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Article(id=1250834201700942606, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1250834186500784538, articleNumber=null, orderNo=null, doi=10.13343/j.cnki.wsxb.20251017, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1767110400000, receivedDateStr=2025-12-31, revisedDate=null, revisedDateStr=null, acceptedDate=1770566400000, acceptedDateStr=2026-02-09, onlineDate=1776151713061, onlineDateStr=2026-04-14, pubDate=1775232000000, pubDateStr=2026-04-04, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1776151713061, onlineIssueDateStr=2026-04-14, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1776151713061, creator=13701087609, updateTime=1776151713061, updator=13701087609, issue=Issue{id=1250834186500784538, tenantId=1146029695717560320, journalId=1192105938417971205, year='2026', volume='66', issue='4', pageStart='1471', pageEnd='2021', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1776151709437, creator=13701087609, updateTime=1776152261216, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1250836500921922256, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1250834186500784538, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1250836500926116561, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1250834186500784538, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=1929, endPage=1940, ext={EN=ArticleExt(id=1250834202493666118, articleId=1250834201700942606, tenantId=1146029695717560320, journalId=1192105938417971205, language=EN, title=Isolation of plant growth-promoting rhizobacteria and arbuscular mycorrhizal fungi from the rhizosphere of Bupleurum chinense and characterization of their synergistic effects on plant growth, columnId=1192149543992045670, journalTitle=Acta Microbiologica Sinica, columnName=Research Article, runingTitle=null, highlight=null, articleAbstract=

Objective The rhizosphere microbial community plays a critical role in plant growth, development, and quality formation. Therefore, this study systematically isolated plant growth-promoting microbial resources from the rhizosphere of Bupleurum chinense and evaluated their application potential, aiming to provide excellent strains for the development of microbial fertilizers to reduce the use of chemical fertilizers and pesticides. Methods Plant growth-promoting rhizobacteria (PGPR) and arbuscular mycorrhizal fungi (AMF) were isolated and identified from the rhizosphere of B. chinense by the culture-dependent methods. Functional traits of PGPR strains were screened through in vitro assays, and the synergistic growth-promoting effects of PGPR and AMF were subsequently evaluated by a pot experiment. Results A