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Shunan Zhang, Yuyi Zhang, Kangning Li, Ming Yan, Jinfei Zhang, Ming Yu, Shuo Tang, Luyang Wang, Hongye Qu, Le Luo, Wei Xuan, Guohua XuCurrent Biology, 2021, 31(4): 671-683 DOI: 10.1016/j.cub.2020.10.095; 追溯原文......本站官方QQ群:62473826florigen; flowering time; heading date; nitrogen fertilization; nitrogen use efficiency; photoperiod; rice; single-nucleotide polymorphism1 State Key Laboratory of Crop Genetics and Germplasm Enhancement, Key Laboratory of Plant Nutrition and Fertilization in Low-Middle Reaches of the Yangtze River, Ministry of Agriculture, Nanjing Agricultural University, Nanjing 210095, China;
2 Shanghai Agrobiological Gene Center, Shanghai 201106, China
High nitrogen (N) fertilization for maximizing crop yield commonly leads to postponed flowering time (heading date in rice) and ripening, thus affecting resources use efficiency and followed planting time. We found that N-mediated heading date-1 (Nhd1) can directly activate florigen gene OsHd3a in rice. Inactivation of either Nhd1 or OsHd3a results in delay and insensitivity to N supply of flowering time. Knockout of Nhd1 increases N uptake and utilization efficiency at low-to-moderate N level under both short- and long-day field conditions. Increasing glutamine, the product of N assimilation, can upregulate expression of Nhd1, which in turn downregulates OsFd-GOGAT expression and OsFd-GOGAT activity, displaying a Nhd1-controlled negative feedback regulatory pathway of N assimilation. Moreover, N fertilization effect on rice flowering time shows genetically controlled diversity, and single-nucleotide polymorphism in Nhd1 promoter may relate to different responses of flowering time to N application. Nhd1 thus balances flowering time and N use efficiency in addition to photoperiod in rice.
为实现作物产量最大化而进行的高氮施肥通常会导致开花期(水稻抽穗期)和成熟期推迟,从而影响资源利用效率和随后的播种期。我们发现,氮介导的抽穗期1(Nhd1)可以直接激活水稻成花素基因OsHd3a。Nhd1或OsHd3a失活导致抽穗期的延迟和对供氮不敏感。在短日和长日田间条件下,敲除Nhd1可提高低至中等氮水平下的氮吸收和利用效率。增加氮同化产物谷氨酰胺可上调Nhd1的表达,Nhd1又可下调OsFd-GOGAT的表达和OsFd-GOGAT活性,表现出Nhd1控制的氮同化负反馈调节途径。另外,氮肥对水稻开花时间的影响表现出遗传多样性,Nhd1启动子单核苷酸多态性可能与开花时间对施氮反应的不同有关。因此,除了光周期外,Nhd1还平衡了水稻开花时间和氮利用效率。
氮素对水稻抽穗开花的影响十分复杂。贪青晚熟是农作物生产中的一种普遍现象,过量施氮情况下,作物的生育期延长,其产量和品质均会受到不良影响,但背后的机制一直不为人所了解。为了深入研究这一现象,徐国华带领团队对800多个水稻种质材料或品种进行了分析对比实验。结果发现,高氮肥条件下贪青晚熟的材料近60%;高氮肥条件下依旧提前开花的占10%~20%;还有20%~30%的水稻材料不为高氮肥所动,按时开花。谷氨酰胺(Gln)而不是无机铵(NH4+)直接诱导水稻的开花,氮素过少和过多均会抑制开花。通过对水稻花器官形成部位的转录组分析,结合遗传分析,该研究鉴定到一个受Gln影响并能调控开花时间的转录因子,将其命名为“首个氮素介导的抽穗开花因子” (N-mediated heading date-1,Nhd1)。
氮肥是农作物高产优质的重要保障,但是氮肥的大量生产和施用不仅增加能源消耗和生产成本,加剧土壤酸化、水体富营养化和温室气体排放等资源生态环境问题,而且导致作物开花延迟和贪青晚熟,病虫害增加,降低产量和氮肥利用效率。开花时间早晚是衡量作物生育期的关键指标之一,是氮肥及多种环境因素共同作用的结果,对作物产量和品质形成起着至关重要的作用。因此,探究氮素(N)调控开花的网络通路对于明晰植物营养与环境互作,培育花期适宜、高产-稳产-氮肥高效品种均具有十分重要的理论和应用价值。然而,至今为止,对氮素调控植物开花的分子遗传机制所知甚少。
该研究发现,Nhd1通过正向调控,直接激活开花基因OsHd3a,从而调控水稻的抽穗期。Nhd1或OsHd3a失活,在不同日照条件下,水稻开花时间均往后推迟,且开花失去对氮肥的响应,增加氮的累积吸收量。与此同时,Nhd1通过负向调控,抑制氮素同化基因OsFd-GOGAT,降低其酶的活性和改变谷氨酰胺含量。因此,缺失Nhd1增强氮的同化效率,同时提高产量和氮素利用效率。此外,从水稻核心种质资源中,发现了分别为高氮促进开花、不影响开花的不同材料,Nhd1启动子区的自然变异与其受氮素介导的表达调控和花期关联。这些结果表明,在不同供氮和日照条件下,Nhd1具有平衡水稻生育期和氮素效率的功能。
►基因列表◄
昼夜节律调节因子; 生物钟调节因子; MYB转录因子; 氮肥利用率; 氮素介导的抽穗开花因子 OsCCA1; OsLHY; Nhd1
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