Article(id=1319334578385437251, tenantId=1146029695717560320, journalId=1316781310003478668, issueId=1319334515705754160, articleNumber=null, orderNo=null, doi=10.11983/CBB24133, pmid=null, cstr=32102.14.CBB24133, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1724947200000, receivedDateStr=2024-08-30, revisedDate=null, revisedDateStr=null, acceptedDate=1737302400000, acceptedDateStr=2025-01-20, onlineDate=1792483475537, onlineDateStr=2026-10-20, pubDate=1746806400000, pubDateStr=2025-05-10, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1792483475537, onlineIssueDateStr=2026-10-20, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1792483475537, creator=13701087609, updateTime=1792483475537, updator=13701087609, issue=Issue{id=1319334515705754160, tenantId=1146029695717560320, journalId=1316781310003478668, year='2025', volume='60', issue='3', pageStart='315', pageEnd='482', issueExtLink='null', onlineDate='null', pubDate='1746806400000', pubDateStr='2025-05-10', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1792483460594, creator='13701087609', updateTime=1792483678409, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1319335429350994324, tenantId=1146029695717560320, journalId=1316781310003478668, issueId=1319334515705754160, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1319335429355188629, tenantId=1146029695717560320, journalId=1316781310003478668, issueId=1319334515705754160, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=472, endPage=482, ext={EN=ArticleExt(id=1319334580423868996, articleId=1319334578385437251, tenantId=1146029695717560320, journalId=1316781310003478668, language=EN, title=Research Progress on Factors and Molecular Mechanisms Regulating Plant Organ Abscission, columnId=1317485676040409248, journalTitle=Chinese Bulletin of Botany, columnName=SPECIAL TOPICS, runingTitle=null, highlight=null, articleAbstract=

The organs abscission is a phenomenon in which parts of organs are separated from the body. It is an adaptive strategy evolved by plants in the process of growth and response to environmental changes, which ensures the normal growth and adaptation of plants to the environment. This process involves the abscission zone formation, signal activation, cell separation, and is regulated by both internal (in vivo) physiological processes and external environmental factors including light, temperature, and humidity. In agricultural production, the abscission of plant organs directly affects crop yield. Understanding the regulatory mechanisms of plant organ abscission