total of 25 PGPR species and 2 AMF species (Funneliformis mosseae and Entrophospora etunicata) were isolated from the rhizosphere of B. chinense. Functional screening of PGPR revealed that Lysobacter antibioticus, Pseudomonas germanica, Rhodococcus corynebacterioides, and Methylobacterium marchantiae exhibited outstanding abilities in indole-3-acetic acid production, organic phosphorus solubilization, inorganic phosphorus solubilization, and nitrogen fixation, respectively. The pot experiment showed that co-inoculation with PGPR and AMF significantly enhanced the plant growth, biomass accumulation, and nutrient uptake of B. chinense, with plant growth-promoting effects markedly greater than single inoculation treatments. Conclusion This study isolated and identified some plant growth-promoting microorganisms from the rhizosphere of B. chinense and demonstrated the synergistic effects between PGPR and AMF, providing valuable microbial resources and theoretical bases for the sustainable cultivation of B. chinense.

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目的 根际微生物群落对植株生长发育和品质形成具有重要调控作用。因此，本研究旨在系统分离柴胡(Bupleurum chinense DC.)根际促生微生物资源，评价其应用潜力，为开发微生物菌肥、减少化肥和农药投入提供优良菌株。 方法 采用传统分离培养方法，从柴胡根际分离鉴定植物根际促生细菌(plant growth-promoting rhizobacteria, PGPR)和丛枝菌根真菌(arbuscular mycorrhizal fungi, AMF)；通过体外功能试验筛选高效PGPR菌株，并在盆栽条件下评估PGPR和AMF的协同促生效应。 结果 从柴胡根际共分离获得25种PGPR，以及摩西管柄囊霉(Funneliformis mosseae)和幼套内养囊霉(Entrophospora etunicata) 2种AMF。PGPR的功能筛选结果表明，抗生素溶杆菌(Lysobacter antibioticus)、德国鸢尾假单胞菌(Pseudomonas germanica)、类棒杆菌红球菌(Rhodococcus corynebacterioides)和地钱甲基杆菌(Methylobacterium marchantiae)分别在吲哚乙酸分泌、有机磷溶解、无机磷溶解和固氮方面表现突出。盆栽试验显示，PGPR和AMF联合接种显著促进柴胡植株生长、生物量积累和养分吸收，其促生效应明显优于单一接种处理。 结论 本研究挖掘并鉴定了一些柴胡根际关键促生微生物资源，明确了PGPR和AMF之间的协同促生效应，为柴胡绿色高效栽培提供了可利用的菌剂资源和理论依据。

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作者贡献声明

余晶：论文撰写，数据收集及分析；马亚秀：论文修改，数据收集；曾静：论文修改；刘永俊：实验与论文指导、修改润色，获取基金。

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The species names and strain numbers in bold correspond to those obtained in this study, with species names representing the molecularly identified species and K-2, N-2, etc. referring to the strain numbers. The GenBank accession numbers are shown in parentheses. Numbers at the nodes indicate bootstrap support values, and the scale bar represents 0.05 nucleotide substitutions per site. The vertical line area on the right side of the phylogenetic tree indicates the categories of each terminal branch at the phylum level., figureFileSmall=0GJl2fyj8YjkWkjSy3RfIQ==, figureFileBig=qB6aHitnMYmBF7Y2XE1XwA==, tableContent=null), ArticleFig(id=1250879409645896698, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1250834201700942606, language=CN, label=图1, caption=本研究获得的柴胡根际促生细菌基于16S