水稻种子发育的分子调控机制

Molecular regulation mechanisms of seed development in rice (Oryza sativa L.)

  • 摘要: 水稻种子发育是一个复杂而精细的过程,从受精到成熟经历多个关键阶段,涉及大量基因的时空表达调控网络和多重激素信号通路的精密协同调控。作为双受精的直接产物,胚和胚乳分别承担着遗传信息传递和营养储备的生物学功能。根据形态学特征,胚发育过程可分为原胚期、球形胚期、盾片期、胚芽鞘期四个阶段;而胚乳发育则经历了游离核期、细胞化期、储藏物质累积期和成熟期四个关键时期。胚和胚乳在不同发育阶段有其独特的基因表达和调控模式。此外,随着表观遗传学深入研究,越来越多的证据表明,种子在正常发育以及环境胁迫下,通过DNA甲基化、组蛋白修饰或非编码RNA等表观遗传调控,动态重塑染色质三维结构,改变基因表达模式,从而调节种子发育的适应性。植物激素在水稻种子发育中也起着重要作用,种子早期发育阶段生长素的极性分布对胚的顶-基轴建立至关重要,生长素与细胞分裂素协同调控细胞分裂与器官分化。本文系统综述了水稻种子的胚和胚乳发育关键基因及其调控机制,同时探讨了种子发育过程中表观遗传调控、以及植物激素的作用。最后指出水稻胚和胚乳发育的分子机制仍有许多未解之谜,并展望未来探索焦点。

     

    Abstract: Rice seed development is a complex and precisely orchestrated process that progresses through multiple critical stages from fertilization to maturity, involving intricate spatiotemporal gene regulatory networks and the coordinated interplay of multiple hormonal signaling pathways. As direct products of double fertilization, the embryo and endosperm fulfill distinct biological roles: the embryo transmits genetic information for progeny propagation, while the endosperm serves as a nutrient reservoir. Morphologically, embryo development is categorized into four stages-proembryo, globular embryo, scutellum formation, and coleoptile emergence-whereas endosperm development sequentially transitions through syncytial nuclear proliferation, cellularization, storage substance accumulation, and maturation. The embryo and endosperm exhibit distinct gene expression profiles and regulatory patterns at different developmental stages. In addition, with the advancing research in epigenetics, an increasing body of evidence demonstrates that under both normal developmental conditions and environmental stress, seeds dynamically reprogram the three-dimensional chromatin structure and alter gene expression patterns via epigenetic mechanisms such as DNA methylation, histone modification, and non-coding RNA regulation, thereby modulating the adaptability of seed development. Phytohormones also play pivotal roles in rice seed development: auxin polar distribution during early embryogenesis is essential for establishing the apical-basal axis, while its synergistic interaction with cytokinin coordinates cell division and organ differentiation. In this review, we conducted a systematic analysis of the key genes and their regulatory mechanisms involved in embryo and endosperm development in rice seeds. Additionally, we explored the epigenetic regulation during seed development as well as the roles of plant hormones. Finally, we highlighted that numerous unresolved questions remain regarding the molecular mechanisms underlying rice embryo and endosperm development, and outlined potential directions for future research.

     

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