is greatly important for improving crop yield. In recent years, tremendous progress has been made in the study of organ abscission mechanisms. The studies showed that the mechanisms of organ abscission in plants are largely conservative between species, but also with significant variations. This review delved into the physiological and biochemical mechanisms of plant organ abscission, and summarized the effects of both the environmental factors and the hormones and enzymes in the process. The review provides a solid theoretical support and practical guidance for crop genetic breeding and agricultural production improvement in connection with organ abscission.

, authors=Liu Xupeng, Wang Min, Han Shou'an, Zhu Xuehui, Wang Yanmeng, Pan Mingqi, Zhang Wen*, authorsList=Liu Xupeng, Wang Min, Han Shou'an, Zhu Xuehui, Wang Yanmeng, Pan Mingqi, Zhang Wen, authorCompany=null, correspAuthors=Zhang Wen, authorNote=null, correspAuthorsNote=
*E-mail:
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植物器官脱落是植物体的器官部分从母体脱离的现象, 是植物在生长发育和响应环境变化过程中进化出的一种适应性策略, 以确保植物正常生长和适应环境。该过程涉及离层区形成、特定信号激活以及细胞分离, 同时还受自身生理进程以及光照、温度和湿度等外部环境因素的显著调节。在农业生产实践中, 植物器官脱落直接影响作物的产量。研究植物器官脱落的调控机理对于进一步提高作物产量具有重要意义。近年来, 关于器官脱落机理的研究取得了显著进展。研究表明, 植物器官在脱落机制上具有保守性, 但不同物种间也表现出显著差异。该文深入探讨了植物器官脱落的生理生化机制, 分析了不同环境因素以及激素和酶对其的影响, 旨在为作物遗传育种和农业生产提供理论支撑和实践指导。

, authors=刘旭鹏, 王敏, 韩守安, 朱学慧, 王艳蒙, 潘明启, 张雯*, authorsList=刘旭鹏, 王敏, 韩守安, 朱学慧, 王艳蒙, 潘明启, 张雯, authorCompany=null, correspAuthors=张雯, authorNote=null, correspAuthorsNote=
*张雯, 博士, 研究员, 国家葡萄产业技术体系果实品质调控岗位科学家。现任新疆园艺学会理事, 新疆农业大学及石河子大学硕士生导师, 西北农林科技大学校外合作导师。其研究团队主要围绕葡萄品质形成机理和调控技术开展研究与示范推广工作, 经多年攻关, 确定了葡萄裂果、落粒、色泽、酚含量以及香气等重要品质的关键影响因子, 开发了配套调控技术; 构建了西北产区葡萄顺沟倾斜龙干树形及配套机械化出埋土技术、制干葡萄高架少蔓龙干树形改造技术和优质快速制干技术等多项综合技术, 并在我国最大的葡萄产区新疆推广应用, 提高了优质果率, 有效促进了标准化栽培, 显著降低了生产成本。主持国家级和省部级科研项目10余项。发表论文58篇。授权发明和实用新型专利23项。制定地方标准19项。获自治区科技进步二等奖2项, 地州科技进步二等奖和三等奖各1项。E-mail:
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Cloned genes for grain shattering in rice

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基因名称 基因编号 编码蛋白 染色体定位 参考文献
qSH1 Os01g0848400 BEL1型同源异型蛋白 1 Konishi et al., 2006
OsSh1 Os03g0650000 YABBY转录因子 3 Lin et al., 2012
ObSH3 OBART03G27740 YABBY转录因子 3 Lv et al., 2018
Sh4/SHA1 Os04g0670900 Myb3转录因子 4 Li et al., 2006; Lin et al., 2007
SHAT1 Os04g0649100 AP2转录因子 4 Zhou et al., 2012
SH5 Os05g0455200 BEL1型同源异型蛋白 5 Yoon et al., 2014
Sh-h/OsCPL1 Os07g0207700 CTD磷酸化酶 7 Ji et al., 2010
OsGRF4/PT2 Os02g0701300 生长调控因子 2 Sun et al., 2016
OsNPC1 Os03g0826600 磷酸酯酶C1 3 Cao et al., 2016
GL4 ORGLA04G0254300 Myb3转录因子 4 Wu et al., 2017
