植物学报 ›› 2020, Vol. 55 ›› Issue (5): 623-633.DOI: 10.11983/CBB20034
张卫勤1, 邹杭2,3, 张妮娜1, 林雪媛1, 陈娟1,2,*(
)
首页收稿日期:2020-03-03
接受日期:2020-06-05
出版日期:2020-09-01
发布日期:2020-09-03
通讯作者:
陈娟
作者简介:E-mail: chenjuan@nwsuaf.edu.cn基金资助:
Weiqin Zhang1, Hang Zou2,3, Nina Zhang1, Xueyuan Lin1, Juan Chen1,2,*()
Received:2020-03-03
Accepted:2020-06-05
Online:2020-09-01
Published:2020-09-03
Contact:
Juan Chen
摘要： 豆科植物-根瘤菌共生过程受双方基因复杂且精细的调控, 能够产生特异的根瘤结构并可将大气中的惰性氮气(N2)转化为可被植物直接利用的氨态氮。结瘤与固氮受多种因素影响, 其中, 一氧化氮(NO)作为一种自由基反应性气体信号分子, 可参与调节植物的许多生长发育过程, 如植物的呼吸、光形态建成、种子萌发、组织和器官发育、衰老以及响应各种生物及非生物胁迫。在豆科植物中, NO不仅影响寄主与菌共生关系的建立, 还参与调控根瘤菌对氮气的固定并提高植株氮素营养利用效率。该文主要从豆科植物及共生菌内NO的产生、降解及其对结瘤、共生固氮的影响和对环境胁迫的响应, 阐述了NO调控豆科植物共生体系中根瘤形成和共生固氮过程的作用机制, 展望了NO信号分子在豆科植物共生固氮体系中的研究前景。
张卫勤, 邹杭, 张妮娜, 林雪媛, 陈娟. 一氧化氮对豆科植物结瘤及固氮的影响机制. 植物学报, 2020, 55(5): 623-633.
Weiqin Zhang, Hang Zou, Nina Zhang, Xueyuan Lin, Juan Chen. Influence Mechanisms of Nitric Oxide on Nodulation and Nitrogen Fixation in Legumes. Chinese Bulletin of Botany, 2020, 55(5): 623-633.
图1 共生体系中NO的产生与降解示意图(改自Hichri et al., 2016a) 图中包含上下2部分, 分别对应植物和共生体NO的产生与降解。其中, 七角星图示代表氧化途径, 椭圆形图示代表还原途径。实线表示已有研究证实, 虚线表示还有待考证。ETC: 线粒体电子传递链; GSNOR: 亚硝基谷胱甘肽还原酶; Hmp: 黄素血红蛋白; Lb: 豆血红蛋白; NnrS: 含血红素和铜的膜蛋白; Nor: NO还原酶; NOS: NO合酶; ns-Hb: 非共生血红蛋白; NR: 硝酸还原酶; PAOX: 多胺氧化酶; sd-Hb: 单域血红蛋白; Tr-Hb: 截短血红蛋白; TrxR: 硫氧还蛋白还原酶; XOR: 黄嘌呤氧化还原酶
Figure 1 The schematic diagram of NO production and degradation in the symbiotic system (modified from Hichri et al., 2016a) The figure contains the upper and lower parts, which correspond to the production and degradation of NO from plants and symbiotes, respectively. The seven horns star diagrams refer to the oxidation pathway and the oval diagrams refer to the reduction pathway. The lines indicate that studies have been confirmed, and the dashed lines indicate that it is yet to be studied. ETC: Mitochondrial electron transport chain; GSNOR: S-nitrosoglutathione reductase; Hmp: Flavin hemoglobin; Lb: Leghemoglobin; NnrS: Haem- and copper-con- taining membrane protein; Nor: NO reductase; NOS: NO synthase; ns-Hb: Nonsymbiotic hemoglobin; NR: Nitrate reductase; PAOX: Polyamine oxidase; sd-Hb: Single domain hemoglobin; Tr-Hb: Truncated hemoglobin; TrxR: Thioredoxin reduction enzymes; XOR: Xanthine oxidoreductase
图2 NO在共生固氮中的作用示意图(改自Boscari et al., 2013; Hichri et al., 2015, 2016b) 一方面, NO抑制固氮和C、N代谢; 另一方面, NO调控细胞氧化还原和保持低氧水平下的能量状态。带+的细线表示NO的活化、诱导和保持效果; 带有-的细线表示NO的抑制作用。椭圆形粗线箭头表示NO主要的代谢途径。爆炸型图示指来自植物和菌共生体的酶, 闪电型图示表示根瘤菌内的基因。ACO: 乌头酸; CS: 柠檬酸合酶; Gln: 谷氨酰胺; Glu: 谷氨酸; GS: 谷氨酰胺合成酶; GSH: 谷胱甘肽; GSHS: 谷胱甘肽合成酶; GSNO: S-亚硝基谷胱甘肽; Hb: 血红蛋白; IDH: 异柠檬酸脱氢酶; MDH: 苹果酸脱氢酶; NH4+: 铵根离子; Nif: 固氮酶; SDH: 琥珀酸脱氢酶; γ-EC: γ-谷氨酰半胱氨酸; γ-ECS: γ-谷氨酰半胱氨酸合成酶
Figure 2 Schematic diagram of the role of NO in symbiotic nitrogen fixation (modified from Boscari et al., 2013; Hichri et al., 2015, 2016b) On the one hand, NO inhibits nitrogen fixation and carbon and nitrogen metabolism; on the other hand, it regulates cellular redox status and maintains the energy state under low oxygen levels. A thin line with + indicate the activation, induction, and retention effects of NO; a thin line with - indicate the inhibition of NO. The oval thick line arrows indicate the main metabolic pathways of NO. Explosive type diagrams refer to enzymes from plants and bacterial symbionts, and lightning type diagram represents genes within rhizobium. ACO: Aconitic acid; CS: Citrate synthase; Gln: Glutamine; Glu: Glutamic acid; GS: Glutamine synthetase; GSH: Glutathione; GSHS: Glutathione synthetase; GSNO: S-nitrosoglu- tathione; Hb: Hemoglobin; IDH: Isocitrate dehydrogenase; MDH: Malate dehydrogenase; NH4+: Ammonium ion; Nif: Nitrogenase; SDH: Succinate dehydrogenase; γ-EC: γ-glu- tamylcysteine; γ-ECS: γ-glutamyl cysteine synthetase
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