荒漠植物内生菌多样性及其增强农作物抗旱和耐盐性的研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2021-1525

摘要/Abstract

摘要：

与内生菌共生是荒漠植物应对极端环境的重要策略。内生菌可促进宿主植物抗旱和耐盐生长，且荒漠植物内生菌可以相似方式促进非宿主农作物的抗逆生长，有望解决极端环境下粮食减产问题。本文综述了近年来荒漠植物来源的内生菌种类及其宿主植物多样性，并从植物-微生物共生互作的生理（激素、代谢物、抗氧化物及渗透势调节）到分子（酶和基因）水平，系统分析了内生菌对宿主及非宿主农作物发挥协同抗旱和耐盐的作用机制。结合本课题组多年从事荒漠植物内生菌的研究进展，在获得大量具有促生抗逆功能的菌株并探明其作用方式的基础上，为荒漠生态植被恢复及农业可持续发展探索备选方案。

关键词: 荒漠植物, 内生菌, 农作物, 抗旱耐盐, 抗逆响应基因

Abstract:

Endophytes symbiosis is an important strategy for desert plant to resist to extreme environments. Endophytes promotes the drought resistance and salt tolerance of host plants，and can also improve the stress resistance of non-host crops in the similar way，which is expected to solve the problem of grain yield reduction in extreme environments. This paper summarized the species of endophytes from desert plants and the diversity of their host plants in recent years. From the physiological level of plant-microbial symbiotic interaction（hormone，metabolite，antioxidant and osmotic potential regulation）to the molecular（enzyme and gene）level，it systematically analyzed the mechanism of synergistic drought resistance and salt tolerance of endophytes on host and non-host crops. Combined with the research progress of desert plant endophytes engaged in by our research group for many years，on the basis of obtaining a large number of strains with growth promoting and stress resistance function and exploring their mode of action，we put forward positive alternative schemes for desert ecological vegetation restoration and agricultural sustainable development.

Key words: desert plant, endophyte, crops, drought and salt tolerance, genes associated with stress

伊帕热·帕尔哈提, 祖力胡玛尔·肉孜, 田永芝, 朱艳蕾, 李远婷, 马晓林. 荒漠植物内生菌多样性及其增强农作物抗旱和耐盐性的研究进展[J]. 生物技术通报, 2022, 38(12): 88-99.

YIPARE·Paerhati , ZULIHUMAER·Rouzi , TIAN Yong-zhi, ZHU Yan-lei, LI Yuan-ting, MA Xiao-lin. Research Progress in Diversity of Endophytes Microbial Communities Isolated from Desert Plants and Their Strengthening Effects on Drought and Salt Tolerance in Crops[J]. Biotechnology Bulletin, 2022, 38(12): 88-99.

