禾本科植物联合固氮的研究现状及应用前景

胡梦媛; 李雅颖; 葛超荣; 张迎迎; 姚槐应

doi:10.13930/j.cnki.cjea.210317

禾本科植物联合固氮的研究现状及应用前景

doi: 10.13930/j.cnki.cjea.210317

胡梦媛^{1, 2,},
李雅颖²,
葛超荣^1, ,,
张迎迎²,
姚槐应²

1.
武汉工程大学　武汉　430074
2.
中国科学院城市环境研究所　厦门　361021

基金项目: 国家重点研发计划项目(2017YFD0200102)和国家自然科学基金项目(41877051)资助

详细信息

作者简介:
胡梦媛, 主要研究方向为土壤氮素循环。E-mail: hmy.222@foxmail.com

通讯作者:
葛超荣, 主要研究方向为环境微生物。E-mail: chaorongge@wit.edu.cn

中图分类号: S144.5
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出版历程
- 收稿日期: 2021-05-25
- 录用日期: 2021-06-25
- 网络出版日期: 2021-08-21
- 刊出日期: 2021-11-10

Research status and application prospects of combined nitrogen fixation in gramineous plants

1.
Wuhan Institute of Technology, Wuhan 430074, China
2.
Institute of Urban Environment, Chinese Academy of Sciences, Xiamen 361021, China

Funds: The study was supported by the National Key Research and Development Project of China (2017YFD0200102) and the National Natural Science Foundation of China (41877051)

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Corresponding author: E-mail: chaorongge@wit.edu.cn

摘要

摘要: 氮是限制农业生产的最重要因素之一。随着人工固氮技术的发展, 氮肥的施用在提高作物产量、解决人类温饱问题的同时, 导致了土壤板结、酸化、氮素流失及温室气体排放(N₂O)等环境问题。与人工合成氨相比, 生物固氮是一种绿色经济的固氮方式, 其包括共生固氮和非共生(自生固氮及联合固氮)固氮, 且每年固定的氮可占总固定量的50%以上。与共生固氮相比, 非共生固氮存在范围广, 如甘蔗、水稻、玉米和小麦等禾本科作物均能进行非共生固氮(联合固氮)。本文主要从禾本科植物的联合固氮菌种类及其作用机理、固氮活性及调控方式以及联合固氮菌的资源及应用3个方面进行综述, 发现相比较共生固氮而言, 联合固氮菌易受到土著微生物、氮素水平等环境因素影响, 其研究难度更大, 需要筛选纯化更多的联合固氮菌, 为其固氮机制研究提供良好材料; 氮、磷、钼、铁等肥料的适量添加可有效促进固氮菌的固氮效率; 固氮菌不仅可以提高土壤固氮量, 而且有利于植物根系激素调节, 从而增加植物抗病抗逆能力, 促进植物更健康的生长。本文最后对禾本科植物联合固氮的农艺管理措施及固氮菌剂的实际应用方面做了展望, 以期为提高禾本科植物联合固氮效率及推动生物固氮菌在农业生产中的应用提供理论依据。
- 禾本科植物 /
- 联合固氮 /
- 固氮菌种类 /
- 固氮活性 /
- 固氮菌应用
Abstract: Nitrogen is one of the most important factors restricting agricultural production. With the development of artificial nitrogen fixation technology, the application of nitrogen fertilizers can increase crop yields and solve problems related to the fulfilment of the basic human needs of food and clothing. However, it has also caused environmental problems, such as soil compaction, acidification, nitrogen loss, and greenhouse gas emissions (e.g., nitrous oxide, N₂O). Compared with synthetic ammonia, biological nitrogen fixation is a green and economical nitrogen fixation method, which entails symbiotic nitrogen fixation and non-symbiotic nitrogen fixation (autogenous nitrogen fixation and combined nitrogen fixation, respectively). Annually, biologically fixed nitrogen can account for more than 50% of the total fixed amount. Compared with symbiotic nitrogen fixation, non-symbiotic nitrogen fixation exists in many plants, for example, sugarcane, rice, maize, wheat, and other gramineous crops that carry out non-symbiotic nitrogen fixation (combined nitrogen fixation). This article reviewed the species of combined nitrogen-fixing bacteria in gramineous plants and their mechanism of action and nitrogen-fixing activity and regulation methods, as well as the resources and applications of these combined nitrogen-fixing bacteria. Compared with symbiotic nitrogen fixation, combined nitrogen-fixing bacteria are more vulnerable to indigenous microorganisms. Research on combined nitrogen-fixing bacteria is more difficult owing to the influence of environmental factors, such as nitrogen levels. It is necessary to screen and purify more combined nitrogen-fixing bacteria to provide optimum materials for research into the nitrogen fixation mechanism. Appropriate levels of nitrogen, phosphorus, molybdenum, iron, and other fertilizers can promote the nitrogen fixation efficiency of bacteria. Nitrogen-fixing bacteria not only increase the extent of soil nitrogen fixation but also facilitate the regulation of plant root hormones, thereby increasing plant disease resistance and stress resistance, promoting healthier plant growth. Finally, agronomic management measures for combined nitrogen fixation through gramineous plants and the practical application of the nitrogen-fixing bacteria are proposed to provide a theoretical basis for improving the efficiency of combined nitrogen fixation through gramineous plants and to promote the application of the nitrogen-fixing bacteria in agricultural production.
- Gramineae /
- Combined nitrogen fixation /
- Species of nitrogen-fixing bacteria /
- Nitrogen fixation activity /
- Application of nitrogen-fixing bacteria

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图 1 根瘤菌固氮系统

A为固氮酶结构图; B为慢生根瘤菌属在非豆科植物根际固氮方式。Figure A shows the structure of nitrogenase; and Figure B shows the nitrogen fixation mode of Bradyrhizobium spp. in the rhizosphere of non-leguminous plants.

