植物花分生组织活性的调控机制

doi:10.13560/j.cnki.biotech.bull.1985.2025-0924

摘要/Abstract

摘要：

花是被子植物传代重要的繁殖器官，在演化过程中形成了繁复的多样化性状。花分生组织（floral meristem, FM）中的干细胞为花器官（floral organ）的发育提供了细胞来源，对FM活性的调控直接影响FM的大小和花器官的数目。外在环境信号、植物激素和内源调控因子构成的调控网络决定了花原基的起始、花分生组织的建立与适时终止以及花器官身份特化的过程。FM活性直接影响作物的产量，因此众多作物育种改良均关注FM调控这一过程。本文以模式植物拟南芥为主，系统梳理了生长素信号及下游转录因子调控花原基起始与FM建立的分子机制，“ABCDE”模型决定花器官身份特性的原理，以CLAVATA-WUSCHEL负反馈通路为核心的FM活性维持与抑制机制，以及多级转录因子介导的FM程序性终止通路。同时，本文也总结了单子叶作物（包括水稻与玉米等）花发育以及FM活性调控的最新进展。通过单子叶植物与拟南芥、番茄等双子叶植物间的比较，揭示了“ABCDE”模型对花器官身份的决定作用，以及CLE（CLV3/ENDOSPERM SURROUNDING REGION）小肽通路抑制FM活性在被子植物中的保守性。在此基础上，猜测C类同源异型基因-C2H2/YABBY转录因子模块是调控被子植物FM适时终止的关键因子。此外，本文亦综述了近年来FM活性研究中涉及的新兴技术及其应用场景，特别探讨了快速发展的单细胞（核）转录组、空间转录组以及单细胞多组学测序等技术在多组学时代背景下，克服FM高度异质性与细胞多样性，以高效、深入解析FM活性调控网络的应用潜力。最后展望了FM发育相关研究的未来方向及可能面临的挑战，旨在为深入解析FM活性调控机制与基于FM活性设计的分子育种策略提供理论依据。

关键词: 花, 花分生组织, 花器官原基, 花分生组织活性, WUSCHEL, 植物激素, 单细胞转录组

Abstract:

Flowers, the reproductive organs of angiosperms, have evolved highly diverse forms. The floral meristem (FM) provides the cellular basis for the development of floral organs, and its activity determines the FM size and floral organ numbers. The precise regulatory network including environmental signals, phytohormones, and endogenous regulatory factors determines the initiation of floral primordia, the establishment and timed termination of FM, and floral organ identity specification. Since the regulation of floral development and FM activity directly affects the yield of crops, it has been a hotspot for crop breeding. With the model plant Arabidopsis thaliana, this review systematically summarizes the molecular mechanisms governing floral primordium initiation and FM establishment mediated by auxin signaling and downstream transcription factors. Besides, this review also details the determining role of “ABCDE” model for floral organ identity, the CLAVATA-WUSCHEL negative feedback loop centered FM activity maintenance and suppression, and the timed FM termination pathway orchestrated by multiple transcription factors. This review also summarizes the latest progress in floral development and FM activity regulation in monocotyledonous crops (including rice (Oryza sativa) and maize (Zea mays)).By comparing mechanisms of flower development and FM regulation in monocot crops (e.g., rice, maize) with those in dicots (e.g., Arabidopsis, tomato), this review reveals the determining roles of the “ABCDE” model in floral organ identity, and the conservation of CLE (CLV3/ ENDOSPERM SURROUNDING REGION) peptide pathways in FM suppression across angiosperms. On this basis, we speculate that the C-class homeotic gene-C2H2/YABBY transcription factor module is a key regulator of timed FM termination in angiosperms. The review also covers emerging technologies in FM research, particularly highlighting the application potential of rapidly advancing single-cell (nucleus) transcriptomics, spatial transcriptomics, and single-cell multi-omics sequencing. These technologies overcome the challenges of FM heterogeneity and cellular diversity, enabling efficient and in-depth dissection of FM regulatory networks in the multi-omics era. Finally, we outline future research directions and potential challenges in FM development, aiming to provide a theoretical basis for elucidating FM regulatory mechanisms and designing molecular breeding strategies based on FM activity.