rRNA基因序列构建的最大似然系统发育树, figureFileSmall=0GJl2fyj8YjkWkjSy3RfIQ==, figureFileBig=qB6aHitnMYmBF7Y2XE1XwA==, tableContent=null), ArticleFig(id=1250879409943691278, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1250834201700942606, language=EN, label=Figure 2, caption=The maximum-likelihood phylogenetic tree based on the SSU-ITS-LSU sequences of AMF isolated from the rhizosphere of Bupleurum chinense. The species names and strain numbers in bold correspond to those obtained in this study, with species names representing the molecularly identified species and c-8, c-7, etc. referring to the strain numbers. The GenBank accession numbers are shown in parentheses. 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Control, PGPR, AMF, and PGPR+AMF represent non-inoculated, PGPR inoculated, AMF inoculated, and PGPR and AMF co-inoculated treatments, respectively. Values are presented as mean±SE. Different letters indicate significant differences among treatments (P<0.05)., figureFileSmall=3lJqjQHnycwCQ/EoLKoN5A==, figureFileBig=Tv0jILqOobxYj74zmfn3yA==, tableContent=null), ArticleFig(id=1250879410346344492, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1250834201700942606, language=CN, label=图3, caption=接种根际促生微生物对柴胡生物量、根冠比、比根长和地上组织氮磷比的影响, figureFileSmall=3lJqjQHnycwCQ/EoLKoN5A==, figureFileBig=Tv0jILqOobxYj74zmfn3yA==, tableContent=null), ArticleFig(id=1250879410551865408, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1250834201700942606, language=EN, label=Table 1, caption=

Functional characterization of plant growth-promoting rhizobacteria isolated from the rhizosphere of Bupleurum chinense

, figureFileSmall=null, figureFileBig=null, tableContent=
PGPR performance parametersStrainsResults
IAA concentration (μg/mL)K-1 (Pseudomonas chlororaphis)5.96±0.16b
K-2 (Agrobacterium tumefaciens)6.85±0.44b
K-3 (Microbacterium imperiale)25.80±2.33ab
K-4 (Erwinia tasmaniensis)12.70±0.37ab
K-6 (Microbacterium phyllosphaerae)21.34±0.63ab
K-7 (Lysobacter antibioticus)30.90±1.00a
K-9 (Streptomyces caeruleatus)16.42±2.87ab
K-10 (Anoxybacillus flavithermus)10.04±1.56ab
Organic P solubilization index (%)M-2 (Methylobacterium marchantiae)118.67±4.10ab
M-3 (Bacillus litoralis)119.00±5.00ab
M-5 (Novosphingobium barchaimii)120.67±5.81ab
M-7 (Pseudomonas germanica)239.00±10.03a
M-9 (Arthrobacter celericrescens)125.33±5.78ab
M-10 (Streptomyces novaecaesareae)144.33±8.09ab
M-11 (Streptomyces aureoverticillatus)110.67±0.33b
Inorganic P concentration (μg/mL)N-1 (Anoxybacillus flavithermus)0.60±0.05bc
N-2 (Mesorhizobium amorphae)0.49±0.02c
N-3 (Variovorax paradoxus)0.72±0.03bc
N-4 (Rhodococcus oxybenzonivorans)0.84±0.05ab
N-5 (Caulobacter soli)0.78±0.03ab
