Article(id=1321790801563206034, tenantId=1146029695717560320, journalId=1316781310003478668, issueId=1321790773536866550, articleNumber=null, orderNo=null, doi=10.11983/CBB25021, pmid=null, cstr=32102.14.CBB25021, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1739203200000, receivedDateStr=2025-02-11, revisedDate=null, revisedDateStr=null, acceptedDate=1746547200000, acceptedDateStr=2025-05-07, onlineDate=1793069084777, onlineDateStr=2026-10-27, pubDate=1752076800000, pubDateStr=2025-07-10, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1793069084777, onlineIssueDateStr=2026-10-27, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1793069084777, creator=13701087609, updateTime=1793069084777, updator=13701087609, issue=Issue{id=1321790773536866550, tenantId=1146029695717560320, journalId=1316781310003478668, year='2025', volume='60', issue='4', pageStart='489', pageEnd='663', issueExtLink='null', onlineDate='null', pubDate='1752076800000', pubDateStr='2025-07-10', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1793069078096, creator='13701087609', updateTime=1793069146158, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1321791059068310443, tenantId=1146029695717560320, journalId=1316781310003478668, issueId=1321790773536866550, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1321791059068310444, tenantId=1146029695717560320, journalId=1316781310003478668, issueId=1321790773536866550, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=489, endPage=498, ext={EN=ArticleExt(id=1321790801898750356, articleId=1321790801563206034, tenantId=1146029695717560320, journalId=1316781310003478668, language=EN, title=“Next-generation Green Revolution” Genes: Toward New “Climate-Smart” Crop Breeding, columnId=1321790801751949715, journalTitle=Chinese Bulletin of Botany, columnName=SPOTLIGHT, runingTitle=null, highlight=null, articleAbstract=

In recent years, significant progresses have been made in plant stress biology, particularly in elucidating the mechanisms underlying responses to extreme temperatures and salinity-alkalinity stresses. These advancements have not only broadened our understanding of plant resilience but also provided a wealth of potential targets for molecular breeding, paving new avenues for developing climate-resilient crop varieties that maintain high yield potential under both optimal and adverse conditions. This review concisely summarizes current knowledge on signal perception and transduction mechanisms during plant adaptation to extreme temperature and saline-alkali stresses, discusses the balance regulation between growth/development and stress tolerance, particularly highlights recent breakthroughs by Chinese scientists in discovering key genes and deciphering mechanisms that synergistically improve crop stress resistance and yield. We also provide future prospects for breeding strategies.

