TGATCGTCTTTCCCGTAAGC 实时定量qPCR
BD-F TTTGTAATACGACTCACTATAGGGCG 通用载体Vector
BD-R TTTTCGTTTTAAAACCTAAGAGTCAC 通用载体Vector
T7 TAATACGACTCACTATAGGGCG 通用载体Vector
3°AD AGATGGTGCACGATGCACAG 通用载体Vector
TUB2-F ATCCGTGAAGAGTACCCAGAT 实时定量qPCR
TUB2-R AAGAACCATGCACTCATCAGC 实时定量qPCR
FT-F CTTGGCAGGCAAACAGTGTATGCAC 实时定量qPCR
FT-R GCCACTCTCCCTCTGACAATTGTAGA 实时定量qPCR
SOC1-F AGCTGCAGAAAACGAGAAGCTCTCTG 实时定量qPCR
SOC1-R GGGCTACTCTCTTCATCACCTCTTCC 实时定量qPCR

图1

棉花不同抗病品种中STK基因的表达"

图2

在农大601中沉默STK基因对棉花黄萎病抗性的影响 a: 沉默棉花GhCLA1基因; b: TRV::GbSTK植株中STK基因的沉默效率检测; c: 接菌后25 d对照植株与沉默植株的黄萎病抗性表现; d和e: 接菌后25 d对照植株与沉默植株的发病级别调查和病情指数。"

图3

过表达GbSTK拟南芥表型鉴定及开花标志基因的表达分析 a~c: 过表达GbSTK基因促进拟南芥提早开花; d: 开花时转基因植株与对照植株莲座叶片数比较; e~g: 转基因植株与对照植株中SOC1、FT和LFY基因的表达。误差线表示标准差, *, ***分别表示在0.05和0.001水平显著差异。WT: 野生型拟南芥; OE: 转GbSTK基因拟南芥; OE1和OE2: 转GbSTK基因拟南芥株系。"

图4

诱饵载体pGBKT7-GbSTK的构建与毒性检测 M: DNA marker DL5000; 1~2: GbSTK-T双酶切; 3~6: pGBKT7双酶切; 7: pGBKT7-GbSTK阳性克隆。d: 诱饵载体pGBKT7- GbSTK毒性检测。"

图5

诱饵载体pGBKT7-GbSTK自激活检测 a: SD/-Leu/-Trp培养基酵母生长情况; b: SD/-Ade/-His/-Leu/-Trp/X-α-Gal培养基酵母生长情况; c: 诱饵载体pGBKT7-GbSTK在培养基中生长情况。"

图6

筛选阳性克隆"

表2

与GbSTK互作的候选蛋白"

编号
No.		 蛋白描述
Protein description		 功能报道
Function reported		 功能分类
Functional classification
1 Cytochrome c oxidase subunit 6b-1 Regulate ROS production[22] ROS production
2 Malate dehydrogenase Related to production of ROS and PCD[23]
3 GTPase-activating protein GYP7 Defense response[24] Involved in defense and abiotic/biotic stresses
4 Cytochrome P450 71D8 Defense response[25]
5 Short-chain dehydrogenase reductase 3b Involved in plant defense responses[26]
6 Cysteine proteinase RD21a Involved in plant defense[27]
7 Vacuolar protein sorting-associated protein 55 Involved in plant defense[28]
8 Pathogenesis-related protein STH-2 Defense response[29]
9 (-)-alpha-terpineol synthase Involved in defense and abiotic stresses[30]
10 Glyceraldehyde-3-phosphate dehydrogenase 2 Negatively regulate autophagy and immunity[31]
11 Phosphoglycerate Ubiquitin-conjugating
enzyme E2		 Involved in plant immunity[32]
12 Dirigent protein 16 Responsive to biotic stress[33]
13 Major allergen Pruar 1 Novel Pathogenesis-related protein[34]
14 S-Adenosylmethionine synthase Responsive to abiotic stress[35]
15 S-adenosylmethionine synthase 2 Responsive to abiotic stress[36]
16 Probable inorganic phosphate transporter 1-5 Involved in phosphate homeostasis[37]
17 Glycerophosphodiester phosphodiesterase GDPD1 Response to Pi deficiency[38]
18 Delta-cadinene synthase isozyme C2 Gossypol biosynthetic[39] Secondary metabolism and
synthesis
19 UDP-galactose/UDP-glucose transporter 3 Synthesis of cell walls[40]
20 Aldehyde dehydrogenase family 2 member C4 Regulator of the growth-defense trade-off[41] Plant development and
stress response
21 Probable 2-oxoglutarate-dependent dioxygenase Related to vascular disease/involved in
gibberellin synthesis[42]
22 Patellin-5 Function in plant development and stress
response[43]
23 Triphosphate tunel metalloenzyme 3 Catalyze diverse enzymatic reactions[44]
24 Chloroplastic 2,3-bisphosphoglycerate-independent mutase Involved in photosynthesis and chloroplast development[45] Plant growth and development
25 Synaptotagmin-2 Participates in pollen germination and tube growth[46]
26 Mediator of RNA polymerase II transcription subunit —
27 Alpha-taxilin —
28 Aconitate hydratase, cytoplasmic —
29 Putative zinc transporter At3g08650 —
30 40S ribosomal protein S5 —

附图2

SAMS蛋白和陆地棉GhSAMS蛋白同源性分析"

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