植物硒生物强化及微生物在其中的应用潜力

doi:10.17521/cjpe.2022.0163

摘要/Abstract

摘要：

硒是人体必需的微量营养元素, 其含量对人体“有益”和“有害”之间的界限非常窄, 通过植物进行生物强化是人体补充硒的安全有效途径。该文综述了植物对硒吸收、转运、代谢的过程。植物主要从土壤中吸收硒酸盐、亚硒酸盐、有机硒, 根系对于不同形态的硒有不同的吸收机制。吸收过程由不同的转运蛋白参与, 吸收后的硒主要以硒酸根离子的形态在植物体内转运, 通过木质部和韧皮部向地上部分运输, 并在各种酶的作用下进行代谢。最终, 根系吸收的硒一部分以有机硒的形式储存在植物体内, 另一部分以硒化物的形式释放到大气中。该文重点关注不同类型根际微生物对植物硒生物强化的影响, 丛枝菌根真菌、外生菌根真菌、根际促生细菌对植物的硒吸收有一定促进作用, 但其内在的作用机制尚不清楚。基于当前的研究现状, 提出了今后的研究重点: 1)植物对硒的吸收过程及其基因调控; 2)微生物对植物硒生物强化的影响机制及应用潜力。

关键词: 硒, 根际微生物, 吸收, 转运, 生物强化

Abstract:

Selenium is an essential micronutrient element for humans. The range between “beneficial” and “harmful” levels of selenium is very narrow, so biofortification through plants is a safe and effective way to supplement selenium. This article reviewed the processes of selenium uptake, transport, and metabolism in plants. Plants mainly absorb selenate, selenite, and organic selenium from soil. The root system has different absorption mechanisms for different forms of selenium, and the absorption process is participated by different transporters. The absorbed selenium is mainly transported in plants in the form of selenate ion, transported to the aboveground through the xylem and the phloem, and metabolized under the action of various enzymes. Finally, part of the selenium absorbed by the roots is stored in the plant as organic selenium, and the other part is released into the atmosphere in the form of selenide. This article also focuses on the effects of different types of rhizosphere microorganisms on plant selenium biofortification. Arbuscular mycorrhizal fungi, ectomycorrhizal fungi, and rhizosphere promoting bacteria can promote the absorption of selenium in plants to a certain extent, but their internal mechanisms are still unclear. Based on the current research status, the future research focuses are put forward: 1) the process of selenium absorption by plants and its gene regulation; 2) the underlying mechanism and application potential of microorganisms on selenium biofortification in plants.

Key words: selenium, rhizosphere microorganism, absorption, transportation, biofortification

李柳, 刘庆华, 尹春英. 植物硒生物强化及微生物在其中的应用潜力. 植物生态学报, 2023, 47(6): 756-769. DOI: 10.17521/cjpe.2022.0163

LI Liu, LIU Qing-Hua, YIN Chun-Ying. Selenium biofortification in plants and application potential of microorganisms in selenium biofortification. Chinese Journal of Plant Ecology, 2023, 47(6): 756-769. DOI: 10.17521/cjpe.2022.0163

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https://www.plant-ecology.com/CN/Y2023/V47/I6/756