, authors=Liang Ma, Yongqing Yang, Yan Guo, authorsList=Liang Ma, Yongqing Yang, Yan Guo, authorCompany=null, correspAuthors=Yan Guo, authorNote=null, correspAuthorsNote=
*E-mail:
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近年来, 植物逆境生物学领域在极端温度与盐碱胁迫响应机制研究方面取得了重要进展, 不仅拓宽了我们对植物抗逆性的理解, 还为分子育种提供了丰富的靶点, 为培育顺境高产、逆境稳产的“气候智能”作物品种开辟了新途径。该文简要总结了植物感知和转导极端温度与盐碱胁迫信号的分子机制, 讨论了植物生长发育与胁迫耐受的平衡调控, 着重介绍了我国科学家近期在协同提高作物耐逆与产量的关键基因挖掘和机制解析方面的突破性成果, 并提出了未来的育种策略。

, authors=马亮, 杨永青, 郭岩*, authorsList=马亮, 杨永青, 郭岩, authorCompany=null, correspAuthors=郭岩, authorNote=null, correspAuthorsNote=
*郭岩, 博士, 教授, 中国农业大学生物学院、未来技术学院院长。1999年获德国科隆大学遗传学系博士。2009年起在中国农业大学生物学院工作。2010年获得“国家杰出青年科学基金”资助, 同年被聘为教育部“长江学者”特聘教授, 2012年获聘科技部973“作物应答盐碱胁迫的分子调控机理”项目首席科学家, 2013年入选“国家百千万人才工程”。兼任Plant Cell Physiology、Journal of Genetics and Genomics和Journal of Integrative Plant Biology等期刊编委。郭岩教授团队围绕植物盐碱胁迫应答的分子机制及调控网络开展系统性研究工作, 取得了一系列重要研究进展, 先后在Developmental Cell、Nature Communications、Proc Natl Acad Sci USA、Plant Cell、Molecular Plant及EMBO Journal等国际知名期刊发表论文60余篇。E-mail:
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“后绿色革命”基因——助力培育“气候智能”作物新品种
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马亮 , 杨永青 , 郭岩 *
植物学报 | 热点评述 2025,60(4): 489-498
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植物学报 |热点评述 2025 , 60 (4) : 489 -498
“后绿色革命”基因——助力培育“气候智能”作物新品种
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马亮, 杨永青, 郭岩*
作者信息
  • 中国农业大学生物学院, 北京 100193
通讯作者:
*郭岩, 博士, 教授, 中国农业大学生物学院、未来技术学院院长。1999年获德国科隆大学遗传学系博士。2009年起在中国农业大学生物学院工作。2010年获得“国家杰出青年科学基金”资助, 同年被聘为教育部“长江学者”特聘教授, 2012年获聘科技部973“作物应答盐碱胁迫的分子调控机理”项目首席科学家, 2013年入选“国家百千万人才工程”。兼任Plant Cell Physiology、Journal of Genetics and Genomics和Journal of Integrative Plant Biology等期刊编委。郭岩教授团队围绕植物盐碱胁迫应答的分子机制及调控网络开展系统性研究工作, 取得了一系列重要研究进展, 先后在Developmental Cell、Nature Communications、Proc Natl Acad Sci USA、Plant Cell、Molecular Plant及EMBO Journal等国际知名期刊发表论文60余篇。E-mail:
“Next-generation Green Revolution” Genes: Toward New “Climate-Smart” Crop Breeding
Liang Ma, Yongqing Yang, Yan Guo
Affiliations
  • College of Biological Sciences, China Agricultural University, Beijing 100193, China
  • Corresponding Author:
出版时间: 2025-07-10 doi: 10.11983/CBB25021