OsLG1 Os04g0656500 SQUAMOSA启动子结合蛋白 4 Lee et al., 2007; Ishii et al., 2013
SSH1 Os07g0235800 AP2转录因子 7 Jiang et al., 2019
), ArticleFig(id=1319334589080912508, tenantId=1146029695717560320, journalId=1316781310003478668, articleId=1319334578385437251, language=CN, label=表1, caption=

水稻中已克隆的落粒性相关基因

, figureFileSmall=null, figureFileBig=null, tableContent=
基因名称 基因编号 编码蛋白 染色体定位 参考文献
qSH1 Os01g0848400 BEL1型同源异型蛋白 1 Konishi et al., 2006
OsSh1 Os03g0650000 YABBY转录因子 3 Lin et al., 2012
ObSH3 OBART03G27740 YABBY转录因子 3 Lv et al., 2018
Sh4/SHA1 Os04g0670900 Myb3转录因子 4 Li et al., 2006; Lin et al., 2007
SHAT1 Os04g0649100 AP2转录因子 4 Zhou et al., 2012
SH5 Os05g0455200 BEL1型同源异型蛋白 5 Yoon et al., 2014
Sh-h/OsCPL1 Os07g0207700 CTD磷酸化酶 7 Ji et al., 2010
OsGRF4/PT2 Os02g0701300 生长调控因子 2 Sun et al., 2016
OsNPC1 Os03g0826600 磷酸酯酶C1 3 Cao et al., 2016
GL4 ORGLA04G0254300 Myb3转录因子 4 Wu et al., 2017
OsLG1 Os04g0656500 SQUAMOSA启动子结合蛋白 4 Lee et al., 2007; Ishii et al., 2013
SSH1 Os07g0235800 AP2转录因子 7 Jiang et al., 2019
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植物器官脱落调控因素及分子机理研究进展
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刘旭鹏 , 王敏 , 韩守安 , 朱学慧 , 王艳蒙 , 潘明启 , 张雯 *
植物学报 | 专题论坛 2025,60(3): 472-482
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植物学报 |专题论坛 2025 , 60 (3) : 472 -482
植物器官脱落调控因素及分子机理研究进展
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刘旭鹏, 王敏, 韩守安, 朱学慧, 王艳蒙, 潘明启, 张雯*
作者信息
  • 新疆农业科学院园艺作物研究所, 农业农村部新疆地区果树科学观测试验站, 新疆特色果蔬基因组研究与遗传改良重点实验室, 乌鲁木齐 830091
通讯作者:
*张雯, 博士, 研究员, 国家葡萄产业技术体系果实品质调控岗位科学家。现任新疆园艺学会理事, 新疆农业大学及石河子大学硕士生导师, 西北农林科技大学校外合作导师。其研究团队主要围绕葡萄品质形成机理和调控技术开展研究与示范推广工作, 经多年攻关, 确定了葡萄裂果、落粒、色泽、酚含量以及香气等重要品质的关键影响因子, 开发了配套调控技术; 构建了西北产区葡萄顺沟倾斜龙干树形及配套机械化出埋土技术、制干葡萄高架少蔓龙干树形改造技术和优质快速制干技术等多项综合技术, 并在我国最大的葡萄产区新疆推广应用, 提高了优质果率, 有效促进了标准化栽培, 显著降低了生产成本。主持国家级和省部级科研项目10余项。发表论文58篇。授权发明和实用新型专利23项。制定地方标准19项。获自治区科技进步二等奖2项, 地州科技进步二等奖和三等奖各1项。E-mail:
Research Progress on Factors and Molecular Mechanisms Regulating Plant Organ Abscission
Liu Xupeng, Wang Min, Han Shou'an, Zhu Xuehui, Wang Yanmeng, Pan Mingqi, Zhang Wen*
Affiliations
  • Key Laboratory of Genome Research and Genetic Improvement of Xinjiang Characteristic Fruits and Vegetables, Scientific Observing and Experimental Station of Pomology (Xinjiang), Ministry of Agriculture and Rural Affair, Institute of Horticultural Crops, Xinjiang Academy of Agricultural Sciences, Urumqi 830091, China
  • Corresponding Author:
出版时间: 2025-05-10 doi: 10.11983/CBB24133
