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作物学报 ›› 2023, Vol. 49 ›› Issue (12): 3143-3153.doi: 10.3724/SP.J.1006.2023.32020

• 综述 •    下一篇

植物有性生殖对高温胁迫的响应机制

陈赛华*(), 仲伟杰, 薛明   

  1. 江苏省作物基因组学和分子育种重点实验室 / 植物功能基因组学教育部重点实验室 / 江苏省作物遗传生理重点实验室, 扬州大学农学院, 江苏扬州 225009
  • 收稿日期:2023-05-06 接受日期:2023-06-28 出版日期:2023-12-12 网络出版日期:2023-07-11
  • 通讯作者: * 陈赛华, E-mail: chensaihua@yzu.edu.cn
  • 基金资助:
    崖州湾种子实验室与中国种子集团联合资助项目(B23YQ1510);江苏省种业振兴揭榜挂帅项目(JBGS[2021]002);江苏省种业振兴揭榜挂帅项目(CX(21)3100)

Advances in heat-stress responses at sexual reproduction stage in plants

CHEN Sai-Hua*(), ZHONG Wei-Jie, XUE Ming   

  1. Key Laboratory of Plant Functional Genomics of the Ministry of Education / Co-Innovation Center for Modern Production Technology of Grain Crops, Yangzhou University, Yangzhou 225009, Jiangsu, China
  • Received:2023-05-06 Accepted:2023-06-28 Published:2023-12-12 Published online:2023-07-11
  • Contact: * E-mail: chensaihua@yzu.edu.cn
  • Supported by:
    Hainan Yazhou Bay Seed Laboratory and China National Seed Group(B23YQ1510);Jiangsu Province Government Funding(JBGS[2021]002);Jiangsu Agricultural Science and Technology Innovation Fund(CX(21)3100)

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摘要：

极端高温天气频发, 严重威胁农作物的生产。高温胁迫对有性生殖过程的影响与作物减产密切相关, 解析其中的分子机制对于指导作物耐高温遗传改良具有重要意义。然而, 与模式植物拟南芥相比, 目前有关作物有性生殖过程中耐高温的相关研究十分有限。本文从植物有性生殖过程出发, 概述了在减数分裂、花粉绒毡层降解、小孢子发育、花粉管萌发与授精以及籽粒发育等一系列生殖发育过程中响应高温胁迫的分子机制。据此, 我们提出了作物耐高温改良的可行策略, 以期为耐高温品种的遗传改良提供理论依据。

关键词: 植物, 有性生殖, 热胁迫, 响应, 分子机制, 耐热

Abstract:

The occurrence of extreme hot weather poses a threat to crop production. Heat stress suffered at reproductive stage in crops is always correlated with crop yield losses, and thus the underlying molecular mechanisms are of great significance in crop thermotolerance improvement. However, relevant studies are mainly focused on Arabidopsis and less is known in crops. From the perspective of plants, here, we reviewed the heat-stress responses at reproductive stage, including meiosis process, tapetum degradation, microspore development, pollen-tube germination, and fertilization, as well as seed development. Based on these advances, we proposed feasible strategies for thermotolerance improvement, which will pave a way for the breeding of heat-tolerant crop varieties.

Key words: plants, sexual reproduction, heat stress, response, molecular mechanism, thermotolerance

图1

植物生殖发育阶段对高温胁迫的分子响应 图中{ }表示其中响应高温的为蛋白、蛋白复合物或者信号通路, →表示促进, ┫表示抑制, -表示互作, /表示或者。{ }↑表示整体响应高温后上调; { }↓表示整体响应高温后下调。"

表1

不同生殖发育阶段高温响应的基因，蛋白或信号通路及相应的高温应对策略"

生殖阶段
Reproductive stage		 物种
Species		 基因/蛋白/信号通路
Gene/Protein/Signal pathway		 高温的响应
Responses to heat stress		 效应
Effects		 参考文献
Reference		 应对高温的策略(高温下)
Strategies for heat resistance
(Under heat stress)
减数分裂
Meiosis		 拟南芥
Arabidopsis		 SPO11, DMC1, ASY4, ASY1, and ZYP1 被抑制
Repressed		 遗传交叉的数量显著减少，联会复合体组装受到影响Decrease CO number, affect the SC assembly [14] 增强蛋白的稳定性
To increase their stability
拟南芥
Arabidopsis		 ATM → MRE11-RAD50-NBS1 被诱导
Upregulated		 确保DSB修复成功，维护高温下减数分裂染色体的完整性 Ensure the recovery of DSB [17] 增强ATM的表达
To increase ATM expression
大麦
Barley		 ASY1 → DMC1-RecA 被诱导
Upregulated		 促进同源染色体序列交换
Promote the exchange of		 [19] 增强ASY1的表达
玉米
Corn		 HSP101 → RAD51 被诱导
Upregulated		 促进DSB修复与同源重组
Ensure the recovery of DSB and recombination		 [20] 增强HSP101的表达
To increase HSP101 expression
玉米
Corn		 INVAN6 → Tre6P/SnRK1 被诱导
Upregulated		 调控花药中糖的稳态
Regulate the sugar homeostasis in anthers		 [21] 增强INVAN6的表达
To increase INVAN6 expression
水稻
Rice		 cCu/Zn-SOD1/OsCATB ┫ROS/MDA 被抑制
Repressed		 花粉中的ROS/MDA含量上升，花粉活力下降
The ROS content elevated and the pollen viability decreased		 [22, 23] 增强 SOD1和CATB的酶活
To enhance SOD1 and CATB
e
杨树
Populus		 PtCYCA1;2/CYCA1;2 被抑制
Repressed		 减数分裂I期到减数分裂II期的转变失败
Hinder the transition from meiosis Ⅰ to meiosis Ⅱ		 [25] 增强 PtCYCA1;2的表达
To increase PtCYCA1;2
绒毡层降解
Tapetal
degradation		 拟南芥
Arabidopsis		 MYB80 → UNDEAD 被抑制
Repressed		 绒毡层的降解提前和花粉败育
Exhibit premature tapetal degradation and pollen abortion		 [29] 增强此通路
To enhance this pathway
棉花
Cotton		 GhMYB66/GhMYB4 → GhCKI 被诱导
Upregulated		 延迟绒毡层的程序性细胞死亡和花药开裂
Delay tapetum programmed cell death and anther dehiscence		 [30, 31, 32] 减弱此通路
To attenuate this pathway
小孢子发育Microspore development 拟南芥
Arabidopsis		 bZIP17/bZIP28/bZIP60 → UPR 被诱导
Upregulated		 UPR的感受与信号转导，促进UPR
As sensor and signal transducer of UPR, promote UPR		 [34, 35] 增强此通路
To enhance this pathway