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近年来, 植物逆境生物学领域在极端温度与盐碱胁迫响应机制研究方面取得了重要进展, 不仅拓宽了我们对植物抗逆性的理解, 还为分子育种提供了丰富的靶点, 为培育顺境高产、逆境稳产的“气候智能”作物品种开辟了新途径。该文简要总结了植物感知和转导极端温度与盐碱胁迫信号的分子机制, 讨论了植物生长发育与胁迫耐受的平衡调控, 着重介绍了我国科学家近期在协同提高作物耐逆与产量的关键基因挖掘和机制解析方面的突破性成果, 并提出了未来的育种策略。

极端温度  /  土壤盐碱化  /  作物抗逆性  /  信号转导  /  气候智能

In recent years, significant progresses have been made in plant stress biology, particularly in elucidating the mechanisms underlying responses to extreme temperatures and salinity-alkalinity stresses. These advancements have not only broadened our understanding of plant resilience but also provided a wealth of potential targets for molecular breeding, paving new avenues for developing climate-resilient crop varieties that maintain high yield potential under both optimal and adverse conditions. This review concisely summarizes current knowledge on signal perception and transduction mechanisms during plant adaptation to extreme temperature and saline-alkali stresses, discusses the balance regulation between growth/development and stress tolerance, particularly highlights recent breakthroughs by Chinese scientists in discovering key genes and deciphering mechanisms that synergistically improve crop stress resistance and yield. We also provide future prospects for breeding strategies.

extreme temperature  /  soil salinization  /  crop resilience  /  signal transduction  /  Climate-Smart
马亮, 杨永青, 郭岩. “后绿色革命”基因——助力培育“气候智能”作物新品种. 植物学报, 2025 , 60 (4) : 489 -498 . DOI: 10.11983/CBB25021
Liang Ma, Yongqing Yang, Yan Guo. “Next-generation Green Revolution” Genes: Toward New “Climate-Smart” Crop Breeding[J]. Chinese Bulletin of Botany, 2025 , 60 (4) : 489 -498 . DOI: 10.11983/CBB25021
全球气候变化造成的局部温度异常、降水模式改变以及土壤盐渍化加剧, 已成为限制植物生长的主要因素, 对全球粮食安全构成了严重威胁(Allen and Ort, 2001; Matsuoka et al., 2002; Wheeler and Von Braun, 2013; Yang and Guo, 2018)。自1900年以来, 全球平均气温已上升1.2°C, 预计在未来10年内仍将持续上升。气温每上升2°C可能导致作物产量下降3%-13% (Wang et al., 2020)。由于气候变化造成的全球变暖以及淡水资源短缺的综合影响, 加剧了土壤水分蒸发, 进一步加速土壤盐碱化。据联合国粮农组织(FAO, 2015; https://www.fao.org/3/i5199e/i5199e.pdf)统计, 目前全球有超过1.0×109 hm2土地受到盐碱化影响, 其中约60%的地区为碱性土壤, 这对全球粮食安全构成严峻挑战。预计到2050年, 全球人口将达到100亿, 而农业生产力必须提高60%才能满足粮食需求(van Dijk et al., 2021)。在全球粮食需求刚性增长与气候变化胁迫持续加剧的双重压力下, 传统作物生产体系已呈现产能-抗逆性双重不足的结构性矛盾, 突破环境适应性育种瓶颈成为农业科技攻关的战略重点。为了确保全球粮食安全并突破育种瓶颈, 科学家亟须深入解析植物应对和适应环境胁迫的分子机制, 培育顺境高产、逆境稳产的“气候智能”作物新品种, 这对于保障气候变化背景下的全球粮食安全至关重要。