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植物器官脱落是植物体的器官部分从母体脱离的现象, 是植物在生长发育和响应环境变化过程中进化出的一种适应性策略, 以确保植物正常生长和适应环境。该过程涉及离层区形成、特定信号激活以及细胞分离, 同时还受自身生理进程以及光照、温度和湿度等外部环境因素的显著调节。在农业生产实践中, 植物器官脱落直接影响作物的产量。研究植物器官脱落的调控机理对于进一步提高作物产量具有重要意义。近年来, 关于器官脱落机理的研究取得了显著进展。研究表明, 植物器官在脱落机制上具有保守性, 但不同物种间也表现出显著差异。该文深入探讨了植物器官脱落的生理生化机制, 分析了不同环境因素以及激素和酶对其的影响, 旨在为作物遗传育种和农业生产提供理论支撑和实践指导。

器官脱落  /  离层区  /  分子机理  /  酶  /  植物激素

The organs abscission is a phenomenon in which parts of organs are separated from the body. It is an adaptive strategy evolved by plants in the process of growth and response to environmental changes, which ensures the normal growth and adaptation of plants to the environment. This process involves the abscission zone formation, signal activation, cell separation, and is regulated by both internal (in vivo) physiological processes and external environmental factors including light, temperature, and humidity. In agricultural production, the abscission of plant organs directly affects crop yield. Understanding the regulatory mechanisms of plant organ abscission is greatly important for improving crop yield. In recent years, tremendous progress has been made in the study of organ abscission mechanisms. The studies showed that the mechanisms of organ abscission in plants are largely conservative between species, but also with significant variations. This review delved into the physiological and biochemical mechanisms of plant organ abscission, and summarized the effects of both the environmental factors and the hormones and enzymes in the process. The review provides a solid theoretical support and practical guidance for crop genetic breeding and agricultural production improvement in connection with organ abscission.

organ abscission  /  abscission zone  /  molecular mechanism  /  enzyme  /  phytohormone
刘旭鹏, 王敏, 韩守安, 朱学慧, 王艳蒙, 潘明启, 张雯. 植物器官脱落调控因素及分子机理研究进展. 植物学报, 2025 , 60 (3) : 472 -482 . DOI: 10.11983/CBB24133
Liu Xupeng, Wang Min, Han Shou'an, Zhu Xuehui, Wang Yanmeng, Pan Mingqi, Zhang Wen. Research Progress on Factors and Molecular Mechanisms Regulating Plant Organ Abscission[J]. Chinese Bulletin of Botany, 2025 , 60 (3) : 472 -482 . DOI: 10.11983/CBB24133
为了保障新生器官或重要功能器官能够获得充足的水分和养分, 植物会调控不需要或不重要的器官开始脱落, 以适应不同的发育阶段。当器官受损、受感染或面临不利于生长的外界刺激时, 植物也会诱导其脱落, 以适应环境(Patharkar and Walker, 2016)。植物器官的自然脱落对于其自身的生存具有积极意义, 能够延长自身生命周期或促进种族扩散。但对于人类生产活动来说, 果实脱落或种子落粒会产生负面效果, 其中最显著的是作物产量下降(Patharkar and Walker, 2018)。水稻(Oryza sativa)种子脱落会导致在收获过程中产量减少; 葡萄(Vitis vinifera)落粒会影响果实的收获、运输和销售。
