光信号和光合产物调控豆科植物根瘤形成发育的研究进展

Research progress on regulation of root nodule formation and development of legume by light signals and photosynthetic products

  • 摘要: 豆科植物与根瘤菌共生形成根瘤, 并通过共生固氮满足植株氮素营养需求。充分发挥豆科固氮能力对减少氮肥消耗及潜在的环境污染、实现农业生产可持续发展有重要意义。根瘤菌与豆科植物共生固氮极其耗能, 因此植株的能量供应调控着共生及固氮过程。本文综述了近些年关于光通过植株的光形态建成和光合作用控制根瘤形成发育的研究进展。植株叶片通过蓝光受体(CRY1s)感知蓝光信号, 并产生从地上移到地下的光受体依赖性转录因子(STFs/FTs), 同时整合根毛发育和根瘤菌诱导的共生信号(NIN)途径形成根瘤, 环境中根瘤菌的运动和附着能力也受光的刺激。植物与根瘤菌形成共生关系后, 植物用光合产物交换根瘤菌同化的NH4+, 光合产物(蔗糖)经韧皮部长距离运输至根瘤, 在根瘤内经糖酵解等途径代谢成有机酸, 为根瘤菌固氮提供碳骨架、能量及还原剂, 光合产物的中间代谢产物如葡萄糖、葡萄糖-6-磷酸、海藻糖-6-磷酸等结合糖信号代谢调控蛋白TOR、SnRK1启动复杂的根瘤皮层细胞生长发育通路。在低氮条件下, 光合产物还以类黄酮、有机酸的形式分泌到根际, 招募根瘤菌与植物形成共生关系。

     

    Abstract: Legumes coexist with rhizobia to form nodules to meet plant nitrogen requirements through symbiotic nitrogen fixation. Legume nitrogen fixation capacity is of great significance to reduce nitrogen fertilizer consumption and potential environmental pollution and achieve sustainable development of agricultural production. The symbiotic nitrogen fixation of legumes consumes a large amount of energy; therefore, the energy supply of plants regulates the process of symbiosis and nitrogen fixation. This study reviews the recent research progress on the control of root nodule formation and development by light through plant photomorphogenesis and photosynthesis. Plant leaves sense blue light signals through blue light receptors (CRY1b) and produce light receptor-dependent transcription factors (STFs/FTs) to move from ground to ground. In addition, they integrate root hair development and the rhizobia-induced symbiotic signal (NIN) pathway to form root nodules. The movement and attachment ability of rhizobia in the environment are stimulated by light. In the symbiosis between plants and rhizobia, plants exchange photosynthetic products to produce NH4+, which is assimilated by rhizobia. In addition, photosynthetic products (sucrose) are transported to root nodules through the phloem for a long distance and are metabolized into organic acids through glycolysis and other pathways in root nodules, providing a carbon skeleton, energy, and reductant for nitrogen fixation of rhizobia. The intermediate metabolites of photosynthates, such as glucose, glucose-6-phosphate, and trehalose 6-phosphate, combine with the regulatory proteins of glucose signal metabolism proteins TOR and SnRK1 to initiate the complex growth and development pathway of nodule cortex cells. Under low-nitrogen conditions, photosynthates are also secreted into the rhizosphere in the form of flavonoids and organic acids, recruiting rhizobia to form symbiotic relationships with plants.

     

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