作物(如水稻(Oryza sativa)、玉米(Zea mays)、小麦(Triticum aestivum)、番茄(Solanum lycopersicum)和大豆(Glycine max))对环境胁迫表现出高度敏感性, 因而在农业生产中, 盐碱、干旱、涝害以及极端温度等非生物胁迫严重威胁作物产量。植物细胞进化出多层级感应机制以应对上述胁迫, 使其能够在复杂多变的环境中快速适应并存活。定位于质膜的受体作为第一道防线, 能够特异性识别各类环境胁迫信号, 激活第二信使系统, 如Ca2+信号以及ROS (reactive oxygen species)和cAMP (cyclic adenosine mono- phosphate), 最终触发特异性逆境响应以增强植物的胁迫耐受性(Yang and Guo, 2018)。目前, 对植物细胞受体的定义标准包括3个关键因素。(1) 膜定位。质膜是感知外部信号的最初部位, 可以将初级外部物理/化学信号转化为内部生理反应。(2) 诱导膜电位变化。感知外部信号后, 受体可以诱导膜电位变化, 如Ca2+的流入。(3) 触发特异性第二信使并激活下游响应基因。例如, 干旱胁迫可以引发持续的Ca2+峰值, 而盐胁迫通常会诱导快速且短暂的Ca2+振荡。这些Ca2+信号被多种钙依赖性蛋白解码, 如钙调蛋白(calmodulin, CaM)、类钙调蛋白(calmodulin-like proteins, CMLs)、钙调磷酸酶B类蛋白(calcineurin B-like proteins, CBLs)/CBL相互作用蛋白激酶(CBL-inter- acting protein kinases, CIPKs)以及钙依赖性蛋白激酶(calcium-dependent protein kinases, CDPKs)(Ma et al., 2022)。
在水稻中, 基于其膜定位、低温诱导的Ca2+信号和电生理反应以及耐寒表型, 跨膜蛋白COLD1 (chilling- tolerance divergence 1)被鉴定为冷受体。COLD1与拟南芥G蛋白α亚基互作蛋白GTG1/2 (GPCR-type G protein 1/2)具有高度同源性, 能够与水稻G蛋白α亚基RGA1 (rice G protein alpha subunit 1)互作并增强其GTP (guanosine triphosphate)酶活性, 从而通过调控Ca2+信号赋予水稻耐冷性(Ma et al., 2015)。最新研究发现, 膜蛋白COLD6通过与OMS1 (osmotin-like 1)互作感知冷胁迫, 触发2′,3′-cAMP信号级联, 最终增强水稻的耐冷能力(Luo et al., 2024)。质膜定位的COG1 (chilling tolerance in Gengdao/ japonica rice 1)-OsSERL2 (Oryza sativa somatic embryogenesis receptor kinases-like 2)受体激酶复合体感知低温信号并激活丝裂原活化蛋白激酶(mitogen-activated protein kinase, MAPK)级联途径以增强耐冷性(Xia et al., 2023)。Gγ亚基TT2 (thermotolerance 2)通过与转录因子SCT1 (sensing Ca2+ transcription factor 1)和钙调蛋白(CaM)相互作用, 在高温下负调控水稻的耐热性。在热敏感水稻品种中, 高温以TT2依赖性方式诱导Ca2+, 激活CaM-SCT1/ SCT2分子模块, 然后抑制OsWR2 (Wax synthesis regulatory 2)和其它蜡质合成相关基因的表达, 最终降低植株的耐热性。由于TT2突变导致高温诱导的Ca2+水平降低, 高温下蜡质合成增加, 水稻耐热性提高(Kan et al., 2022)。TT3.1也参与温度感知, 细胞膜定位的TT3.1通过改变定位将高温物理信号解码为生物信号传递给叶绿体前体蛋白TT3.2, 并促进其在液泡中降解, 参与调控水稻高温胁迫(Zhang et al., 2022)。研究表明, 水稻钙网蛋白OsCRT3 (calreticulin 3)与蛋白激酶OsCIPK7形成蛋白复合体, 可通过蛋白构象变化感知低温信号, 参与调控冷胁迫诱导的钙信号并增强水稻OsCIPK7的激酶活性, 提高水稻的耐冷性(Zhang et al., 2019; Guo et al., 2023)。
在拟南芥(Arabidopsis thaliana)中, 光敏色素B (phytochrome B, phyB)通过活性形式Pfr向非活性形式Pr转换以响应高温信号, 释放bHLH (basic Helix-Loop-Helix)类转录因子PIF4 (phytochrome interacting factor 4)和PIF7, 从而调控植物的生长发育(Jung et al., 2016; Legris et al., 2016)。蛋白质相变在温度感知中也发挥关键作用: ELF3 (early flowering 3)通过其PrD结构域介导液-液相分离(liquid-liquid phase separation, LLPS), 导致EC (evening complex)解离并促进开花基因表达, 从而引发高温条件下的早花表型(Jung et al., 2020)。最新发现的温度感知转录共调控因子TWA1 (thermo-with ABA- response 1)通过相变在核内形成凝聚体, 与JAM2 (jasmonate-associated myc-like 2)转录因子、TPL (TOPLESS)和TPR (TOPLESS-related)形成抑制复合物, 重编程高温条件下的基因表达网络(Bohn et al., 2024)。此外, mRNA二级结构构象也参与温度感知: PIF7 mRNA的5′UTR区域具有发夹结构, 在高温条件下发生构象改变, 显著提升PIF7蛋白的翻译水平(Chung et al., 2020)。然而, 未来的研究可能需要阐明这些感知复合体如何整合到信号通路中, 包括靶标这些受体创制“气候智能”型作物新种质。