Key words: flower, floral meristem, floral organ, floral meristem activity, WUSCHEL, phytohormones, single cell transcriptomics

王鑫, 陈炜, 孙博. 植物花分生组织活性的调控机制[J]. 生物技术通报, 2026, 42(9): 1-13.

WANG Xin, CHEN Wei, SUN Bo. Regulatory Mechanism of Plant Floral Meristem Activity[J]. Biotechnology Bulletin, 2026, 42(9): 1-13.

图/表 3

图1 拟南芥花原基起始和花分生组织活性建立的基因调控网络植物的花原基起始和分生组织活性的建立是由环境信号、激素和内源基因表达形成的复杂调控网络决定的。在茎尖分生组织周边区（PZ）形成的生长素最大值通过激活下游转录因子MP，启动花原基起始程序。MP及其靶基因LFY通过表观修饰依赖的方式调控众多靶基因表达，激活细胞分裂素信号，从而限定器官边界，确定花身份特性，促进花原基生长，建立FM活性

Fig. 1 Regulatory network for the establishment of floral primordium and floral meristem in ArabidopsisThe establishment of floral primordia initiation and meristem activity is determined by a complex regulatory network formed by environmental signals, hormones, and endogenous gene expression. Auxin maxima formed in the peripheral zone (PZ) of the inflorescence meristem (IM) leads to floral primordia initiation through the activation of downstream transcription factor MP. MP, together with its target gene LFY, regulate the expressions of numerous targeted genes and activate cytokinin signaling in an epigenetic modification-dependent manner, thereby defining floral identities, delimiting organ boundaries, promoting floral primordia growth, and establishing FM activity

图2 花器官决定的“ABCDE”模型示意图A：双子叶植物拟南芥的花器官决定的“ABCDE”模型；从外向内依次为4枚萼片、4枚花瓣、6枚雄蕊和1枚融合的雌蕊；B：单子叶植物水稻的花器官决定的“ABCDE”模型；从外向内依次为1枚外稃、1枚内稃、2枚浆片、6枚雄蕊和1枚雌蕊

Fig. 2 The ABCDE model of floral organ identity specificationA: The ABCDE model of floral organ identity specification in the dicot Arabidopsis thaliana. From the outermost to the innermost whorl, the flower consists of four sepals, four petals, six stamens, and one fused gynoecium. B: The ABCDE model of floral organ identity specification in the monocot rice (Oryza sativa). From the outermost to the innermost whorl, the floret consists of one lemma, one palea, two lodicules, six stamens, and a single pistil

图3 拟南芥花分生组织活性的抑制与适时终止的分子调控机制植物中存在精密的调控网络以抑制过高的FM活性，并在合适的时间终止FM活性以促进雌蕊原基的分化和发育。在花发育早期阶段，对生长素和细胞分裂素等主要植物激素和干细胞决定基因WUS的调控是抑制过量FM活性的主要途径。拟南芥中，MADS结构域蛋白AG一方面从花发育第3期起直接对WUS起到抑制作用，也在花发育第6期，诱导C2H2型锌指转录因子KNU的表达，并由KNU启动表观修饰依赖的转录抑制途径，快速抑制WUS和干细胞标志基因CLV3表达，实现对FM活性的适时终止。ag、knu和knu crc突变体中未终止的FM活性产生额外轮次的花器官[72-74]

Fig. 3 Molecular regulatory mechanisms underlying the inhibition and timely termination of FM termination in ArabidopsisFine-tuned regulatory networks exist in plants to suppress excessive FM activity and to timely terminate FM activity for gynoecium differentiation and development. In the early phases of floral development, phytohormonal regulation (encompassing auxin and cytokinin) and the function of the stem cell identity gene WUS together act as key regulators to constrain hyperactivation of the floral meristem (FM). In Arabidopsis, the MADS-domain protein AG not only could directly repress WUS at floral stage 3, but also activates the C2H2-type zinc finger transcription repressor KNU at floral stage 6. KNU subsequently initiates the epigenetic-dependent transcriptional repression, and rapidly suppresses the expressions of WUS and the stem cell marker gene CLV3, thereby timely terminates the FM activity. Unterminated FM activity in ag, knu, and knu crc mutants produces additional whorls of floral organs [72-74]

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