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微生物种类 Microbial type	寄主植物 Host plant	微生物名称 Microbes		施肥方式 Fertilization method		硒形态 Selenium form		结论 Result	参考文献 Reference
AMF	玉米 Zea mays	摩西球囊霉 Glomus mosseae		土壤施肥 Soil fertilization		Se(IV) Se(VI)		用Se(IV)处理时, 接种AMF减少了有机硒在玉米中的积累, 用Se(VI)处理时, 接种AMF对玉米中的硒含量无明显影响 When treated with Se(IV), inoculation with AMF reduced organic selenium accumulation in maize, and when treated with Se(VI), inoculation with AMF had no significant effect on selenium content in maize	Yu et al., 2011
	莴苣 Lactuca sativa	根内根孢囊霉和摩西管柄囊霉 Rhizophagus intraradices and Funneliformis mosseae		土壤施肥 Soil fertilization		Se(IV) SeCH₃ SeU		接种AMF减少了硒在莴苣叶片中的积累 Inoculation with AMF reduced selenium accumulation in lettuce leaves	Goicoechea et al., 2015
	小麦 Triticum aestivum	变形球囊霉或摩西管柄囊霉 Glomus versiforme or Funneliformis mosseae		土壤施肥 Soil fertilization		Se(IV) Se(VI) SeMet		接种AMF提高了冬小麦对Se(IV)、Se(VI)吸收, 对SeMet的吸收无影响 Inoculation with AMF increased the absorption of Se(IV) and Se(VI) in winter wheat, but had no effect on the absorption of SeMet	Luo et al., 2019
	石刁柏 Asparagus officinalis	根内根孢囊霉和摩西管柄囊霉 Rhizophagus intraradices and Funneliformis mosseae		叶面施肥 Foliage fertilization		Se(VI)		接种AMF使石刁柏中硒水平提高11.8倍, 提升了叶面施用硒肥的效率 Inoculation with AMF increased the level of selenium in spears of Asparagus officinalis 11.8 times, and significantly improved the efficiency of foliar selenium application	Conversa et al., 2019
	紫花苜蓿 Medicago sativa	摩西球囊霉 Glomus mosseae		土壤施肥 Soil fertilization		Se (IV)		在低磷条件下, 接种AMF促进了苜蓿根系对硒的吸收 Under low phosphorus conditions, inoculation with AMF promoted the uptake of selenium by alfalfa roots	Peng et al., 2020
	水稻 Oryza sativa	变形球囊霉和摩西管柄囊霉 Glomus versiforme and Funneliformis mosseae		土壤施肥 Soil fertilization		Se(IV) Se(VI)		接种AMF可能会改变微生物群落, 将土壤中的无效硒转变为有效硒而提高硒的吸收, 接种AMF能促进水稻籽粒中有机硒的积累 Inoculation with AMF may changethe microbial community, convert unavailable selenium to available selenium in soil and improve selenium absorption. Inoculation with AMF promote organic selenium accumulation in rice grain	Chen et al., 2020
	大蒜 Allium sativum	变形球囊霉 Glomus versiforme		叶面施肥 Foliage fertilization		Se(IV)		在高浓度(50 mg·L^-1)硒含量下, 接种AMF明显降低了植物对硒的吸收和转运 Under high selenium content (50 mg·L^-1), inoculation with AMF significantly reduced the absorption and transport of selenium by plants	Yang et al., 2021