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序号No.	促生功能Function of growth promoting	内生菌Endophyte	参与调控的物质Substances involved in regulation				植物来源Plant origin	参考文献Reference
序号No.	促生功能Function of growth promoting	内生菌Endophyte	代谢产物Metabolite	植物类激素Phytohormone	酶类Enzymes	渗透势物质Osmotic potential material	植物来源Plant origin	参考文献Reference
1	增加根长，茎长Increase root length and stem length	短小芽孢杆、克雷伯氏菌 B. pumilus，Klebsiella sp.	总黄酮，总多糖，总皂苷，亚精胺Total flavonoids，total polysaccharides，total saponins，and spermidine	吲哚乙酸 IAA	过氧化氢酶、谷胱甘肽过氧化物酶CAT and GPX		甘草、阿尔哈吉Glycyrrhiza uralensis，and Fisch Alhagi sparsifolia	[10，22]
2	增加生物量Increase biomass	不等弯孢、镰刀、新孢子菌、格孢腔菌、单格孢菌、奇氏新孢霉、新卡玛孢菌 Curvularia inaequali，Fusariumavenaceum，Neocamarosporium sp.，Pleosporales，Ulocladium sp.，Neocamarosporium chichastianum，and Neocamarosporium goegapense			过氧化物酶、苯丙氨酸解氨酶、过氧化氢酶POD，PAL，and CAT	可溶性糖，脯氨酸 Soluble sugar and proline	桑托林丝兰、沙棘、盐地碱蓬、小叶罗汉果，迷迭香、鹰嘴豆、梭梭Ceratocarpus arenarius，Seriphidium santolinum，Suaeda salsa，Eragrostis minor，Seidlitzia rosmarinus，Chick peas，and Sacsaoul	[21，23]
3	固氮 Fix nitrogen	克雷伯氏菌、解磷细菌、芸苔假单胞菌 Klebsiella sp.，Pseudomonas frederickbergensis，and Pseudomonas brassicacearum	亚精胺，胞外多糖Spermidine and exopolysaccharides	吲哚乙酸 IAA	ACC脱氨酶、固氮酶ACC deaminase，and nitrogenase		阿尔哈吉、椰枣、黑果枸杞、多枝柽柳Alhagi sparsifolia，Phoenix dactylifera L.，Lycium ruthenicum，and Tamarix ramosissima	[22，31-32]
4	增加鲜重、干重Increase fresh and dry weight	嗜热真菌、克雷伯氏菌 Thermomyces lanuginosus，Klebsiella sp.	总多糖，总黄酮和抗坏血酸，亚精胺Total polysaccharides，total flavonoids，ascorbic acid，and spermidine	吲哚乙酸 IAA	苯丙氨酸解氨酶、过氧化氢酶、过氧化物酶PAL，CAT，and POD		皱襞植物、阿尔哈吉Cullen plicata Delile，and Alhagi sparsifolia	[22，35]
5	增加根重，茎重Increase root weight and stem weight	短芽孢杆菌属、解磷细菌、芸苔假单胞菌 Brevibacillus sp.，Pseudomonas frederickbergensis，and Pseudomonas brassicacearum	胞外多糖Exopolysaccharides	吲哚乙酸 IAA	纤维素酶、果胶酶、ACC脱氨酶Cellulase，pectase，and ACC deaminase		翅果油树、桂花、椰枣Fagonia mollis Delile，Achillea fragrantissima（Forssk）Sch. Bip.，and Phoenix dactylifera L.	[28，31]
