质体发育调控果实成熟与品质的研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2025-1308

摘要/Abstract

摘要：

果实颜色是影响果实商品价值与品质形成的关键农艺性状，而其色彩差异主要受果实质体发育的影响。质体作为半自主性细胞器，其发育分子机理研究对全面解析果实发育、成熟与品质调控具有重大意义。本文系统综述了质体的分类与转变特点、其在果实内源物质代谢中的功能，以及质体发育与转化调控的分子机制与环境影响因素。质体主要包括原质体、叶绿体、有色体和淀粉体等类型，在果实发育不同阶段可发生相互转化，最受关注的是果实成熟过程中叶绿体向有色体的转变，直接影响果皮色泽与营养成分的积累。而质体内叶绿素、类胡萝卜素等色素的合成与积累受到多层次的调控，包括关键转录因子（如GLK、KNOX、APRR2等）的转录调控、植物激素（如生长素、脱落酸等）的信号传导，以及蛋白质翻译后修饰和质体内RNA编辑等表观遗传机制。此外，光照、温度等环境因素通过影响光信号通路和温度响应基因，调节叶绿体结构与色素代谢，进而影响果实采后品质。迄今，尽管对果实叶绿体发育以及向有色体转变的分子机理有了较深入的研究，但对有色体形成机理的解析仍相对有限，未来研究需进一步整合多组学技术与分子调控网络，解析质体在果实发育与环境适应中的协同作用机制，为果实品质改良与贮藏保鲜提供理论依据。

关键词: 果实质体, 质体转变, 叶绿体发育, 有色体形成, 质体物质代谢, 番茄果实, 植物激素, 环境因子

Abstract:

Fruit color is a key agronomic trait that influences both commercial value and quality formation. The variation in fruit color is primarily governed by plastid development. As semi-autonomous organelles, elucidating the molecular mechanisms underlying plastid development is of great significance for comprehensively understanding fruit development, ripening, and quality regulation. This article systematically reviews the classification and dynamic interconversion of plastids, their functions in endogenous substance metabolism in fruits, and the molecular mechanisms and environmental factors that regulate plastid development and transformation. In general, plastids mainly include proplastids, chloroplasts, chromoplasts, and amyloplasts, which can interconvert during different stages of fruit development. The most notable conversion is the transformation of chloroplasts into chromoplasts during fruit ripening, which directly affects peel coloration and the accumulation of nutritional components. The synthesis and accumulation of pigments within plastids, such as carotenoids and chlorophylls, are regulated at multiple regulation levels. This includes transcriptional regulation by key transcription factors (e.g., GLK, KNOX, APRR2), signal transduction by plant hormones (e.g., auxin, abscisic acid), and epigenetic mechanisms such as post-translational protein modifications and RNA editing within plastids. Furthermore, environmental factors such as light and temperature modulate chloroplast structure and pigment metabolism by affecting light signaling pathways and cold-responsive genes, thereby influencing postharvest fruit quality. Although significant progress has been made in understanding the molecular mechanisms of chloroplast development and their conversion to chromoplasts in fruits, the understanding of chromoplast biogenesis remains relatively limited. In future, multi-omics technologies and molecular regulatory networks should be integrated to decipher the synergistic mechanisms of plastids in fruit development and environmental adaptation, providing a theoretical basis for fruit quality improvement and storage preservation.

Key words: fruit plastid development, plastid interconversion, chloroplast development, chromoplast biogenesis, plastid metabolism, tomato fruit, phytohormones, environmental factors

杜丹, 郭翔, 胡鑫, 潘宇. 质体发育调控果实成熟与品质的研究进展[J]. 生物技术通报, 2026, 42(3): 48-59.

DU Dan, GUO Xiang, HU Xin, PAN Yu. Advances in the Regulatory Mechanisms of Plastid Development on Fruit Ripening and Quality[J]. Biotechnology Bulletin, 2026, 42(3): 48-59.