拟南芥
Arabidopsis		 IRE1-AtbZIP60 → SEC31A → UPR 被诱导
Upregulated		 激活UPR，维持高温下花粉的发育与活力
Activate UPR, maintain the pollen viability		 [36] 增强此通路
To enhance this pathway
番茄/拟南芥
Tomato/ Arabidopsis		 HsfA1a, HsfA2, AtREN1 被诱导
Upregulated		 增加了花粉高温抗性
Improve the thermotolerance of pollen		 [37, 38, 39] 增强这些基因的表达
To increase these genes
水稻
Rice		 OsHSP60-3B-FLO6 被诱导
Upregulated		 稳定FLO6蛋白，调控后期淀粉体的发育 Stabilize FLO6 and modulate starch granule biogenesis [40] 增强OsHSP60对FLO6的稳定
To stabilize FLO6 by enhancing OsHSP60
番茄
Tomato		 SlACS3/SlACS11 → ethylene synthesis 被诱导
Upregulated		 促进乙烯合成
Promote ethylene synthesis		 [42] 促进 SlACS3/SlACS11的表达
To promote SlACS3/SlACS11
番茄
Tomato		 SlETR3/SlCTR2 → ethylene signal transduction 被诱导
Upregulated		 促进乙烯信号转导
Promote ethylene signal transduction		 [42] 促进SlETR3/SlCTR2的表达
To promote SlETR3/SlCTR2
拟南芥, 大麦
Arabidopsis, barley		 YUC2, YUC6, TAA1/TIR2 被抑制
Repressed		 降低了内源生长素的含量，导致花粉败育
Decrease the Auxin content and make pollen abortion		 [43] 调节生长素合成基因表达
Modulate genes in Auxin biosynthesis
棉花
Cotton		 miR160 ┫ARF10/ARF17 被诱导
Upregulated		 激活生长素信号，增强高温敏感性
Activate the auxin signaling pathway and enhance heat sensitivity		 [46] 抑制miR160的表达
To repress the expression of miR160
花粉管萌发与授精
Pollen tube growth and fertilization		 拟南芥
Arabidopsis		 CNGC16 → HsfA2/HsfB1d 被诱导
Upregulated		 感受钙离子信号，激活热激反应
Sensitive to calcium, activate heat shock response		 [49] 增强此信号通路
To enhance this pathway
拟南芥
Arabidopsis		 TMS1 被诱导
Upregulated		 促进高温下花粉管的正常生长
Promote the pollen tube growth under heat stress		 [50] 增强 TMS1的表达
To promote TMS1 gene expression
拟南芥
Arabidopsis		 CLE45-SKM1/SKM2 被诱导
Upregulated		 改善高温下花粉管的生长缺陷
Compensate the deficiency of pollen tube growth		 [51] 增强CLE45的表达
To enhance the expression of CLE45
籽粒发育
Seed development		 拟南芥
Arabidopsis		 SnRK1-FUSCA3(FUS3) 被诱导
Upregulated		 SnRK1介导了胚胎发育阶段FUS3的体内磷酸化
The phosphorylation of FUS3 mediated by SnRK1		 [53] 稳定SnRK1的活性
To stabilize the activity of SnRK1
二穗短柄草
Brachypodium		 APR6 → H2A.Z 被抑制
Repressed		 H2A.Z在染色体上占据的规律被扰乱
Impair the profile and deposition of H2A.Z		 [56] 增强 APR6的表达
To enhance APR6
其他
Others		 番茄
Tomato		 F3H → Flavonols ┫ROS 被诱导
Upregulated		 黄酮醇物质作为抗氧化剂抑制ROS的积累
Reduce ROS accumulation by flavonols		 [52] 增强F3H等基因的表达
To promote F3H gene
水稻
Rice		 HTH5 ┫ROS 被诱导
Upregulated		 抑制ROS的积累，提高花粉活力
Reduce ROS accumulation, improve pollen viability		 [57] 增强HTH5的表达
To promote HTH5
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