6	增加叶片叶绿素浓度，提高光合作用Increase leaf chlorophyll concentration and improve photosynthesis	奇氏新孢霉、新卡玛孢菌、巨大芽孢杆菌、解淀粉芽孢杆菌、阴沟肠杆菌、肺炎克雷伯菌 Neocamarosporium chichastianum，Neocamarosporium goegapense，Periconia macrospinosa，Bacillus megaterium，B.amylolique faciens，Enterobacter cloacae，and Klebsiella pneumoniae	胞外多糖Exopolysaccharides		苯丙氨酸解氨酶、过氧化氢酶、过氧化物酶、抗坏血酸过氧化物酶、谷胱甘肽过氧化物酶、超氧化歧化酶PAL，CAT，POD，APX，GPX，and SOD	脯氨酸Proline	迷迭香、鹰嘴豆、梭梭、仙人掌Seidlitzia rosmarinus，Chick peas，Sacsaoul，and Euphorbia trigonas	[23，33]
7	提高种子萌发率Improve seed germination rate	芽孢杆菌、葡萄球菌属、假单孢菌属、地衣芽孢杆菌、莫哈韦芽孢杆菌、枯草芽孢杆菌 Bacillus，Staphylococcus，Pseudomonas，B. licheniformis，B. mojavensi，and B. subtilis		吲哚乙酸 IAA		脯氨酸Proline	银砂槐、抱茎独行菜Ammodendron bifolium，and Lepidium perfoliatum L.	[18，24]
8	提高植株磷、氮的含量Increase plant phosphorus and nitrogen content	短芽孢杆菌属 Brevibacillus sp		吲哚乙酸 IAA	纤维素酶、果胶酶、ACC脱氨酶Cellulase，pectase，and ACC deaminase		翅果油树、桂花、椰枣Fagonia mollis Delile，Achillea fragrantissima（Forssk）Sch. Bip.，and Phoenix dactylifera L.	[27]
9	促进植物生长Promote plant growth	泛菌属、深色有隔内生菌、芽孢杆菌（pantoea）的Pantoea alhagi sp.nov，Dark septate endophytes，Bacillus	铁载体，胞外多糖，氨，糖原，蛋白质，多聚磷酸盐，磷Siderophores，exopolysaccharides， ammonia，glycogen，protein，polyphosphate，and phosphorus	吲哚乙酸 IAA	蛋白酶、纤维素酶，果胶酶、木聚糖酶Protease，cellulase，and pectase，pentopan mono	可溶性糖、丙二醛、脯氨酸、钙、钾Soluble sugar， malondiald- ehyde，proline，calcium，and kalium	骆驼刺、裸果木、紫茎泽兰Alhagi sparsifolia Shap，Gymnocarpos przewalskii，and Teucrium polium	[62-64]
10	提高抗干旱能力Improve drought resistance	芽孢杆菌、葡萄球菌属、假单孢菌属、泛菌属、深色有隔内生菌、巨大芽孢杆菌、解淀粉芽孢杆菌、阴沟肠杆菌、肺炎克雷伯菌（Bacillus，Staphylococcus，Pseudomonas，pantoea）的Pantoea alhagi sp.nov，Dark septate endophytes，Bacillus megaterium，B.amylolique faciens，Enterobacter cloacae，Klebsiella pneumoniae	铁载体，胞外多糖，氨，糖原，蛋白质，多聚磷酸盐，磷 Siderophores，exopolysaccharides， ammonia，glycogen，protein，polyphosphate，and phosphorus	吲哚乙酸 IAA	蛋白酶、抗坏血酸过氧化物酶、谷胱甘肽过氧化物酶、过氧化氢酶、超氧化歧化酶Protease，APX，GPX，CAT，and SOD	脯氨酸 Proline	银砂槐、裸果木、骆驼刺、仙人掌Ammodendron bifolium，Gymnocarpos przewalskii，Alhagi sparsifolia Shap，and Euphorbia trigonas	[24，33，62-163]
11	溶磷 Solve phosphorus	解磷细菌、芸苔假单胞菌、克雷伯菌 Pseudomonas frederickbergensis，Pseudomonas brassicacearum，Klebsiella	胞外多糖Exopolysaccharides	吲哚乙酸 IAA	ACC脱氨酶、固氮酶ACC deaminase，and nitrogenase		椰枣、黑果枸杞、多枝柽柳Phoenix dactylifera L.，Lycium ruthenicum，and Tamarix ramosissima	[31-32]
12	形成生物膜 Form biofilms	地衣芽孢杆菌、莫哈韦芽孢杆菌、枯草芽孢杆菌 B.licheniformis，B.mojavensi，B.subtilis					抱茎独行菜Lepidium perfoliatum L.	[18]