器官脱落贯穿于植物生长发育的整个生命周期, 通常发生在植株母体与器官的连接处, 即离层区(abscission zone, AZ), 一般在侧根(Kumpf et al., 2013)、叶基部(Nakano et al., 2012)、花器官(Mao et al., 2000)、果实果柄(Botton et al., 2011)、种子(Zhou et al., 2012)及开裂区(Lewis et al., 2006)等部位。为研究器官脱落现象的形成与内在机理, 科研人员主要针对番茄(Solanum lycopersicum)、水稻及陆地棉 (Gossypium hirsutum)等模式作物, 研究光照、温度和水分等外部环境因素对植物器官脱落的影响。随着研究的深入, 学者们开始聚焦于器官脱落过程中酶活性、激素水平以及相关代谢产物的合成与积累变化, 以揭示其生物学机制, 近几年更是将焦点转向不同物种中基因调控器官离层区脱落的分子机制。利用转录组学等技术对离层区的信号转导途径进行了广泛而细致的研究, 结果表明, 尽管不同植物中离层区信号的响应与调控模式在机制上具有相似性, 但尚未形成一个普遍适用的调控网络(Meir et al., 2010; Botton et al., 2011)。在不同物种中, 虽然已经鉴定出一系列参与调控离层区发育过程的基因, 但每个物种都有其特有的调控基因和调控模式。这些发现为深入理解植物生命活动提供了新视角, 并为未来的研究指明了方向。
本文系统梳理了植物器官脱落的生理生化机制以及调控植物器官脱落的分子机制, 以期为器官脱落研究与农业生产中预防落粒落果提供参考。
器官脱落一般发生在植株母体与器官的连接处, 该位置由于受到脱落信号的诱导, 母体与器官的连接处会形成离层区。离层区一般由多层细胞构成, 导致器官脱离母体的主要原因是离层区的中胶层细胞壁被水解酶溶解, 进而使器官与母体分离(Sexton and Roberts, 1982; Taylor and Whitelaw, 2001; Olsson and Butenko, 2018)。
植物器官脱落包括离层区的形成、脱落信号通路的激活、离层区中层细胞的分离以及分离后离层区的保护层形成4个阶段(图1)。在器官脱落前, 离层区细胞会减少水分吸收, 导致该区域细胞的水势降低, 水分流失使得细胞进一步收缩, 细胞间隙增大, 从而促进器官脱落; 同时, 乙烯(ethylene, ET)和脱落酸(abscisic acid, ABA)等激素在离层区细胞中积累, 激活细胞壁降解酶(如纤维素酶、果胶酶和木葡聚糖酶)的活性, 导致离层区细胞壁成分发生改变, 细胞壁中果胶和半纤维素等物质降解, 细胞间的黏合力下降, 使细胞层之间的连接变松弛, 为器官脱落创造条件。随着离层区细胞壁的分解和水分流失, 该区域的机械强度显著降低, 最终导致断裂。在器官脱落后, 离层区细胞还会形成保护层, 以封闭伤口, 防止病原体侵入和水分过度蒸发, 保护植物不受进一步损害(Patterson, 2001; Patterson et al., 2016; Patharkar and Walker, 2018)。
导致植物器官脱落的环境因素包括光照强度、温度、水分以及营养供应; 内在因素有植物不同时期的生理生化变化等(Wilmowicz et al., 2018)。光照对于植物发育必不可少, 日照的长短会改变植株光合作用, 进而影响植物的生长发育, 并对开花时间产生影响, 或导致某些器官开始脱落。在拟南芥(Arabidopsis thaliana)中, 光照时间延长能够诱导拟南芥提前开花(Pittendrigh and Minis, 1964; Andrés and Coupland, 2012); 而在光照强度降低时, 会导致拟南芥叶片脱落(付欣, 2023)。在水稻中, 光周期核心基因Hd1、Ghd7、DTH8和PRR37能够调节Ehd1-Hd3a/RFT1通路, 进而促进水稻提前抽穗(Zong et al., 2021)。
温度过高或过低均导致植物器官提前脱落。当温度过高时, 植物叶片的光合作用变弱, 碳水化合物合成受抑制, 同时消耗大量能量, 诱导植物器官脱落; 当温度过低时, 植株的正常代谢受到抑制, 导致主动脱落一部分器官(Tudela and Primo-Millo, 1992)。器官脱落也受光照与温度共同调控(Nakano et al., 2014)。对受到环境胁迫的番茄进行研究, 发现由于受到光照时长变短和温度过低的影响, 其花柄离层区细胞提前形成, 进而使花朵脱落(Reichardt et al., 2020)。
当植物受到干旱胁迫时, 一方面由于水分缺失, 植物体内水分平衡出现异常, 诱导脱落酸合成, 并促使离层区脱落信号通路被激活, 进而引发脱落(Iglesias et al., 2003; Florkiewicz et al., 2020); 另一方面, 由于气孔开度减小和叶片叶绿素含量降低, 导致CO2运输与新陈代谢速率降低, 进而影响光合效率, 使植株叶片变小或直接脱落, 最终影响植物的生长发育和产量(Barickman, 2021; Farouk and Al-Ghamdi, 2021; 姜籽竹, 2022)。而当水分过多时, 同样导致植物光合效率降低, 代谢出现异常, 造成叶片枯萎和脱落(叶龙华等, 2012; 吕艳伟等, 2013; 张旭东等, 2016)。在受涝渍胁迫4天后, 番茄和杨树(Populus sp.)的叶片会出现枯萎和脱落(李德明等, 2010; 孙慧等, 2020; 高巍, 2021)。
能量对器官的脱落也起较大作用。当碳水化合物供给不足时, 不同器官之间相互竞争能量, 一些器官由于得不到充足的能量供给导致脱落(Liang et al., 2020)。对石斛(Dendrobium sp.)进行修剪, 剪下来的花枝由于无法补充消耗的营养物质, 其花器官与叶片发生脱离; 当人为施加蔗糖溶液后, 由于营养得到补充, 石斛花枝的叶片和花器官的衰败大幅延缓, 在很大程度上防止了器官脱落(张蝶, 2011; Pattaravayo et al., 2013)。
环境因素对植物器官脱落的影响是一个复杂而精细的过程。光照、温度、水分和营养物质等通过影响酶活性和激素的合成、运输及信号转导等过程, 间接调控植物器官脱落。这些复杂的相互作用有助于植物应对环境变化, 确保其能够适应不利条件, 从而维持正常的生长发育, 保障植物的生存和繁衍。