N-8 (Rhodococcus corynebacterioides)0.98±0.04a
Acetylene reduction activity [nmol C2H4/(h·mL)]F-1 (Anoxybacillus flavithermus)0.81±0.07b
F-2 (Streptomyces plumbiresistens)2.93±0.16b
F-4 (Methylobacterium marchantiae)8.19±1.58a
F-5 (Agromyces aureus)3.37±1.29b
F-6 (Williamsia muralis)2.25±1.27b
F-7 (Streptomyces peucetius)0.75±0.02b
F-8 (Streptomyces rameus)0.71±0.12b
), ArticleFig(id=1250879410698666060, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1250834201700942606, language=CN, label=表1, caption=

柴胡根际促生细菌的性能评估

, figureFileSmall=null, figureFileBig=null, tableContent=
PGPR performance parametersStrainsResults
IAA concentration (μg/mL)K-1 (Pseudomonas chlororaphis)5.96±0.16b
K-2 (Agrobacterium tumefaciens)6.85±0.44b
K-3 (Microbacterium imperiale)25.80±2.33ab
K-4 (Erwinia tasmaniensis)12.70±0.37ab
K-6 (Microbacterium phyllosphaerae)21.34±0.63ab
K-7 (Lysobacter antibioticus)30.90±1.00a
K-9 (Streptomyces caeruleatus)16.42±2.87ab
K-10 (Anoxybacillus flavithermus)10.04±1.56ab
Organic P solubilization index (%)M-2 (Methylobacterium marchantiae)118.67±4.10ab
M-3 (Bacillus litoralis)119.00±5.00ab
M-5 (Novosphingobium barchaimii)120.67±5.81ab
M-7 (Pseudomonas germanica)239.00±10.03a
M-9 (Arthrobacter celericrescens)125.33±5.78ab
M-10 (Streptomyces novaecaesareae)144.33±8.09ab
M-11 (Streptomyces aureoverticillatus)110.67±0.33b
Inorganic P concentration (μg/mL)N-1 (Anoxybacillus flavithermus)0.60±0.05bc
N-2 (Mesorhizobium amorphae)0.49±0.02c
N-3 (Variovorax paradoxus)0.72±0.03bc
N-4 (Rhodococcus oxybenzonivorans)0.84±0.05ab
N-5 (Caulobacter soli)0.78±0.03ab
N-8 (Rhodococcus corynebacterioides)0.98±0.04a
Acetylene reduction activity [nmol C2H4/(h·mL)]F-1 (Anoxybacillus flavithermus)0.81±0.07b
F-2 (Streptomyces plumbiresistens)2.93±0.16b
F-4 (Methylobacterium marchantiae)8.19±1.58a
F-5 (Agromyces aureus)3.37±1.29b
F-6 (Williamsia muralis)2.25±1.27b
F-7 (Streptomyces peucetius)0.75±0.02b
F-8 (Streptomyces rameus)0.71±0.12b
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柴胡根际植物促生细菌与丛枝菌根真菌资源的挖掘及协同促生效应
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余晶 , 马亚秀 , 曾静 , 刘永俊
微生物学报 | 研究报告 2026,66(4): 1929-1940
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微生物学报 | 研究报告 2026, 66(4): 1929-1940
柴胡根际植物促生细菌与丛枝菌根真菌资源的挖掘及协同促生效应
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余晶, 马亚秀, 曾静, 刘永俊
作者信息
  • 兰州大学 生态学院，甘肃 兰州
Isolation of plant growth-promoting rhizobacteria and arbuscular mycorrhizal fungi from the rhizosphere of Bupleurum chinense and characterization of their synergistic effects on plant growth
Jing YU, Yaxiu MA, Jing ZENG, Yongjun LIU
Affiliations
  • College of Ecology, Lanzhou University, Lanzhou, Gansu, China
出版时间: 2026-04-04 doi: 10.13343/j.cnki.wsxb.20251017