植物响应温度变化的调控网络研究已取得重要进展。拟南芥CBF/DREB1 (C-repeat binding factor/dehydration-responsive element binding protein 1)被确认是低温驯化过程的核心调控因子。低温可快速诱导CBF基因高表达, 进而激活COR (cold regulated)基因的表达以增强植物的抗冻性(Ding et al., 2024)。CBFs的表达水平及其蛋白积累分别受转录因子和翻译后修饰的精确调控。转录因子ICE1/2 (inducer of CBF expression 1/2)及转录激活因子CAMTA (calmodulin-binding transcription activator)通过促进冷胁迫下CBFs的表达增强植物的抗冻性(Doherty et al., 2009; Kim et al., 2013)。蛋白激酶OST1 (open stomata 1)通过磷酸化修饰ICE1、BTF3 (basic transcription factor 3)、BTF3L (BTF3-like protein)、PUB25 (plant U-box 25)和PUB26来调控其蛋白的稳定性, 从而正调控植株的抗冻性。相反, BIN2 (brassinosteroid-insensitive 2)、MPK3 (mitogen-activated protein kinase 3)和MPK6通过磷酸化作用降低ICE1蛋白的稳定性, 负调控拟南芥的抗冻性(Ding et al., 2024)。类受体胞质激酶CRPK1 (cold-responsive protein kinase 1)通过磷酸化并调控14-3-3蛋白的核质分配, 促进CBF蛋白降解, 从而负调控CBF信号通路(Liu et al., 2017)。在高温胁迫响应中, HsfA1s (heat stress transcription factors A1s)被认为是增强植物耐热性调控网络的核心枢纽, 在高温条件下其表达量显著上调。在常温条件下, HsfA1与HSP70 (heat shock protein 70)和HSP90解离后被激活, 进一步通过钙调素结合蛋白激酶CBK3 (calmodulin-binding protein kinase 3)的磷酸化修饰增强其与下游靶基因启动子的结合能力。然而, 细胞周期依赖性激酶CDC2a (cyclin-dependent kinase 2a)介导的HSF1 (heat shock transcription factor 1)磷酸化会抑制其DNA结合活性(Reindl et al., 1997)。已鉴定到的诸多调控元件, 为将来的分子育种提供了丰富的基因资源和靶点。
前期研究多聚焦于模式植物拟南芥。近年来, 对于玉米响应极端温度的分子机制及适应性研究有了一定的进展。玉米起源于热带地区, 对低温十分敏感; 低温严重影响玉米的生长发育和产量。玉米的耐冷性是由多个数量性状位点(quantitative trait locus, QTL)共同控制的复杂性状(Zhang et al., 2020; Zhou et al., 2022)。通过全基因组关联分析(genome-wide association study, GWAS)鉴定出ICE1和HSF21等关键调节因子, 这些转录因子分别通过调控氮代谢和脂质代谢途径增强玉米幼苗期以及萌发期的低温适应能力(Jiang et al., 2022; Gao et al., 2024a, 2024b)。此外, 研究还发现与膜转运系统、Ca2+通道及转录调控相关的多个基因位点与玉米耐冷性显著关联(Zhou et al., 2022)。利用反向遗传学筛选同样鉴定出多个玉米耐冷性相关组分。DREB1作为进化保守的调控因子, 在植物低温响应中发挥核心作用(Yang et al., 2023)。A型响应调节因子ZmRR1 (response regulator 1)和转录因子bZIP68 (basic leucine zipper transcription factor 68)通过DREB1依赖途径调控玉米的耐冷性, 其蛋白稳定性受激酶MPK8动态调控。值得注意的是, ZmRR1和bZIP68的优异等位基因已被证实可显著增强玉米的耐冷性(Zeng et al., 2021; Li et al., 2022)。上述研究在揭示玉米耐冷性的分子调控网络方面取得重要进展, 但对于玉米适应高纬度低温环境的分子遗传机制仍不清楚。
近期, Zeng等(2025)揭示了调控玉米耐冷性的关键基因COOL1 (cold-responsive operation locus 1)。通过全基因组关联分析鉴定出位于第3号染色体上的主效耐冷基因COOL1。COOL1编码一个bHLH转录因子, 负调控玉米的低温胁迫响应。进一步分析COOL1启动子区域, 发现9个与低温胁迫耐受性密切相关的单核苷酸多态性(single nucleotide polymorphisms, SNPs)位点, 并将玉米自交系分为2种单倍型: HapA (Haplotype A)和HapB。HapA型表现出较低的COOL1表达和更强的低温胁迫耐受性, 进一步表明COOL1的表达与低温胁迫耐受性呈负相关。该研究进一步创建了2种单倍型的近等基因型系(near- isogenic lines, NILs): NILHapA和NILHapB。结果表明, NILHapA系具有较低的COOL1表达和更强的低温胁迫耐受性。研究还发现, 4个关键SNPs位于COOL1启动子中的A-box基序。转录因子HY5 (bZIP61)受低温胁迫诱导上调表达, 并且结合到A-box基序, 负调控COOL1的表达, 从而促进HapA型的低温胁迫耐受性。相反, HapB型中A-box基序的破坏阻遏了HY5的结合, 导致COOL1表达升高, 低温胁迫耐受性降低。RNA测序(RNA-seq)和染色质免疫沉淀测序(ChIP- seq)实验进一步表明, COOL1直接调控关键低温胁迫响应基因的表达, 如DREB1/CBF基因和TPS基因, 通过负调控这些基因的表达, COOL1在玉米应对低温胁迫中发挥关键作用。此外, 低温诱导激活的钙依赖性蛋白激酶CPK17转移至细胞核内, 磷酸化并稳定COOL1, 进一步促进COOL1的低温胁迫负调控作用(Zeng et al., 2025)。