图/表 3

图1 果实中的质体发育与相互转变情况模式图

Fig. 1 Schematic diagram of plastid development and interconversion in fruits

表1 参与叶绿体发育的转录因子

Table 1 Transcription factors involved in chloroplast development

基因名 Gene name	物种 Species	调控方式 Regulation way	参考文献 Reference
GLK2	苹果、猕猴桃和番茄等	正调控果实质体发育	［29-32］
KNOXs	番茄	正调控叶绿体发育	［15， 36］
SlBEL2	番茄	负调控果实绿肩的形成	［37］
SlBLH7	番茄	负调控果实质体发育	［40］
APRR2s	番茄、黄瓜和辣椒等	正调控果实质体发育	［36， 39］
SlGRAS9	番茄	负调控果实质体叶绿素合成	［18］
SlZHD17	番茄	负调控果实质体叶绿素合成	［18］
SlGRAS38	番茄	正调控果实番茄红素合成	［42］
SlTGA2.2-SRDX	番茄	果实质体数目与结构	［43］
PTOX	番茄、拟南芥	正调控质体结构发育	［44-45］
TAGL1	番茄	正调控质体结构发育	［46］

图2 质体发育与分化过程中的转录调控模式图果实从发育到成熟的过程，也是叶绿体发育到转变为有色体的过程，其间主要是叶绿体的结构转变以及伴随着相关的物质代谢两个方面。其中，SlGLK2与SlAPRR2主要参与果实早期叶绿体发育的调控。SlKN2与SlKN4均能激活SlGLK2与SlAPRR2的表达，而另一个起转录激活作用的因子是BR响应基因BZR1-1D。而SlKN2与SlKN4的表达却分别受到HP1、SlBL4和SlBEL11的抑制。与SlKN2与SlKN4的激活作用不同，SlKN5则通过与SlBLH7互作，并抑制SlAPRR2以及叶绿素合成相关基因SlGUN4和光系统II相关基因SlCAB-1C的转录。此外，生长素也在质体发育中起到重要调控作用。生长素响应因子SlARF4可抑制SlGLK2与SlAPRR2的转录，而SlARF6和SlARF10均可激活GLK1基因的表达，并分别激活CAB与RbcS，以及PORCBP1与CBP2的转录表达。而SlZHD17和SlbHLH95等基因主要参与有色体的发育过程，主要通过激活色素合成基因（如PSY1、PDS等）以及抑制糖分代谢相关基因（如SUS1）的表达参与有色体的发育。尽管如此，SlZHD17却能与SlARF4互作，并抑制SlGLK2的转录。而作为半自主性的细胞器，SlSP1和SlSAD8主要参与Toc蛋白的CHLORAD途径，参与叶绿体到有色体结构的转变，而REC1a则通过参与叶绿体内蛋白翻译的过程参与叶绿体向有色体的转变过程。而果实成熟相关转录因子TAGL1和RIN等则多在于协同有色体色素的积累过程

Fig. 2 Schematic diagram of transcriptional regulation during plastid development and differentiationThe development and maturation of fruit is also the process of chloroplasts developing into chromoplasts, mainly involving two aspects: structural transformation of chloroplasts and related metabolic processes. SlGLK2 and SlAPRR2 are primarily involved in regulating early chloroplast development in fruit. Both SlKN2 and SlKN4 can activate the expressions of SlGLK2 and SlAPRR2, while another transcriptional activator is the BR-responsive gene BZR1-1D. However, the expression of SlKN2 and SlKN4 is inhibited by HP1, SlBL4, and SlBEL11, respectively. Unlike the activation effects of SlKN2 and SlKN4, SlKN5 interacts with SlBLH7 and inhibits the transcription of SlAPRR2, as well as the chlorophyll synthesis-related gene SlGUN4 and the photosystem II-related gene SlCAB-1C. Furthermore, auxin also plays an important regulatory role in plastid development. The auxin response factor SlARF4 can inhibit the transcription of SlGLK2 and SlAPRR2, while SlARF6 and SlARF10 can activate the expression of the GLK1 gene and activate the transcriptional expression of CAB and RbcS, PORCBP1 and CBP2, respectively. Genes such as SlZHD17 and SlbHLH95 are mainly involved in chromoplast development, primarily by activating pigment synthesis genes (such as PSY1 and PDS) and inhibiting the expressions of sugar metabolism-related genes (such as SUS1). Despite this, SlZHD17 can interact with SlARF4 and inhibit the transcription of SlGLK2. As semi-autonomous organelles, SlSP1 and SlSAD8 mainly participate in the CHLORAD pathway of Toc proteins, involved in the transition from chloroplast to chromoplast structure, while REC1a participates in the chloroplast-chromoplast transition by participating in protein translation within chloroplasts. Fruit ripening-related transcription factors such as TAGL1 and RIN are mainly involved in the process of synergistic pigment accumulation in chromoplasts

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