Figure 1. Nitrogen fixation system of rhizobia

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表 1 禾本科植物中联合固氮菌种类

Table 1. Species of associated nitrogen-fixing bacteria in grasses

宿主 Parasitifer	联合固氮菌种类 Combined nitrogen-fixing bacteria species	参考文献 Reference
甘蔗 Sugarcane	伯克霍尔德氏菌属、肠杆菌属、假单胞菌属、泛菌属、寡养单胞菌、芽孢杆菌属 Burkholderia sp., Enterobacter sp., Pseudomonas sp., Pantoea sp., Stenotrophomonas sp., Bacillus sp.	[13]
	克雷伯氏菌属、柠檬酸杆菌属 Klebisiella sp., Citrobacter sp.	[26]
	草螺菌属、固氮螺菌属、葡糖醋杆菌 Herbaspirillum sp., Azospirillum sp., Gluconacetobacter sp.	[27]
	链霉菌属、小双孢菌属、马杜拉放线菌属、不动杆菌属、小单胞菌属、类芽孢杆菌、葡萄球菌、赖氨酸芽孢杆菌属、微球菌 Streptomyces, Microbispora, Actinomadura, Acinetobacter, Micromonospora, Paenibacillus, Staphylococcus, Lysinibacillus, Micrococcus	[14]
水稻 Rice	鞘丝藻属、念珠藻属、蓝丝菌属、固氮螺菌属、慢生根瘤菌属、甲基孢囊菌属、地杆菌属、脱硫叶菌属、脱硫弧菌属 Leptolyngbya, Nostoc, Cyanothece, Azospirillum, Bradyrhizobium sp., Methylocystis, Geobacter, Desulfobulbus, Desulfovibrio	[28]
水稻 Rice	甲基弯曲菌、根瘤菌属 Methylosinus trichosporium, Rhizobium sp.	[29]
玉米 Maize	短小芽孢杆菌、枯草芽孢杆菌、泛菌属、微杆菌属、不动杆菌属、红球菌属、微球菌属、杆菌属、棒形杆菌属、短小杆菌属、肠杆菌属、金黄杆菌属 Bacillus pumilus, Bacillus subtilis, Pantoea, Microbacterium, Acinetobacter, Rhodococcus, Micrococcus, Brachybacterium, Clavibacter, Curtobacterium, Enterobacter, Chryseobacterium	[23]
小麦 Wheat	慢生根瘤菌属、伯克霍尔德氏菌属、地杆菌属、脱硫杆菌属、太阳杆菌属、磁螺菌属、甲基球菌属、固氮弧菌属、解纤维素菌属 Bradyrhizobium, Burkholderia, Geobacter, Desulfobacter, Heliobacterium, Magnetospirillum, Methylococcus, Azoarcus, Cellulosilyticum	[21]

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表 2 施加固氮菌对禾本科植物固氮酶活性或产量的影响

Table 2. Effects of nitrogen-fixing bacteria on nitrogenase activity or yield of Poaceae plants

植株 Plant	施加菌株 Bacteria	固氮酶活性提高率 Increase rate of nitrogenase activity (%)	作物产量提高率 Increase rate of crop yield (%)	参考文献 Reference
甘蔗 Sugarcane	巨大芽孢杆菌、蕈状芽孢杆菌 Bacillus megaterium, Bacillus mycoides	21~35	—	[60]
甘蔗 Sugarcane	枯草芽孢杆菌B9 Bacillus subtilis B9	—	29.84~95.51	[61]
水稻 Rice	固氮鱼腥藻 Anabaena azotica (FACHB-119)	—	25	[62]
水稻 Rice	固氮菌 Azotobacter sp. strain Avi2 (MCC 3432)	—	6.3~10.7	[63]
玉米 Maize	多粘芽孢杆菌 Paenibacillus triticisoli BJ-18	12.9~36.4	—	[65]
	固氮螺菌属 Azospirillum brasilense Ab-V5和Ab-V6	—	27	[66]
	多粘芽孢杆菌 Paenibacillus triticisoli BJ-18	—	16.9	[36]
小麦 Wheat	溶磷菌、内生固氮菌、假单胞菌 Paenibacillus sp. B1, Klebsiella, Pseudomonas	26~163	—	[56]
小麦 Wheat	巴西固氮螺菌Ab-V5和Ab-V6 Azospirillum brasilense Ab-V5 and Ab-V6	—	31	[66]

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