植物通过酶活性的变化响应光照、温度、水分和营养物质等环境因素, 这种响应机制能够帮助植物适应环境, 维持代谢平衡。在器官脱落过程中, 细胞壁修饰酶的激活, 特别是多聚半乳糖醛酸酶(polygalacturonases, PG)和纤维素酶(cellulase, Cel)的活性增强起重要作用。在植物器官脱落的离层区, 这2种酶均表现出高活性(Sundaresan et al., 2020)。
多聚半乳糖醛酸酶的功能包括分解果胶或果胶酸, 并促进细胞壁的解体, 使果实成熟软化(Yang et al., 2022)。当PG的表达量升高时, 叶片、花和果实等器官的脱落进程加快。在拟南芥中, PG基因家族的QRT2调控花器官脱落, ADPG1参与花药开裂, ADPG2诱导角果开裂(Ogawa et al., 2009)。
纤维素酶是一组复合酶, 又名β-1,4-葡聚糖-4-葡聚糖水解酶, 其成分包括内切β-葡聚糖酶、外切β-葡聚糖酶、β-葡萄糖苷酶和木聚糖酶等, 能够降解纤维素生成葡萄糖。研究表明, Cel编码基因在植物不同器官离层区的表达量差异显著。例如, Cel1主要在花柄离层区和花药中表达, 而Cel2则在果柄中表达(Brummell et al., 1999)。此外, Cel3也被发现在辣椒(Capsicum frutescens)的花器官与叶片离层区高表达(Trainotti et al., 1998; Sakamoto et al., 2008)。在对番茄器官脱落的研究中, 发现Cel1、Cel2和Cel5在果柄离层区大量富集, 而在叶柄离层区仅Cel5大量富集(del Campillo and Bennett, 1996)。
器官脱落是一个复杂且高度连续的过程, 其依赖于多种激素之间的密切配合与协同调节。乙烯作为关键的植物激素之一, 通过促进细胞壁中果胶降解进而诱导器官脱落。脱落酸(ABA)通过提高乙烯前体物质的含量, 促进乙烯生物合成, 进而加速器官脱落。生长素(auxin)和赤霉素(gibberellin, GA)对器官脱落的调控具有浓度依赖性, 根据浓度以及环境条件的不同对器官脱落产生促进或抑制作用。细胞分裂素(cytokinin, CTK)通过促进细胞分裂和延缓植物衰老进程抑制植物器官脱落。目前, 乙烯、ABA、生长素、赤霉素以及细胞分裂素等激素对植物器官脱落的调控作用受到广泛关注。
器官脱落主要是由于离层区中层果胶的损失所致, 乙烯能够促进细胞壁中果胶的降解, 进而促进植物器官脱落。在番茄中, 乙烯诱导PG的表达, 导致叶柄离层区细胞壁中果胶被降解, 最终使番茄果实脱落(Kalaitzis et al., 1995; Hong and Tucker, 1998)。此外, 参与调控乙烯生物合成的基因通过控制乙烯的合成调控器官脱落。乙烯通过调控活性氧(reactive oxygen species, ROS)爆发促使离层区形成。在烟草(Nicotiana tabacum)和番茄中, ERF家族的TERF1与TERF2通过诱导活性氧产生影响植物体内乙烯的生物合成, 最终调控离层区的形成(Zhang et al., 2009, 2016)。
ABA能够触发特定植物器官的脱落。无论是在器官的近轴端还是远轴端, 施加ABA都能促使器官加速脱落。研究表明, ABA能够提高乙烯前体物质ACC合酶的含量, 进而促进乙烯的生物合成, 乙烯促进植物器官离层区细胞壁的降解, 进而使器官脱落(Gómez- Cadenas et al., 2000)。当ABA存在时, PG和Cel在离层区的活性显著增强, 当乙烯的生物合成受到抑制时, 即使ABA仍然存在, PG和Cel的活性也不会增强。说明ABA通过影响乙烯的生物合成促进器官脱落。
生长素调控植物器官脱落的功能是由浓度差所导致。 具体而言, 较高浓度的生长素可促使细胞分化, 而较低浓度的生长素则促进细胞继续扩张与分化。因此, 生长素在植物体内既能促进又能延缓脱落(Bunya-Atichart et al., 2011; Sundaresan et al., 2020)。当生长素的浓度较高时, ATP活性升高, 进而促使细胞壁结构松散, 最终导致植物器官脱落(Kućko et al., 2020)。当生长素浓度较低时, 无法促进ATP活性提升, 且外源生长素具有抑制PG活性的作用, 进而抑制乙烯的生物合成。生长素还能改变降解细胞壁PG的转运机制, 从而延缓器官脱落(韩静等, 1999)。在番茄中, SlBL4基因参与果实叶绿体发育和细胞壁代谢, 且能够影响离层区对生长素浓度的敏感性。SlBL4基因沉默可导致番茄果柄变大及果实提前脱落(Yan et al., 2021)。此外, 根据生长素浓度梯度学说, 植物器官的脱落由近轴端和远轴端生长素浓度决定。当远轴端生长素浓度高于近轴端时, 离层细胞对乙烯信号的响应受到抑制, 导致植物器官不易脱落; 相反, 在近轴端施用生长素可促进离层的形成, 进而加速器官脱落(Addicott, 1970)。综上表明, 生长素在不同位置的浓度梯度对于植物器官的脱落有重要影响。
常见的生长素能够调控植物器官的脱落。例如, 吲哚-3-乙酸(indole-3-acetic acid, IAA)抑制棉花叶片的脱落(Mishra et al., 2008), 其作用机理是IAA可降低乙烯介导的离层区细胞壁降解(van Doorn and Stead, 1997)。1-萘乙酸(1-naphthyl acetic acid, NAA)是人工合成的植物生长调节剂, 具有与IAA类似的功能, 向番茄花器官喷施NAA可以显著减少番茄落果(Alam and Khan, 2002)。2,4-二氯苯氧乙酸(2,4-dichlo-rophenoxyacetic acid, 2,4-D)同样能调控果实脱落。对甜橙(Citrus sinensis)施加低浓度2,4-D能够抑制其果实脱落, 但2,4-D浓度过高会导致甜橙严重落果(陈乃荣和周溅平, 1989)。