序号No.	促生功能Function of growth promoting	内生菌Endophyte	参与调控的物质Substances involved in regulation				植物来源Plant origin	参考文献Reference
序号No.	促生功能Function of growth promoting	内生菌Endophyte	代谢产物Metabolite	植物类激素Phytohormone	酶类Enzymes	渗透势物质Osmotic potential material	植物来源Plant origin	参考文献Reference
1	增加根长，茎长Increase root length and stem length	短小芽孢杆、克雷伯氏菌 B. pumilus，Klebsiella sp.	总黄酮，总多糖，总皂苷，亚精胺Total flavonoids，total polysaccharides，total saponins，and spermidine	吲哚乙酸 IAA	过氧化氢酶、谷胱甘肽过氧化物酶CAT and GPX		甘草、阿尔哈吉Glycyrrhiza uralensis，and Fisch Alhagi sparsifolia	[10，22]
2	增加生物量Increase biomass	不等弯孢、镰刀、新孢子菌、格孢腔菌、单格孢菌、奇氏新孢霉、新卡玛孢菌 Curvularia inaequali，Fusariumavenaceum，Neocamarosporium sp.，Pleosporales，Ulocladium sp.，Neocamarosporium chichastianum，and Neocamarosporium goegapense			过氧化物酶、苯丙氨酸解氨酶、过氧化氢酶POD，PAL，and CAT	可溶性糖，脯氨酸 Soluble sugar and proline	桑托林丝兰、沙棘、盐地碱蓬、小叶罗汉果，迷迭香、鹰嘴豆、梭梭Ceratocarpus arenarius，Seriphidium santolinum，Suaeda salsa，Eragrostis minor，Seidlitzia rosmarinus，Chick peas，and Sacsaoul	[21，23]
3	固氮 Fix nitrogen	克雷伯氏菌、解磷细菌、芸苔假单胞菌 Klebsiella sp.，Pseudomonas frederickbergensis，and Pseudomonas brassicacearum	亚精胺，胞外多糖Spermidine and exopolysaccharides	吲哚乙酸 IAA	ACC脱氨酶、固氮酶ACC deaminase，and nitrogenase		阿尔哈吉、椰枣、黑果枸杞、多枝柽柳Alhagi sparsifolia，Phoenix dactylifera L.，Lycium ruthenicum，and Tamarix ramosissima	[22，31-32]
4	增加鲜重、干重Increase fresh and dry weight	嗜热真菌、克雷伯氏菌 Thermomyces lanuginosus，Klebsiella sp.	总多糖，总黄酮和抗坏血酸，亚精胺Total polysaccharides，total flavonoids，ascorbic acid，and spermidine	吲哚乙酸 IAA	苯丙氨酸解氨酶、过氧化氢酶、过氧化物酶PAL，CAT，and POD		皱襞植物、阿尔哈吉Cullen plicata Delile，and Alhagi sparsifolia	[22，35]
5	增加根重，茎重Increase root weight and stem weight	短芽孢杆菌属、解磷细菌、芸苔假单胞菌 Brevibacillus sp.，Pseudomonas frederickbergensis，and Pseudomonas brassicacearum	胞外多糖Exopolysaccharides	吲哚乙酸 IAA	纤维素酶、果胶酶、ACC脱氨酶Cellulase，pectase，and ACC deaminase		翅果油树、桂花、椰枣Fagonia mollis Delile，Achillea fragrantissima（Forssk）Sch. Bip.，and Phoenix dactylifera L.	[28，31]
6	增加叶片叶绿素浓度，提高光合作用Increase leaf chlorophyll concentration and improve photosynthesis	奇氏新孢霉、新卡玛孢菌、巨大芽孢杆菌、解淀粉芽孢杆菌、阴沟肠杆菌、肺炎克雷伯菌 Neocamarosporium chichastianum，Neocamarosporium goegapense，Periconia macrospinosa，Bacillus megaterium，B.amylolique faciens，Enterobacter cloacae，and Klebsiella pneumoniae	胞外多糖Exopolysaccharides		苯丙氨酸解氨酶、过氧化氢酶、过氧化物酶、抗坏血酸过氧化物酶、谷胱甘肽过氧化物酶、超氧化歧化酶PAL，CAT，POD，APX，GPX，and SOD	脯氨酸Proline	迷迭香、鹰嘴豆、梭梭、仙人掌Seidlitzia rosmarinus，Chick peas，Sacsaoul，and Euphorbia trigonas	[23，33]
7	提高种子萌发率Improve seed germination rate	芽孢杆菌、葡萄球菌属、假单孢菌属、地衣芽孢杆菌、莫哈韦芽孢杆菌、枯草芽孢杆菌 Bacillus，Staphylococcus，Pseudomonas，B. licheniformis，B. mojavensi，and B. subtilis		吲哚乙酸 IAA		脯氨酸Proline	银砂槐、抱茎独行菜Ammodendron bifolium，and Lepidium perfoliatum L.	[18，24]
8	提高植株磷、氮的含量Increase plant phosphorus and nitrogen content	短芽孢杆菌属 Brevibacillus sp		吲哚乙酸 IAA	纤维素酶、果胶酶、ACC脱氨酶Cellulase，pectase，and ACC deaminase		翅果油树、桂花、椰枣Fagonia mollis Delile，Achillea fragrantissima（Forssk）Sch. Bip.，and Phoenix dactylifera L.	[27]
9	促进植物生长Promote plant growth	泛菌属、深色有隔内生菌、芽孢杆菌（pantoea）的Pantoea alhagi sp.nov，Dark septate endophytes，Bacillus	铁载体，胞外多糖，氨，糖原，蛋白质，多聚磷酸盐，磷Siderophores，exopolysaccharides， ammonia，glycogen，protein，polyphosphate，and phosphorus	吲哚乙酸 IAA	蛋白酶、纤维素酶，果胶酶、木聚糖酶Protease，cellulase，and pectase，pentopan mono	可溶性糖、丙二醛、脯氨酸、钙、钾Soluble sugar， malondiald- ehyde，proline，calcium，and kalium	骆驼刺、裸果木、紫茎泽兰Alhagi sparsifolia Shap，Gymnocarpos przewalskii，and Teucrium polium	[62-64]
10	提高抗干旱能力Improve drought resistance	芽孢杆菌、葡萄球菌属、假单孢菌属、泛菌属、深色有隔内生菌、巨大芽孢杆菌、解淀粉芽孢杆菌、阴沟肠杆菌、肺炎克雷伯菌（Bacillus，Staphylococcus，Pseudomonas，pantoea）的Pantoea alhagi sp.nov，Dark septate endophytes，Bacillus megaterium，B.amylolique faciens，Enterobacter cloacae，Klebsiella pneumoniae	铁载体，胞外多糖，氨，糖原，蛋白质，多聚磷酸盐，磷 Siderophores，exopolysaccharides， ammonia，glycogen，protein，polyphosphate，and phosphorus	吲哚乙酸 IAA	蛋白酶、抗坏血酸过氧化物酶、谷胱甘肽过氧化物酶、过氧化氢酶、超氧化歧化酶Protease，APX，GPX，CAT，and SOD	脯氨酸 Proline	银砂槐、裸果木、骆驼刺、仙人掌Ammodendron bifolium，Gymnocarpos przewalskii，Alhagi sparsifolia Shap，and Euphorbia trigonas	[24，33，62-163]
11	溶磷 Solve phosphorus	解磷细菌、芸苔假单胞菌、克雷伯菌 Pseudomonas frederickbergensis，Pseudomonas brassicacearum，Klebsiella	胞外多糖Exopolysaccharides	吲哚乙酸 IAA	ACC脱氨酶、固氮酶ACC deaminase，and nitrogenase		椰枣、黑果枸杞、多枝柽柳Phoenix dactylifera L.，Lycium ruthenicum，and Tamarix ramosissima	[31-32]
12	形成生物膜 Form biofilms	地衣芽孢杆菌、莫哈韦芽孢杆菌、枯草芽孢杆菌 B.licheniformis，B.mojavensi，B.subtilis					抱茎独行菜Lepidium perfoliatum L.	[18]