为探究耐低温等位基因的地理分布, Zeng等(2025)对1 008个玉米地方品种的COOL1等位基因进行基因分型, 发现COOL1的耐低温等位基因主要分布在高纬度地区品种中。这一发现凸显了这些等位基因在玉米适应寒冷环境中的重要作用。研究还发现, COOL1功能丧失并未影响正常条件下的产量性状, 表明降低COOL1表达有助于增强玉米冷胁迫耐受性但并不影响其它农艺性状。此外, 许多中国主要栽培品种的亲本携带对低温敏感的COOL1等位基因。该研究成果为通过引入有利的COOL1等位基因改良中国玉米的耐冷性提供了新思路, 为培育适应高纬度地区或应对气候变化的耐冷玉米品种提供了宝贵的遗传资源。
盐碱胁迫及其引发的渗透胁迫共同构成限制作物产量的关键逆境因子, 其信号感知过程始于植物细胞对渗透压变化的识别。有研究表明, 甘露醇处理可在数秒内诱导胞质钙离子浓度快速升高(Knight et al., 1997), 说明渗透胁迫受体与钙通道间存在紧密耦合作用。随着渗透胁迫感知与信号转导关键元件的相继发现, 探究其在盐胁迫感知中的潜在功能成为当前的研究重点。
OSCA1 (reduced hyperosmolality-induced [Ca2+]i increase 1)作为定位于质膜的高渗透压门控钙渗透通道, 通过感知渗透压变化引起的细胞膨压改变, 在钙信号转导中发挥核心作用(Yuan et al., 2014)。BON (BONZAI)蛋白作为钙响应型磷脂结合蛋白, 通过调控钙信号、ABA积累、转录重编程及生长发育等过程, 多维度参与植物的渗透胁迫响应(Chen et al., 2020)。鉴于盐胁迫同时引发渗透与离子稳态失衡, OSCA1与BON蛋白可能是植物感知盐的关键元件。最新研究发现, 生物分子凝聚体作为新型调控因子, 在环境胁迫感知中发挥重要作用(Banani et al., 2017)。转录因子SEU (SEUSS)通过形成液态核凝聚体直接感知渗透胁迫, 调控关键抗逆基因的表达, 其固有无序区(intrinsically disordered region, IDR)在体外和体内均响应渗透压引起的细胞体积变化(Wang et al., 2022)。拟南芥DCP5 (Decapping 5)通过构象变化感知胞外高渗环境, 形成渗透胁迫颗粒(DCP5-enriched osmotic stress granules, DOSGs), 实现mRNA与调控蛋白的区室化, 提高植物渗透适应性(Wang et al., 2024)。这些发现系统阐明了相分离机制在细胞渗透胁迫感知中的分子基础。
值得注意的是, NaCl处理而非甘露醇诱导的渗透胁迫可快速激活SOS2 (Salt Overly Sensitive 2)激酶活性(拟南芥钠离子稳态核心调控因子), 表明植物中存在独立于渗透感知的钠离子特异性识别机制(Ma et al., 2019)。细胞表面糖基肌醇磷酸神经酰胺(glycosyl inositol phosphorylceramide, GIPC)鞘脂被鉴定为钠离子感受器, 通过MOCA1 (monocation-induced [Ca2+]i increases 1)依赖途径触发钙振荡(Jiang et al., 2019)。该研究揭示了钠离子直接结合质膜GIPC引起膜去极化, 进而激活钙离子通道产生特异性钙信号, 调控Na+/H+反向转运系统(Jiang et al., 2019)。SOS通路通过质膜锚定膜联蛋白AtANN4反馈调节盐胁迫下特异钙信号的形成, 参与调控植物的盐胁迫耐受性(Ma et al., 2019)。AtANN1被证明具有钙离子通道活性, 参与调控由ROS激活的钙信号产生和转导(Laohavisit et al., 2013)。盐胁迫诱导的钙信号起源于根内皮层与中柱鞘细胞(Kiegle et al., 2000; Tracy et al., 2008), 通过TPC1 (Two Pore Channel 1)离子通道以~400 μm·s-1速度传播(Choi et al., 2014)。研究进一步发现钙信号瞬变幅度与波速表现钠浓度依赖性, 形成钙浓度门控开关机制(Steinhorst et al., 2022), 使植物可以精确解码盐胁迫强度。除钙信号途径外, 植物通过FER (FERONIA )受体激酶与LLG1 (LORELEI (LRE)-LIKE GLYCOSYLPHOSPHATIDYLINOSITOL (GPI)-ANCHORED PROTEIN 1)共受体构成的细胞壁完整性(cell wall integrity, CWI)传感系统, 直接识别盐胁迫引起的细胞壁软化, 激活特异性钙信号(Feng et al., 2018)。该研究揭示了细胞壁组分调控在盐胁迫适应中的关键作用。尽管已有众多渗透及盐胁迫感受器被鉴定, 但其协同调控机制、环境信号交叉调控潜能及作物改良应用仍需深入探究。
