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作物学报 ›› 2020, Vol. 46 ›› Issue (9): 1368-1379.doi: 10.3724/SP.J.1006.2020.94188

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

棉花GhPIF4调控高温下花药败育机制初探

陈淼(), 谢赛, 王超智, 李焱龙, 张献龙, 闵玲*()   

  1. 华中农业大学植物科学技术学院 / 作物遗传改良国家重点实验室, 湖北武汉 430070
  • 收稿日期:2019-11-30 接受日期:2020-03-24 出版日期:2020-09-12 网络出版日期:2020-04-10
  • 通讯作者: 闵玲
  • 作者简介:E-mail: 2857520681@qq.com
  • 基金资助:
    本研究由中央高校基础研究经费(140419001);国家重点研发计划项目资助(2018YFD0100403)

Mechanism of GhPIF4 regulating anther abortion under high temperature stress in cotton

CHEN Miao(), XIE Sai, WANG Chao-Zhi, LI Yan-Long, ZHANG Xian-Long, MIN Ling*()   

  1. National Key Laboratory of Crop Genetic Improvement / College of Plant Science and Technology, Huazhong Agricultural University, Wuhan 430070, Hubei, China
  • Received:2019-11-30 Accepted:2020-03-24 Published:2020-09-12 Published online:2020-04-10
  • Contact: Ling MIN
  • Supported by:
    Funding from Fundamental Research Funds for the Central Universities(140419001);National Key Research and Development Program of China(2018YFD0100403)

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摘要：

PIF4属于PIF家族(PHYTOCHROME INTERACTING FACTORS FAMILY), 是一个响应温度变化的关键性转录因子, 广泛参与植物热信号传导和激素信号通路。本研究从陆地棉‘YZ1’中克隆获得GhPIF4基因, 亚细胞定位结果表明, GhPIF4是一个核蛋白。qRT-PCR分析和proGhPIF4:GUS转基因棉花GUS染色结果显示, GhPIF4基因在棉花花药中高量表达, 且其表达在高温敏感型材料‘H05’中受到高温的显著诱导。在棉花中超量表达GhPIF4, 不同的转基因株系表现不同的育性, 表达量高的3个转基因系(OE5、OE7和OE19)开花当天的花药不开裂, 花粉的活力均低于对照‘YZ1’, 而超表达程度略低的OE10, 花药正常开裂, 花粉活性正常, 表明GhPIF4导致的花药败育具有剂量效应。对OE7和OE19在9~14 mm (绒毡层降解期)、14~19 mm (有丝分裂I期)中生长素的含量以及生长素合成关键基因GhTAA1、GhYUC2、GhCYP71A13的表达分析发现基因的表达与生长素含量下降变化一致。推测超表达GhPIF4在花药后期可能部分模拟棉花的高温响应状态, 而组成型超表达GhPIF4可能改变了营养器官或早期花药中生长素的含量, 亦或花药中生长素含量过低也会导致花药败育。以上结果为深入解析GhPIF4基因功能及了解高温胁迫下棉花花药败育的机制提供参考。

关键词: 棉花, 高温, PIF4, 生长素, 花药/花粉败育

Abstract:

PIF4, which belongs to the phytochrome interacting factor family, is a key transcription factor in response to temperature changes, and is widely involved in plant heat signaling transduction and hormonal signaling pathways. In this study, the GhPIF4 gene was cloned from ‘YZ1’ (Gossypium hirsutum). Subcellular localization results indicated that GhPIF4 is a nuclear located protein. qRT-PCR analysis and GUS staining of proGhPIF4:GUS transgenic cotton plants showed that the GhPIF4 gene was highly expressed in the anther, and its expression was significantly induced by high temperature (HT) in the ‘H05’ (HT sensitive line). With the overexpression of GhPIF4, different cotton transgenic lines showed different male fertility performances. Three transgenic lines with high GhPIF4 expression, OE5, OE7, and OE19, showed indehiscent anthers in the flower on the day of flowering and lower pollen activity than wild type ‘YZ1’. However, the transgenic line OE10 with slighter overexpression showed normal anther dehiscence and pollen activity. These results indicated that GhPIF4 causes anther abortion in an expression level-dependent manner. Furthermore, the IAA content was measured in OE7 and OE19 anthers at the tapetum degradation stage (9-14 mm buds) and mitotic stage (14-19 mm buds). As a result, the IAA content in OE7 and OE19 decreased compared to that in ‘YZ1’. Then, we analyzed the expression of auxin biosynthesis genes, including GhTAA1, GhYUC2, and GhCYP71A13, showing that the expression of these genes was down-regulated in OE7 and OE19, which was consistent with the decrease in IAA content. We hypothesize that overexpression of GhPIF4 might partially simulate the response of late stage anthers to HT, while constitutive overexpression of GhPIF4 may alter the auxin content in vegetative organs or early stage anthers, or the decline of IAA may lead to the anther abortion. These results provide a reference for further analysis of the function of GhPIF4 gene and understanding the mechanism of cotton anther abortion under HT.