ECM	马尾松 Pinus massoniana	褐环乳牛肝菌 Suillus luteus		-		-		在低重金属污染的土壤中, 马尾松接种ECM后, 其植物中的平均硒含量显著增大 In low level heavy metal pollution soil, the mean selenium content in Pinus massoniana increased significantly after inoculation with ECM	Sun et al., 2020
PGPR	小麦 Triticum aestivum	假单胞菌属R8和寡养单胞菌属B19 Pseudomonas sp. R8 and Stenotrophomonas sp. B19		土壤施肥 Soil fertilization		Se(IV)		接种耐硒细菌促进了硒从小麦的根部到叶片的运输 Inoculation with selenium-tolerant bacteria promoted selenium transport from roots to leaves in wheat	Acu?a et al., 2012
	小麦 Triticum aestivum	蜡样芽孢杆菌-YAP6和地衣芽孢杆菌-YAP7 Bacillus Cereus-YAP6 and Bacillus Licheniformis-YAP7		土壤施肥 Soil fertilization		Se(VI)		接种菌株YAP6和YAP7是一种很好的小麦硒生物强化策略 Inoculation with strains YAP6 and YAP7 is a good strategy for selenium biofortification in wheat	Yasin et al., 2015a
	小麦 Triticum aestivum	Bacillus pichinotyi YAM2		土壤施肥 Soil fertilization		Se(VI)		Bacillus pichinotyi YAM2在硒胁迫和缺硒条件下均能促进小麦生长, 并提高小麦中硒的含量 Bacillus pichinotyi YAM2 promoted the growth of wheat and increased the content of selenium in wheat under both selenium stress and selenium deficiency conditions	Yasin et al., 2015b
	甘蓝 Brassica oleracea	假单胞菌属T5-6-I Pseudomonas sp. strain T5-6-I		培养基施肥 Medium fertilization		Se(IV)		Pseudomonas sp. strain T5-6-I的接种能够提高硒在甘蓝中的积累, 但是并没有影响硒在甘蓝中的存在形态 The inoculation of Pseudomonas sp. strain T5-6-I could increase the accumulation of selenium in cabbage, but did not affect the existence of selenium in cabbage	Lusa et al., 2019
	白菜 Brassica campestris	农杆菌属T3F4 Agrobacterium sp. T3F4		土壤施肥 Soil fertilization		Se⁰ Se(IV)		Agrobacterium sp. T3F4能够将土壤中的0价硒转变为+4价的硒, 从而促进白菜对硒的吸收 Agrobacterium sp. T3F4 could convert 0-valent selenium in soil to +4-valent selenium, thereby promoting the absorption of selenium by cabbage	Zhu et al., 2021
	水稻 Oryza sativa	噬几丁质菌属和睾丸酮丛毛单胞菌 Chitinophaga sp. and Comamonas testosteroni	-		-		Chitinophaga sp.和Comamonas testosteroni均能有效还原Se (IV), 生成硒纳米粒子, 都能提高土壤中的有效硒和水稻植株中的硒含量 Both Chitinophaga sp. and Comamonas testosteroni effectively reduced Se(IV) and generated selenium nanoparticles, both of which can increase the available selenium in soil and the selenium content in rice plants		Huang et al., 2021