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摘要
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目的 根际微生物群落对植株生长发育和品质形成具有重要调控作用。因此，本研究旨在系统分离柴胡(Bupleurum chinense DC.)根际促生微生物资源，评价其应用潜力，为开发微生物菌肥、减少化肥和农药投入提供优良菌株。 方法 采用传统分离培养方法，从柴胡根际分离鉴定植物根际促生细菌(plant growth-promoting rhizobacteria, PGPR)和丛枝菌根真菌(arbuscular mycorrhizal fungi, AMF)；通过体外功能试验筛选高效PGPR菌株，并在盆栽条件下评估PGPR和AMF的协同促生效应。 结果 从柴胡根际共分离获得25种PGPR，以及摩西管柄囊霉(Funneliformis mosseae)和幼套内养囊霉(Entrophospora etunicata) 2种AMF。PGPR的功能筛选结果表明，抗生素溶杆菌(Lysobacter antibioticus)、德国鸢尾假单胞菌(Pseudomonas germanica)、类棒杆菌红球菌(Rhodococcus corynebacterioides)和地钱甲基杆菌(Methylobacterium marchantiae)分别在吲哚乙酸分泌、有机磷溶解、无机磷溶解和固氮方面表现突出。盆栽试验显示，PGPR和AMF联合接种显著促进柴胡植株生长、生物量积累和养分吸收，其促生效应明显优于单一接种处理。 结论 本研究挖掘并鉴定了一些柴胡根际关键促生微生物资源，明确了PGPR和AMF之间的协同促生效应，为柴胡绿色高效栽培提供了可利用的菌剂资源和理论依据。

药用植物  /  根际微生物  /  菌根真菌  /  微生物接种  /  生态种植

Objective The rhizosphere microbial community plays a critical role in plant growth, development, and quality formation. Therefore, this study systematically isolated plant growth-promoting microbial resources from the rhizosphere of Bupleurum chinense and evaluated their application potential, aiming to provide excellent strains for the development of microbial fertilizers to reduce the use of chemical fertilizers and pesticides. Methods Plant growth-promoting rhizobacteria (PGPR) and arbuscular mycorrhizal fungi (AMF) were isolated and identified from the rhizosphere of B. chinense by the culture-dependent methods. Functional traits of PGPR strains were screened through in vitro assays, and the synergistic growth-promoting effects of PGPR and AMF were subsequently evaluated by a pot experiment. Results A total of 25 PGPR species and 2 AMF species (Funneliformis mosseae and Entrophospora etunicata) were isolated from the rhizosphere of B. chinense. Functional screening of PGPR revealed that Lysobacter antibioticus, Pseudomonas germanica, Rhodococcus corynebacterioides, and Methylobacterium marchantiae exhibited outstanding abilities in indole-3-acetic acid production, organic phosphorus solubilization, inorganic phosphorus solubilization, and nitrogen fixation, respectively. The pot experiment showed that co-inoculation with PGPR and AMF significantly enhanced the plant growth, biomass accumulation, and nutrient uptake of B. chinense, with plant growth-promoting effects markedly greater than single inoculation treatments. Conclusion This study isolated and identified some plant growth-promoting microorganisms from the rhizosphere of B. chinense and demonstrated the synergistic effects between PGPR and AMF, providing valuable microbial resources and theoretical bases for the sustainable cultivation of B. chinense.