近日, Guo等(2025)发现活性赤霉素浓度的精细调控可增强水稻的碱、热胁迫耐受性及提高产量, 并挖掘出优良的天然等位基因, 可进一步提高广泛栽培的“绿色革命”品种的抗逆性和产量特性, 为日益增长的粮食需求提供了有力保障。该研究团队通过大规模鉴定水稻种质耐碱、耐热表型, 最终定位并克隆到2个耐碱、热的QTLs基因ATT1 (alkali-thermotolerance 1)和ATT2。ATT1/Sd1和ATT2/GNP1编码GA-20氧化酶(GA-20 oxidase) (Wu et al., 2016), 均参与调控活性赤霉素的合成。研究发现, 高浓度的活性赤霉素触发SLR1 (slender rice 1)的降解, 导致活性氧(ROS)过度积累, 造成水稻碱、热胁迫的敏感表型; 而当活性赤霉素保持在低水平时, SLR1蛋白积累, 通过SLR1/DELLA-NGR5 (nitrogen-mediated tiller growth response 5)介导的组蛋白H3K27me3修饰, 抑制耐盐碱或耐热基因(如OsNAAT1和OsHsfA2d)的表达, 水稻同样表现出碱、热胁迫敏感表型。这也解释了为什么水稻在活性赤霉素浓度过高或过低时对碱、热胁迫均表现出较高的敏感性。研究进一步发现, 表达ATT2或施用外源赤霉素(GA), 可以将内源活性赤霉素调节到适中水平, 从而平衡H3K27me3修饰水平和ROS浓度, 进而提高水稻的碱、热耐受性和产量(Guo et al., 2025)。进一步在大田测试中发现, 维持体内中等浓度活性赤霉素, 相比于高浓度和低浓度活性赤霉素, 可以在正常大田环境下提高水稻产量; 在碱性土壤条件下种植, 提高ATT2的表达量, 会显著提高半矮秆“绿色革命”水稻品种在碱胁迫下的产量; 在高温环境下, 中等浓度活性赤霉素含量的水稻株系小区产量显著提升, 进一步说明水稻碱、热抗性新基因ATT1/2具有重要的育种价值。尽管传统“绿色革命”基因(如Sd1)通过赤霉素调控实现矮化与增产, 但其环境适应性的不足在气候变化背景下逐渐显现。Guo等(2025)挖掘的ATT2通过动态平衡活性赤霉素水平, 同步优化抗逆性与产量, 突破了“抗逆-高产”性状拮抗的育种瓶颈(Guo et al., 2025)。这一机制革新标志着从“单一性状改良”向“多维性状协同调控”的范式转变, 可视为“后绿色革命”的代表性基因。
非生物胁迫通过抑制细胞分裂和伸长等关键生理过程, 显著限制植物的生长发育。为应对这一挑战, 植物进化出胁迫触发的信号转导机制, 通过主动抑制生长发育来实现生存策略的优化。这种适应性调控本质上是将有限的资源从生长发育向胁迫防御系统重新分配的生态权衡策略。值得注意的是, 当环境条件改善时, 植物可通过负反馈调控迅速解除胁迫应答基因的过度激活, 重启正常生长程序。解析植物生长与胁迫应答网络的动态互作形成的调控枢纽, 可为培育高抗、高产作物提供潜在的基因靶点。水稻OsMADS57通过功能重编程机制实现器官发育与抗逆响应的精准切换: 在正常条件下, 其与OsTB1 (teosinte branched 1)互作抑制D14 (DWARF14)的表达, 促进分蘖形成(Guo et al., 2013); 而在低温胁迫下则通过构象转变激活OsWRKY94防御通路, 同时上调D14抑制分蘖生长, 并且OsMADS57不同磷酸化修饰状态可能是构成其功能转换的分子开关(Chen et al., 2018)。这种时空特异性的调控模式为解析生长-抗逆平衡机制提供了新范式。为了培育高抗高产理想作物, 需要在一定程度上打破这种平衡。非洲稻STH1 (salt tolerance and heading date 1)的发现揭示了打破传统权衡关系的创新机制, 具有里程碑意义。STH1通过协同调控脂肪酸代谢通路与光周期响应网络, 在提高盐胁迫耐受性的同时显著增加穗粒数(Xiang et al., 2022)。AGO2 (argonaute 2)介导的调控网络通过双重作用机制实现抗逆增产: 一方面通过调控BG3 (Big grain 3)的表达优化细胞分裂素的空间分布, 另一方面维持根冠区激素稳态(Yin et al., 2020)。这些发现印证了通过代谢网络与激素信号的重编程实现“抗逆不减产”的可行性。通过构建胁迫响应元件与发育调控网络的智能互作系统, 可实现环境感知与生长调控的实时耦合。例如, 利用合成生物学技术设计胁迫诱导型启动子驱动生长相关基因的表达, 或开发磷酸化修饰敏感的转录调控开关等, 有望解析胁迫记忆与生长发育的跨代调控机制以及表观遗传修饰在平衡调控中的作用。这些突破将为创制“气候智能”型作物提供理论支撑, 推动农业生产的可持续发展。
近期, Lou等(2025)通过创新的环境智能高产稳产育种技术, 成功培育出顺境下高产、逆境中稳产的作物新种质, 为解决高温引发的主要粮食和蔬菜作物显著减产问题提供了全新的农业生产策略。源库关系(source-sink relationship)最早由Mason和Maskell于1928年提出(Mason and Maskell, 1928)。该概念指出, 光合产物从“源”组织(如叶片)输送到“库”组织(如发育中的花、果实、种子和棉纤维)的过程。高温胁迫限制了“源”组织产生的碳同化物向“库”组织的分配, 从而破坏了源库平衡, 可能导致种子或幼果的选择性流产, 最终造成果实和谷类作物的产量下降。在玉米驯化过程中, “选择性流产”作为一种适应自然环境变化的“权衡”策略得以保留。然而, 在当今良好管理的农业生态系统中, 将碳同化物分配与抗逆性解耦, 已成为减轻全球气候变化对作物生产影响的关键。研究发现, 在高温胁迫条件下, 细胞壁蔗糖转化酶(cell-wall invertase, CWIN)表达下调, 果实中蔗糖含量约为正常条件下的8倍, 而葡萄糖和果糖的含量却显著下降。LIN5编码番茄中的CWIN, 其突变体(lin5CR)在整个发育过程中生长受限, 果实和叶片较小, 果实中的蔗糖积累与己糖缺乏, 近似于受到高温胁迫的野生型番茄果实表型, 表明LIN5对高温胁迫响应的碳分配具有重要影响。LIN5过表达株系也表现出类似lin5CR突变体的表型, 说明LIN5的表达量需要被精细调控, 以优化碳分配。在此基础上, Lou等(2025)提出了一种基于气候响应优化碳分配到“库”器官(climate-responsive optimization of carbon par- titioning to sinks, CROCS)的环境智能育种策略。