不同浓度赤霉素对器官脱落产生促进或抑制作用。与2,4-D相反, 较高浓度的GA通过抑制乙烯的产生或影响乙烯信号转导有效延缓器官的衰老过程, 在甜橙开花期施加一定浓度的GA3能够抑制其果实脱落(Chen et al., 2006)。而GA分布不均匀会增强细胞壁分解酶的活性, 从而导致果实和叶片脱落(Marciniak et al., 2018)。
细胞分裂素具有多重功效, 其不仅具有促进植物细胞分裂的活性, 而且对植物的衰老进程具有延缓作用。CTK通过抑制乙烯的合成或直接激活保护性基因的表达来抑制器官脱落。在水稻中, CTK的高水平表达能够使籽粒变大、穗变长, 并降低落粒率(Sun et al., 2016)。CTK的增加能够促进番茄果实成熟, 并通过提高内源GA和生长素的积累防止落果(Sugiyama and Yamaki, 1995)。
各种激素对植物器官脱落的影响本质上由基因决定。基因对器官脱落起作用主要分为2个方向, 一是通过调控代谢途径中关键酶与激素的合成使其积累量发生变化, 进而影响器官脱落; 二是通过接受酶与激素释放的信号, 对离层区的形成和分化进行调控, 使其生理结构发生改变, 进而导致器官脱离。植物器官脱落相关基因的定位、克隆及其作用的分子机制在品种培育中具有重要意义。目前, 已在不同物种中定位和克隆到一些离层区形成与分化相关基因, 对部分基因的功能也进行了初步研究。
BOP1/2 (BLADE ON PETIOLE 1/2)能够调控拟南芥花器官离层区发育, BOP1/2过表达会延缓花器官的衰老及脱落(McKim et al., 2008)。锌指蛋白(zinc finger protein, ZFP)家族的AtZFP2过表达同样会减少花器官的脱落(Cai and Lashbrook, 2008)。拟南芥果荚开裂也与离层区发育有关, 同源异型盒基因能够以转录因子的方式调控植物的生长发育。在拟南芥中, Homeobox基因家族的RPL基因通过参与乙烯等激素的合成、信号转导或响应基因的表达, 间接调控假隔膜的发育进程, 进而影响种子脱落(Roeder, 2003)。MADS-box家族的AP2 (APETALA2)、SHP1 (Shatterproof1)、SHP2及bHLH家族的IND (INDEHISCENT)和ALC (ALCATRAZ)受到乙烯等激素和环境因素的影响, 在器官脱落的特定时期和特定组织中激活表达, 调控细胞壁降解酶的活性, 导致细胞间连结断裂, 最终影响假隔膜发育, 使拟南芥果荚开裂(Lewis et al., 2006; Ripoll et al., 2011)。
为了方便收获种子, 在水稻人工驯化过程中, 种子是否容易脱落成为非常重要的农艺性状。目前已克隆的水稻落粒相关基因有12个(表1), 其中主效基因7个, 多效基因5个。位于水稻1号染色体上RPL的同源基因qSH1是控制水稻落粒的主效基因之一, 该基因编码一个BEL1型同源异型蛋白。在日本晴中, 由于qSH1基因发生了碱基突变, 使其在小穗基部的表达下调, 进而影响离层区的正常形成, 导致日本晴无法落粒(Konishi et al., 2006)。除qSH1基因外, 位于5号染色体上的同源基因SH5也是控制水稻落粒性的重要基因, 该基因同样编码BEL1型蛋白, 与qSH1基因具有相同的功能(Yoon et al., 2014)。位于水稻4号染色体的sh4基因编码Myb3转录因子, 该转录因子在早期的离层发育中起关键作用, 且能促进种子脱落。而SHA1基因虽然与sh4基因具有高度的相似性, 但SHA1是通过调控离层区的降解而非通过参与离层区的形成影响水稻落粒(Li et al., 2006; Lin et al., 2007)。水稻AP2家族的SHAT1和SSH1基因也是通过调控离层区的降解导致其落粒性下降(Zhou et al., 2012; Jiang et al., 2019)。水稻中编码YABBY因子的OsSh1基因通过影响离层的发育促进种子脱落(Lin et al., 2012)。非洲野生稻(O. glaberrima)中的ObSH3通过调控种子内外颖离层区的发育导致种子脱落(Lv et al., 2018)。
在番茄中, 乙烯生物合成基因ACS2和ACS4在果实成熟过程中高表达, 当ACS2和ACS4失去功能时乙烯合成受阻, 进而抑制花器官脱落(Hoogstrate et al., 2014)。MADS-box蛋白可以与ERFs相互作用,共同对乙烯信号途径下游的基因进行调控, 激活细胞壁降解酶的活性, 进而影响器官脱落。MADS-box家族的J (JOINTLESS)基因能够调控乙烯合成和离层区发育相关基因的表达(Mao et al., 2000; Roberts et al., 2002)。番茄第12号染色体上CPL家族J2 (JOINTLESS2)基因的功能与J基因类似, 不仅能够影响离层区的发育, 也影响花序分生组织的生长(Yang et al., 2005)。J2基因是J基因的下位基因, 其功能受J基因的影响(Roldan et al., 2017)。MADS-box家族的MC (MACROCALYX)是番茄花梗离层区发育所必需的基因, MC通过与J基因互作, 参与调控花梗离层区的发育(Nakano et al., 2012)。SLMBP21基因能够抑制花器官离层区的形成, 其作用机理是通过与J和MC基因互作形成蛋白复合体, 进而影响番茄果柄离层区的发育(Liu et al., 2014)。位于乙烯和生长素信号转导途径下游的SLMYB2转录因子通过调控TAPG1、TAPG2和TAPG4的表达影响番茄果实的脱落(Kalaitzis et al., 1995; Hong and Tucker, 1998; 王晓阳, 2021)。