微生物种类 Microbial type	寄主植物 Host plant	微生物名称 Microbes		施肥方式 Fertilization method		硒形态 Selenium form		结论 Result	参考文献 Reference
AMF	玉米 Zea mays	摩西球囊霉 Glomus mosseae		土壤施肥 Soil fertilization		Se(IV) Se(VI)		用Se(IV)处理时, 接种AMF减少了有机硒在玉米中的积累, 用Se(VI)处理时, 接种AMF对玉米中的硒含量无明显影响 When treated with Se(IV), inoculation with AMF reduced organic selenium accumulation in maize, and when treated with Se(VI), inoculation with AMF had no significant effect on selenium content in maize	Yu et al., 2011
	莴苣 Lactuca sativa	根内根孢囊霉和摩西管柄囊霉 Rhizophagus intraradices and Funneliformis mosseae		土壤施肥 Soil fertilization		Se(IV) SeCH₃ SeU		接种AMF减少了硒在莴苣叶片中的积累 Inoculation with AMF reduced selenium accumulation in lettuce leaves	Goicoechea et al., 2015
	小麦 Triticum aestivum	变形球囊霉或摩西管柄囊霉 Glomus versiforme or Funneliformis mosseae		土壤施肥 Soil fertilization		Se(IV) Se(VI) SeMet		接种AMF提高了冬小麦对Se(IV)、Se(VI)吸收, 对SeMet的吸收无影响 Inoculation with AMF increased the absorption of Se(IV) and Se(VI) in winter wheat, but had no effect on the absorption of SeMet	Luo et al., 2019
	石刁柏 Asparagus officinalis	根内根孢囊霉和摩西管柄囊霉 Rhizophagus intraradices and Funneliformis mosseae		叶面施肥 Foliage fertilization		Se(VI)		接种AMF使石刁柏中硒水平提高11.8倍, 提升了叶面施用硒肥的效率 Inoculation with AMF increased the level of selenium in spears of Asparagus officinalis 11.8 times, and significantly improved the efficiency of foliar selenium application	Conversa et al., 2019
	紫花苜蓿 Medicago sativa	摩西球囊霉 Glomus mosseae		土壤施肥 Soil fertilization		Se (IV)		在低磷条件下, 接种AMF促进了苜蓿根系对硒的吸收 Under low phosphorus conditions, inoculation with AMF promoted the uptake of selenium by alfalfa roots	Peng et al., 2020
	水稻 Oryza sativa	变形球囊霉和摩西管柄囊霉 Glomus versiforme and Funneliformis mosseae		土壤施肥 Soil fertilization		Se(IV) Se(VI)		接种AMF可能会改变微生物群落, 将土壤中的无效硒转变为有效硒而提高硒的吸收, 接种AMF能促进水稻籽粒中有机硒的积累 Inoculation with AMF may changethe microbial community, convert unavailable selenium to available selenium in soil and improve selenium absorption. Inoculation with AMF promote organic selenium accumulation in rice grain	Chen et al., 2020
	大蒜 Allium sativum	变形球囊霉 Glomus versiforme		叶面施肥 Foliage fertilization		Se(IV)		在高浓度(50 mg·L^-1)硒含量下, 接种AMF明显降低了植物对硒的吸收和转运 Under high selenium content (50 mg·L^-1), inoculation with AMF significantly reduced the absorption and transport of selenium by plants	Yang et al., 2021
ECM	马尾松 Pinus massoniana	褐环乳牛肝菌 Suillus luteus		-		-		在低重金属污染的土壤中, 马尾松接种ECM后, 其植物中的平均硒含量显著增大 In low level heavy metal pollution soil, the mean selenium content in Pinus massoniana increased significantly after inoculation with ECM	Sun et al., 2020
PGPR	小麦 Triticum aestivum	假单胞菌属R8和寡养单胞菌属B19 Pseudomonas sp. R8 and Stenotrophomonas sp. B19		土壤施肥 Soil fertilization		Se(IV)		接种耐硒细菌促进了硒从小麦的根部到叶片的运输 Inoculation with selenium-tolerant bacteria promoted selenium transport from roots to leaves in wheat	Acu?a et al., 2012
	小麦 Triticum aestivum	蜡样芽孢杆菌-YAP6和地衣芽孢杆菌-YAP7 Bacillus Cereus-YAP6 and Bacillus Licheniformis-YAP7		土壤施肥 Soil fertilization		Se(VI)		接种菌株YAP6和YAP7是一种很好的小麦硒生物强化策略 Inoculation with strains YAP6 and YAP7 is a good strategy for selenium biofortification in wheat	Yasin et al., 2015a
	小麦 Triticum aestivum	Bacillus pichinotyi YAM2		土壤施肥 Soil fertilization		Se(VI)		Bacillus pichinotyi YAM2在硒胁迫和缺硒条件下均能促进小麦生长, 并提高小麦中硒的含量 Bacillus pichinotyi YAM2 promoted the growth of wheat and increased the content of selenium in wheat under both selenium stress and selenium deficiency conditions	Yasin et al., 2015b
	甘蓝 Brassica oleracea	假单胞菌属T5-6-I Pseudomonas sp. strain T5-6-I		培养基施肥 Medium fertilization		Se(IV)		Pseudomonas sp. strain T5-6-I的接种能够提高硒在甘蓝中的积累, 但是并没有影响硒在甘蓝中的存在形态 The inoculation of Pseudomonas sp. strain T5-6-I could increase the accumulation of selenium in cabbage, but did not affect the existence of selenium in cabbage	Lusa et al., 2019
	白菜 Brassica campestris	农杆菌属T3F4 Agrobacterium sp. T3F4		土壤施肥 Soil fertilization		Se⁰ Se(IV)		Agrobacterium sp. T3F4能够将土壤中的0价硒转变为+4价的硒, 从而促进白菜对硒的吸收 Agrobacterium sp. T3F4 could convert 0-valent selenium in soil to +4-valent selenium, thereby promoting the absorption of selenium by cabbage	Zhu et al., 2021
	水稻 Oryza sativa	噬几丁质菌属和睾丸酮丛毛单胞菌 Chitinophaga sp. and Comamonas testosteroni	-		-		Chitinophaga sp.和Comamonas testosteroni均能有效还原Se (IV), 生成硒纳米粒子, 都能提高土壤中的有效硒和水稻植株中的硒含量 Both Chitinophaga sp. and Comamonas testosteroni effectively reduced Se(IV) and generated selenium nanoparticles, both of which can increase the available selenium in soil and the selenium content in rice plants		Huang et al., 2021