利用自主开发的高效精准引导编辑(prime editing, PE)工具, 向水稻和番茄的CWIN基因启动子中精准敲入10 bp的热激元件(heat-shock element, HSE)。HSE的插入赋予了CWIN基因高温响应上调表达的能力, 从而在高温胁迫下增强碳分配到水稻籽粒和番茄果实中, 显著缓解了高温胁迫下果实的“糖饥饿”。在不同栽培条件下进行的多地点多季节番茄产量测试结果显示, 在顺境下, CROCS策略使番茄产量提高了14%-47%; 在高温胁迫下, 番茄的每单元果实产量较对照组增加了26%-33%, 并且成功挽回了56.4%- 100%的热胁迫导致的果实产量损失。同时, 果实品质指标(如均匀性和糖分含量)较未经改良的对照组也有显著提高。此外, 通过该策略改良的水稻品种在顺境下产量提高了7%-13%, 而在高温条件下则提高了25%, 最多可挽回41%的谷物产量损失(Lou et al., 2025)。
这一创新方法旨在实现顺境下高产, 同时确保胁迫条件下的稳量。CROCS环境智能育种策略整合了一整套快速育种技术, 包括顺式调控元件筛选、靶向位点/基因选择、瞬时表达验证、基因编辑工具优化以及种质产量和性状评估。近年来, 广泛的研究已识别出多个在生物和非生物胁迫响应、营养吸收与利用及共生微生物相互作用中发挥重要作用的顺式调控元件。这些元件能够响应高温、低温、干旱、盐碱、光照不足、病原入侵和营养缺乏等多种胁迫。CROCS策略通过精准引入这些顺式作用元件到靶基因中, 促进了环境抗逆作物的发展, 使其在顺境下保持高产, 逆境下维持产量稳定性。此外, 该研究提供了高效的基因编辑工具和坚实的技术框架, 为植物发育适应机制的基础研究提供了有力支持。
干旱、高温和低温等多重非生物胁迫往往同时发生, 对作物产量产生显著影响。因此, 阐明这些胁迫信号通路之间的协同调控网络, 并应用先进技术开发具有多重胁迫抗性的作物是未来的研究重点。传统育种面临周期长、效率低和遗传多样性窄等限制, 且转基因方法主要用于单基因性状, 这使得复杂性状的品种育成十分困难。育种的本质在于揭示基因型与表型之间的相关性, 当前育种手段从传统方法转向分子设计和智能育种, 从鉴定关键表型开始, 进一步将基因组学、转录组学、代谢组学、表型组学和合成生物学与多样化的种质资源相结合, 同时人工智能提供数据驱动的指导, 以提高精确性和效率, 旨在通过解码复杂性状的全基因组模式和优化多模块组装来开发新一代超级品种(图1)。
依赖基因组编辑技术和独特种质遗传资源进行新的驯化, 是现代智能育种实践发展的基本策略。野生型种质具有强大的胁迫耐受性和遗传多样性, 为改良栽培品种的胁迫抗性提供了宝贵的资源。基因组编辑技术的发展, 特别是通过CRISPR/Cas9 (clustered regularly interspaced short palindromic repeats/CRISPR-associated protein 9)系统、引导编辑和合成生物学, 正在革新作物的耐逆性和产量性状。CRISPR/Cas9系统以及引导编辑工具通过精确编辑耐逆位点, 打破基因连锁并促进新的驯化和大规模位点筛选, 加速了权衡抗逆性与生长发育新位点的发现(图1)。与此同时, 合成生物学基于工程设计原则, 能够构建、重新设计和制造符合实际应用需求的生物模块和系统, 使其成为开发具有增强抗性和高产的作物新品种的关键工具。
马亮: 撰写、修改文章; 杨永青: 参与撰写文章; 郭岩: 指导修改文章。
  • 国家自然科学基金(32000216)
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2025年第60卷第4期
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doi: 10.11983/CBB25021
  • 接收时间:2025-02-11
  • 首发时间:2026-10-27
  • 出版时间:2025-07-10
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  • 收稿日期:2025-02-11
  • 录用日期:2025-05-07
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国家自然科学基金(32000216)
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    中国农业大学生物学院, 北京 100193

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*郭岩, 博士, 教授, 中国农业大学生物学院、未来技术学院院长。1999年获德国科隆大学遗传学系博士。2009年起在中国农业大学生物学院工作。2010年获得“国家杰出青年科学基金”资助, 同年被聘为教育部“长江学者”特聘教授, 2012年获聘科技部973“作物应答盐碱胁迫的分子调控机理”项目首席科学家, 2013年入选“国家百千万人才工程”。兼任Plant Cell Physiology、Journal of Genetics and Genomics和Journal of Integrative Plant Biology等期刊编委。郭岩教授团队围绕植物盐碱胁迫应答的分子机制及调控网络开展系统性研究工作, 取得了一系列重要研究进展, 先后在Developmental Cell、Nature Communications、Proc Natl Acad Sci USA、Plant Cell、Molecular Plant及EMBO Journal等国际知名期刊发表论文60余篇。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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