medicinal plants  /  rhizosphere microorganisms  /  mycorrhizal fungi  /  microbial inoculation  /  ecological cultivation
余晶, 马亚秀, 曾静, 刘永俊. 柴胡根际植物促生细菌与丛枝菌根真菌资源的挖掘及协同促生效应. 微生物学报, 2026 , 66 (4) : 1929 -1940 . DOI: 10.13343/j.cnki.wsxb.20251017
Jing YU, Yaxiu MA, Jing ZENG, Yongjun LIU. Isolation of plant growth-promoting rhizobacteria and arbuscular mycorrhizal fungi from the rhizosphere of Bupleurum chinense and characterization of their synergistic effects on plant growth[J]. Acta Microbiologica Sinica, 2026 , 66 (4) : 1929 -1940 . DOI: 10.13343/j.cnki.wsxb.20251017
正文
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柴胡(Bupleurum chinense DC.)是我国常用的大宗中药材，以干燥根入药，具有疏散退热、疏肝解郁、升举阳气等功效，广泛应用于多种中成药和临床方剂中[1]。近年来，随着野生柴胡资源持续减少以及市场需求快速增长，柴胡人工栽培面积不断扩大，产业化进程显著加快。然而，在集约化生产条件下，柴胡栽培普遍面临土壤退化、病虫害加剧、有益微生物丧失以及多重生物和非生物胁迫叠加等问题，严重制约了植株生长、品质形成和产业可持续发展[2-5]。
在国家大力推进中药材生态种植和绿色发展的背景下，依赖高投入化肥和农药的传统栽培模式已难以满足“化肥减施、提质增效”的政策要求[6]。因此，将微生物技术这一关键生态调控手段引入柴胡栽培体系，被认为是推动柴胡产业高质量发展的重要途径之一[7]。自然条件下，植物根系富集着复杂多样的促生微生物群落，其中以植物根际促生细菌(plant growth-promoting rhizobacteria, PGPR)和丛枝菌根真菌(arbuscular mycorrhizal fungi, AMF)最具代表性[8-9]。这些微生物可通过分泌植物激素、溶磷、固氮以及改善根系吸收能力等多种途径，促进植物生长发育、增强抗逆性，并有助于活性成分积累，在中药材生态化栽培中具有广阔应用前景[10-11]。在此背景下，挖掘柴胡根际有益微生物成为一个有前景的方向。尽管如此，当前柴胡相关研究主要集中在种质资源评价、次生代谢产物解析及药理活性研究等方面，而针对柴胡根际促生微生物资源的分离鉴定、功能筛选及应用潜力评估仍相对不足[12-14]。尤其是PGPR和AMF在柴胡生长中的实际作用效果及其协同作用机制尚缺乏系统实验验证，限制了微生物技术在柴胡生产实践中的推广应用。
本研究以柴胡根际微生物为研究对象，采用平板分离培养和单孢扩繁技术分别获取柴胡根际PGPR和AMF菌株，并分别通过16S rRNA基因和SSU-ITS-LSU rRNA基因序列分析进行分子鉴定；进一步通过体外功能筛选，遴选具有显著吲哚乙酸分泌、溶磷和固氮能力的优良PGPR菌株；最后，在温室盆栽条件下系统评估优良PGPR和AMF对柴胡生长和养分吸收的促生效应。本研究旨在为柴胡生态种植发掘高效的微生物菌剂资源，并为柴胡产业绿色可持续发展提供理论依据和技术支撑。
1 材料与方法
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1.1 样品采集
本研究于2023年7月在甘肃省甘南藏族自治州舟曲县的3个典型柴胡种植区共采集15份生长2年的人工栽培柴胡根际土壤，用于促生微生物的分离与培养。每个种植区随机选取5个地块，在每个地块中随机选取4-5株生长健壮、无明显病害的柴胡植株，去除表层浮土后将植株连根挖出，并在低温条件下运回实验室。随后，采用无菌毛刷轻刷根系表面，收集距离根部约2 mm范围内的附着土壤作为根际土样，用于PGPR的分离培养；其余土壤作为根围土样，用于AMF的分离与扩繁。
1.2 PGPR的分离培养
PGPR的分离培养采用多种选择性和功能性培养基，包括金氏B培养基(King’s B medium)[15]、蒙金娜有机磷培养基(MeHKNHa organic phosphorus agar medium)[16]、国际植物研究所磷酸盐生长培养基(national botanical research institute’s phosphate growth medium)[17]和无氮培养基(nitrogen free medium)[18]。
针对每份根际土壤样品，称取5 g土样加入到含45 mL无菌生理盐水的三角瓶中，28 ℃、180 r/min振荡30 min，使土壤充分分散，制备10-1稀释的土壤菌悬液。随后取1 mL菌悬液加入到9 mL无菌生理盐水中，依次进行10倍梯度稀释，获得10-2和10-3稀释液。取100 μL 10-3稀释液，分别涂布于金氏B培养基、蒙金娜有机磷培养基、国际植物研究所磷酸盐生长培养基和无氮培养基上，每个样品设3个平行重复，于28 ℃恒温培养箱中培养5-7 d。培养结束后，根据菌落颜色、形态、大小及表面特征进行初步分离，并通过反复划线纯化3-4次，直至获得形态稳定的纯培养菌株。
1.3 AMF的分离培养
针对每一份根围土壤样品，称取50 g土样，采用蔗糖梯度离心法分离AMF孢子[19]。将同一采样点分离获得的孢子合并后，在体视显微镜下根据孢子大小、颜色和形态特征进行形态聚类。随后，选取各形态类型的单个孢子，接种于2周龄无菌高粱幼苗(Sorghum bicolor L.)根系，并用无菌滤纸包裹，移栽至装有灭菌基质的花盆中央(容量500 mL)。栽培基质由沙子与沸石按1:1混合组成，经121 ℃灭菌75 min，间隔24 h重复灭菌1次。在每个花盆滤纸周围播种经表面消毒并露白的高粱和苜蓿(Medicago sativa L.)种子，于温室条件下进行单孢扩繁培养。每种孢子形态设置3-8个重复。温室培养条件：光强约为10 340 lx，光暗周期为16 h光照/8 h黑暗，昼夜温度为25 ℃ (光照期)/20 ℃ (黑暗期)。培养期间，每3-4 d浇灌1次蒸馏水，每2周施加1次低磷Hoagland营养液(P浓度为0.5 mmol/L)。培养4个月后停止浇水，自然干燥2周，剪除地上部分收集植物根系和盆栽基质。取30 g混匀基质样品，采用蔗糖梯度离心法检测AMF孢子形成情况，当观察到大量新生孢子时判定培养成功，将用于后续分子鉴定及多孢扩繁。