Key words: cotton, high temperature, PIF4, auxin, anther/pollen abortion

表1

实验中用到的载体和菌株"

载体/菌株 Vector/strain 用途 Purpose
pGEM-T Easy载体 pGEM-T Easy vector TA克隆 TA clone
pDONOR221载体 pDONOR221 vector BP反应 BP reaction
pGBW409载体 pGBW409 vector 35S:GhPIF4:His载体的构建 Construction of 35S:GhPIF4:His vector
pMDC43载体 pMDC43 vector GhPIF4亚细胞定位 Subcelluar localization of GhPIF4
pGWB433载体 pGWB433 vector GhPIF4启动子表达模式分析 Analysis of GhPIF4 promoter expression pattern
大肠杆菌TOP10 E. coli TOP10 载体转化 Vector transformation
农杆菌GV3101 Agrobacterium GV3101 棉花下胚轴的转化及烟草的侵染 Transformation of cotton hypocotyl and tobacco infection

表2

本实验中所用的引物及用途"

引物名称Primer name 引物序列Primer sequence (5°-3°) 用途Purpose
GhPIF4-CDS-F ATGGATCACCAACATGAACAACA Gene amplification
GhPIF4-CDS-R TCAGTTAAATCCCGGATTGGCAG Gene amplification
GhPIF4-CDS-BP-F GGGGACAAGTTTGTACAAAAAAGCAGGCTGGATGGAT
CACCAACATGAACAACA		 BP reaction
GhPIF4-CDS-BP-R GGGGACCACTTTGTACAAGAAAGCTGGGTGTCAGTTA
AATCCCGGATTGGCAG		 BP reaction
proGhPIF4-F TTTTTGTCTCCCATTACAGTATC Gene promoter amplification
proGhPIF4-R GGGTCATAAACTGGAAATTCAGA Gene promoter amplification
GhUB7-qRT-F CTTGACCTTCTTCTTCTTGTGCTTG qRT-PCR
GhUB7-qRT-R GAAGGCATTCCACCTGACCAAC qRT-PCR
GhPIF4-CDS-qRT-F AACTACACCTCAAAGTCCCACGG qRT-PCR
GhPIF4-CDS-qRT-R CCCGGATTGGCAGTGGTC qRT-PCR
GhTAA1-qRT-F GGTCTTAAAAAGGTTGGGGCTTA qRT-PCR
GhTAA1-qRT-R TTAGCTTGGGTATGTGTTTGATTTG qRT-PCR
GhYUC2-qRT-F ACCGATGTGGGTTTTGGCGAAT qRT-PCR
GhYUC2-qRT-R CTCAGCATTTTCCCCGGTAGCA qRT-PCR
GhCYP71A13-qRT-F CGTAAACAGACCGAAACGCAGC qRT-PCR
GhCYP71A13-qRT-R GTGGTTGCGGAAAAGAGTTCGC qRT-PCR

图1

陆地棉中GhPIFs和 GhPIL1s的进化树及GhPIF4基因结构特征 A: 陆地棉中GhPIFs和 GhPIL1s的进化树分析; B: GhPIF4与AtPIF4和AtPIF5的氨基酸序列比对, 红线表示bHLH domain; C: GhPIF4的基因结构。红色方框表示外显子, 黑线表示内含子。"

图2

GhPIF4蛋白的亚细胞定位 35S:GFP: GFP空载在烟草中的亚细胞定位; 35S:GhPIF4:GFP: GFP 标记的GhPIF4蛋白在烟草中的亚细胞定位。Bar = 20 μm。"

图3

GhPIF4 组织表达及高温诱导表达分析 A: GhPIF4在‘H05’中的组织表达分析。B: GhPIF4在‘YZ1’中的组织表达分析。C: proGhPIF4:GUS转基因棉花组织表达情况; a: 开花当天的整体花药; b: 开花当天的单个花药; c: 萼片; d: 花瓣; e: 幼嫩的叶子。D: GhPIF4在‘H05’不同发育时期的花药中高温诱导表达分析。a、c、d、e图, bar = 1 cm; b图bar = 100 μm; HN和HH分别表示正常温度和高温胁迫条件下的‘H05’; < 7 mm表示花蕾长度小于7 mm花药的混合; **P < 0.01。"

图4

35S:GhPIF4:His转基因棉花GhPIF4表达量检测与拷贝数检测 A: T0代可育和败育材料中GhPIF4的表达量分析; B: T0代转基因材料的拷贝数检测。"

图5

35S:GhPIF4:His转基因棉花表型 A: 整体花药, bar = 1 cm; B: TTC染色单个花药, bar = 500 μm。"

图6

超表达GhPIF4败育棉花花药中生长素含量及生长素合成相关基因表达量 A: 超表达GhPIF4败育棉花花药中生长素含量降低; B~D: 超表达GhPIF4败育棉花花药中生长素合成相关基因表达分析; **P < 0.01。"

图7

GhirPIF4和GhPIF4的氨基酸序列比对 红色方框表示APB motif。"

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