在小麦(Triticum aestivum)中, qSH1的同源基因TaqSH1直接影响离层区的发育, 将该基因导入拟南芥中, 发现过表达TaqSH1导致拟南芥长角果假隔膜出现异常, 离层区细胞发育迟缓, 说明该基因既影响果实开裂又影响花器官发育(Zhang et al., 2013)。小麦AP2家族的Q基因能够通过调控穗轴离层的发育控制种子脱落(Simons et al., 2006)。在油菜(Brassica campestris)中, RPL基因同样能够导致假隔膜发育异常, 进而调控其落粒性(Arnaud et al., 2011)。在玉米(Zea mays)中, YABBY家族的同源基因ZmSh1也调控其落粒性。在高粱(Sorghum bicolor)中Sh1同样调控落粒(Lin et al., 2012)。柑橘(Citrus sp.)中乙烯生物合成基因CsACS1和CsACO在叶片和果实中高表达, 进而导致叶片和果实脱落(Yuan et al., 2005)。月季(Rosa hybrida)中2个乙烯反应因子(ethylene response factor, ERF)成员RhERF1和RhERF4在花瓣脱落期间能够协调乙烯和生长素信号, 对果胶代谢进行调节, 进而调控花瓣的脱落(Gao et al., 2019)。这些基因通过参与调控乙烯信号途径影响叶片及果实等器官的脱落。
综上所述, 对于植物离层区的发育导致器官脱落的生理机制已研究得较为透彻, 环境因素影响植物器官脱落的机制也比较清晰。相关研究表明, 当外界环境发生变化时, 植物离层区ABA水平升高, 促进乙烯的生物合成, 而乙烯积累量增加上调PG和Cel的表达水平, 进而导致离层区细胞壁发生降解, 最终引发器官脱落。除乙烯调控网络外, 还有其它器官脱落调控网络独立于乙烯途径, 或完全不同于乙烯途径, 或与乙烯途径有重叠, 但对其它调控途径的研究比较缓慢。
目前相关研究主要聚焦于激素和酶对器官脱落的生理调控, 对器官脱落的分子机制研究也主要集中在模式植物, 如水稻的落粒、拟南芥的开花和果荚开裂以及番茄的落花落果, 对于其它物种的研究还十分匮乏。因此, 在不同物种中定位并克隆器官脱落相关基因, 阐明相关基因的功能及构建分子调控网络, 将对揭示器官脱落的机制和作物品种改良具有重要意义, 也对探究植物的起源与进化有所助益。利用器官脱落相关基因培育落粒性适中的品种, 不仅能适应农业机械化, 还能有效克服性状转育过程中易落粒带来的负面影响, 减少生产和运输过程中的损耗, 增加农民收入。
刘旭鹏: 整理文献并撰写论文初稿; 王敏, 韩守安, 朱学慧, 王艳蒙: 整理文献; 潘明启: 提出指导和建议; 张雯: 构思及修改论文。
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doi: 10.11983/CBB24133
  • 接收时间:2024-08-30
  • 首发时间:2026-10-20
  • 出版时间:2025-05-10
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  • 收稿日期:2024-08-30
  • 录用日期:2025-01-20
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新疆农业科学院科技创新稳定支持项目(XJNKYWDZC-2024003-10)
现代农业产业技术体系专项(CARS-29-ZP-08)
天山英才青年拔尖人才项目(2023TSYCCX0022)
天山英才青年拔尖人才项目(2022TSYCCX0068)
乌鲁木齐北疆葡萄种质资源与果园生态新疆野外观测研究站(XJYWZ2307)
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    新疆农业科学院园艺作物研究所, 农业农村部新疆地区果树科学观测试验站, 新疆特色果蔬基因组研究与遗传改良重点实验室, 乌鲁木齐 830091

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*张雯, 博士, 研究员, 国家葡萄产业技术体系果实品质调控岗位科学家。现任新疆园艺学会理事, 新疆农业大学及石河子大学硕士生导师, 西北农林科技大学校外合作导师。其研究团队主要围绕葡萄品质形成机理和调控技术开展研究与示范推广工作, 经多年攻关, 确定了葡萄裂果、落粒、色泽、酚含量以及香气等重要品质的关键影响因子, 开发了配套调控技术; 构建了西北产区葡萄顺沟倾斜龙干树形及配套机械化出埋土技术、制干葡萄高架少蔓龙干树形改造技术和优质快速制干技术等多项综合技术, 并在我国最大的葡萄产区新疆推广应用, 提高了优质果率, 有效促进了标准化栽培, 显著降低了生产成本。主持国家级和省部级科研项目10余项。发表论文58篇。授权发明和实用新型专利23项。制定地方标准19项。获自治区科技进步二等奖2项, 地州科技进步二等奖和三等奖各1项。E-mail:
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2种不同金属材料的力学参数
科
Family
属数
Number of
genus
种数
Number of
species
占总种数比例
Percentage of
total species (%)
属
Genus
种数
Number of
species
占总种数比例
Percentage of total
species (%)
鹅膏菌科Amanitaceae 2 11 5.26 鹅膏菌属 Amanita 10 4.78
小菇科 Mycenaceae 2 12 5.74 丝盖伞属 Inocybe 5 2.39
多孔菌科 Polyporaceae 8 14 6.70 蜡蘑属 Laccaria 5 2.39
红菇科 Russulaceae 3 23 11.00 小皮伞属 Marasmius 6 2.87
小菇属 Mycena 11 5.26
光柄菇属 Pluteus 5 2.39
红菇属 Russula 17 8.13
栓菌属 Trametes 5 2.39
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