1.4 菌株的分子生物学鉴定
PGPR菌株的分子鉴定以16S rRNA基因为目标序列。采用无菌枪头挑取单个PGPR菌落，置于含10 μL无菌ddH2O的离心管中，在80 ℃水浴中热裂解15 min，随后低速离心，取上清液作为PCR扩增的DNA模板。PCR扩增使用细菌通用引物27F (5ʹ-AGAGTTTGATCCTGGC TCAG-3ʹ)和1492R (5ʹ-GGTTACCTTGTTACGA CTT-3ʹ)[20]。PCR反应体系(20 μL)：Taq Mix 10 μL，上、下游引物(5 μmol/L)各1 μL，DNA模板1 μL，ddH2O补足20 μL。PCR扩增程序：94 ℃ 5 min；94 ℃ 30 s，55 ℃ 30 s，72 ℃ 60 s，共30个循环；72 ℃ 10 min。
针对成功扩繁的AMF纯培养菌株，采用AMF特异性引物组合SSUmAf-LSUmAr和SSUmCf-LSUmBr[21]对SSU-ITS-LSU rRNA基因片段进行巢式PCR扩增。这2对引物均为混合引物，其中SSUmAf (SSUmAf1、SSUmAf2)与LSUmAr (LSUmAr1-LSUmAr4)用于第一轮PCR，SSUmCf (SSUmCf1-SSUmCf3)和LSUmBr (LSUmBr1-LSUmBr5)用于第二轮PCR。AMF孢子DNA的提取方法、PCR反应体系及扩增程序均参照刘佳美的方法[22]。
PCR扩增产物经1%琼脂糖凝胶电泳检测确认后，送至生工生物工程(上海)股份有限公司进行测序。获得的序列在NCBI数据库中进行BLAST比对(https://blast.ncbi.nlm.nih.gov/Blast.cgi)，下载与目标序列相似性最高的已知物种作为参考序列。随后，利用IQ-TREE v1.6.12软件[23]，在最佳替代模型下构建最大似然系统发育树，以确定柴胡根际可培养促生微生物的分类地位。所有获得的序列均已提交至GenBank数据库并获得登录号。
1.5 PGPR菌株的功能潜力测定
吲哚-3-乙酸(indole-3-acetic acid, IAA)分泌能力测定：将初筛获得的具有潜在IAA分泌能力的PGPR菌株接种于金氏B液体培养基中，在28 ℃、180 r/min培养24 h；培养结束后，采用Salkowski比色法测定上清液中IAA含量[24]。解磷能力测定：将初筛获得的具有潜在有机磷或无机磷溶解能力的菌株分别点接于蒙金娜有机磷培养基和国际植物研究所磷酸盐生长培养基上，在28 ℃培养7 d；培养结束后，测定菌落直径(d)和溶磷圈直径(D)，并以溶磷指数[(D/d)×100%]评价菌株的磷溶解能力[25]；鉴于大部分解无机磷菌株在固体培养基上未形成明显透明圈，进而采用钼锑抗比色法进行定量测定[26]。固氮能力测定：将初筛获得的具有潜在固氮能力的菌株接种于含5 mL半固体无氮培养基的培养瓶中，在28 ℃培养48 h；随后，使用无菌注射器抽取培养瓶内1 mL气体，并注入等体积乙炔气体，继续培养48 h；培养结束后，取100 μL瓶内气体注入气相色谱仪，采用乙炔还原法测定菌株固氮酶活性[27]。上述各项功能测定均设置3个生物学重复。
1.6 柴胡根际促生菌对柴胡的促生作用
为评估优良PGPR和AMF对柴胡生长的促进效应，在温室条件下开展微生物接种试验。试验设置4个处理：未接种对照、单接PGPR、单接AMF以及PGPR和AMF联合接种，每个处理设置6个重复，共24盆。
盆栽基质为采自柴胡种植区的林下山地表层土壤，未经灭菌处理，每盆装填1.3 kg土壤。PGPR接种物由1.5节筛选获得的4株高效PGPR组成，分别代表IAA分泌、解有机磷、解无机磷和固氮能力最强的菌株。各菌株经摇瓶扩大培养后4 000 r/min离心10 min收集菌体，使用无菌水洗涤并重悬，将菌液浓度统一调整至1×108 CFU/mL，按等体积混合制备PGPR复合接种液。PGPR处理组按10 mL/盆施加接种液，对照处理施加等体积无菌水。
AMF接种物由分离纯化获得的AMF菌株分别经扩大培养后混合制备(培养方法同1.3节，接种物为单孢诱导培养成功的AMF培养物，花盆容积3 L)。根据基质中孢子密度按比例混合，制备AMF复合菌剂。AMF处理每盆施加约1 000个AMF孢子，其中每种AMF孢子各约500个。对照处理每盆加入等量经灭菌处理的AMF菌剂，并额外补加100 mL AMF菌剂滤液(水:菌剂=2:1，经38 μm孔径筛过滤以去除AMF孢子)，以排除非AMF微生物的影响。
柴胡种子经表面消毒后催芽并播种于花盆中，出苗2周后每盆保留10株幼苗。花盆在温室内随机摆放并进行常规水分管理。植物生长16周后收获柴胡植株。地上部分于80 ℃烘干至恒重，称重后粉碎用于测定地上组织氮、磷浓度。根系洗净后使用扫描仪(Epson公司)成像，并利用WinRHIZO Pro v.2020软件(Regent Instruments Inc.公司)分析根系性状，随后烘干测定地下生物量。
1.7 数据处理
本研究所有数据的统计分析与作图均在R v4.2.3软件中完成，采用stats包进行单因素方差分析，检验不同PGPR菌株在IAA分泌、解磷和固氮能力，以及不同接种处理下柴胡生物量、根系性状和组织氮、磷浓度的差异，并使用agricolae包进行Tukey’s HSD事后多重比较。所有统计检验的显著性水平均设定为P<0.05。
2 结果与分析
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2.1 柴胡根际促生菌的分离培养及分子鉴定
利用4种选择性培养基，从柴胡根际土壤中共分离获得28株PGPR。基于16S rRNA基因序列的分子鉴定和系统发育分析，这些菌株共隶属于25个物种(图1)。在功能类型上，分离菌株中IAA分泌菌有8种，溶解有机磷菌有7种，溶解无机磷菌有6种，固氮菌有7种。在门水平，分离获得的PGPR主要隶属于放线菌门(Actinomycetota)、假单胞菌门(Pseudomonadota)和芽孢杆菌门(Bacillota)，分别占总分离菌株的46.43%、39.29%和14.28%。在属水平，链霉菌属(Streptomyces)占优势地位，占PGPR总分离菌株的21.43%，其次为无氧芽孢杆菌属(Anoxybacillus)，占比为10.72%。
在柴胡根际土壤中共观察到7类典型的AMF孢子形态类型。通过单孢扩繁技术成功分离获得7个AMF纯培养物。基于SSU-ITS-LSU rRNA基因序列的分子鉴定结果表明，这些纯培养物属于2种AMF，分别为摩西管柄囊霉(Funneliformis mosseae)和幼套内养囊霉(Entrophospora etunicata) (图2)。
2.2 PGPR的性能及优良菌株筛选
所有IAA分泌菌株均检测到IAA产生能力，其分泌浓度范围为5.96-30.90 μg/mL (表1)。其中，抗生素溶杆菌(Lysobacter antibioticus) K-7的IAA分泌能力最强。解有机磷菌株的溶磷指数介于110.67%-239.00%之间，德国鸢尾假单胞菌(Pseudomonas germanica) M-7表现出最高的有机磷溶解能力(表1)。无机磷溶解菌培养液中的可溶性磷浓度为0.49-0.98 μg/mL，其中类棒杆菌红球菌(Rhodococcus corynebacterioides) N-8表现出较强的无机磷溶解能力(表1)。乙炔还原实验结果显示，所有候选固氮菌株均检测到乙炔还原活性，乙烯生成速率介于0.71-8.19 nmol C2H4/(h·mL)之间(表1)。其中，地钱甲基杆菌(Methylobacterium marchantiae) F-4的乙炔还原活性最高，表明其具有较强的潜在固氮能力。
综合各菌株在IAA分泌、解磷及固氮相关功能指标上的表现，最终筛选出抗生素溶杆菌(Lysobacter antibioticus) K-7、德国鸢尾假单胞菌(Pseudomonas germanica) M-7、类棒杆菌红球菌(Rhodococcus corynebacterioides) N-8和地钱甲基杆菌(Methylobacterium marchantiae) F-4作为促生功能潜力较强的PGPR菌株，用于后续柴胡促生效应的评价。
2.3 柴胡根际促生菌对柴胡的协同促生作用
单独接种PGPR或AMF均轻微促进柴胡生长，但对地上、地下及总生物量的提升未达到显著水平(图3A-3C)。然而，PGPR和AMF联合接种显著提高了柴胡地上、地下和总生物量，相较于对照，增幅分别为95.28%、505.38%和167.19%，其中对地下生物量的促进效应尤为显著(图3A-3C)。联合接种显著改变了柴胡的生物量分配和根系构型，表现为柴胡根冠比上升而比根长下降(图3D、3E)。在养分吸收方面，联合接种显著提高了柴胡组织磷浓度，从而导致地上部组织氮磷比显著降低(图3F)。
3 讨论与结论
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本研究以药用植物柴胡为研究对象，采用传统微生物培养技术对其根际促生微生物进行系统分离与鉴定，共获得25种PGPR和2种AMF。分离获得的PGPR隶属于3门17属，表明柴胡根际可培养促生细菌在系统发育组成上具有较高多样性。在本研究中，链霉菌属(Streptomyces spp.)和无氧芽孢杆菌属(Anoxybacillus spp.)的菌株分离频率较高，是主要优势类群。已有研究表明，这2个属的微生物在多种植物根际中普遍具有促生潜力，可通过改善养分供应、分泌促生物质或抑制病原微生物来促进植物生长[28-29]。
本研究发现，抗生素溶杆菌(Lysobacter antibioticus) K-7、德国鸢尾假单胞菌(Pseudomonas germanica) M-7、类棒杆菌红球菌(Rhodococcus corynebacterioides) N-8和地钱甲基杆菌(Methylobacterium marchantiae) F-4分别在IAA分泌、有机磷溶解、无机磷溶解和固氮能力方面表现最为突出，显示出良好的植物促生潜力。关于这些优势PGPR菌株的功能特性，已有部分研究报道了其促生作用及应用价值。例如，Daid等[30]研究表明，抗生素溶杆菌(Lysobacter antibioticus)具有IAA合成、溶磷及纤维素水解等多种功能，能够显著促进玉米(Zea mays L.)生长并提高田间产量。假单胞菌属(Pseudomonas spp.)中的多种菌株已被证实具备有机磷矿化或无机磷溶解能力[31]，然而关于德国鸢尾假单胞菌(Pseudomonas germanica)溶磷作用的研究迄今尚未见报道。本研究证实该菌株具有较强的有机磷溶解能力，暗示其可能通过分泌磷酸酶等途径参与有机磷的转化，其具体作用机理仍有待深入阐明。此外，Khan等[32]发现，类棒杆菌红球菌(Rhodococcus corynebacterioides)可通过提升红树林幼苗(Avicennia marina)根际土壤中磷酸酶活性改善土壤养分循环，从而促进幼苗在柴油胁迫条件下的生长。晁红军等[33]的研究表明，许多甲基杆菌(Methylobacterium spp.)能够以N2作为氮源，具有固氮潜力。本研究中分离获得的地钱甲基杆菌(Methylobacterium marchantiae)同样表现出较强的固氮能力，并兼具有机磷溶解功能，显示出其在养分受限土壤条件下的潜在应用价值。
大量研究表明，单独接种PGPR或AMF可显著促进玉米、藏红花(Crocus sativus L.)、棉花(Gossypium hirsutum L.)等植物的生长[30,34-35]。在本研究中，尽管单独接种PGPR或AMF略微促进了柴胡的生长，但未达到统计显著水平，这可能与柴胡的特定生理特点或土壤条件有关(本研究使用的是未灭菌的林地土)。尽管如此，当PGPR和AMF联合接种时柴胡的总生物量相较于未接种处理增幅达1.67倍，地下生物量的增幅更高达5.05倍，表明这两类促生菌之间存在明显的协同增效作用。Duan等[36]提出的植 物-AMF-细菌三界互作连续体理论认为，AMF通过菌丝网络为细菌提供植物源碳，而细菌则通过分泌胞外酶促进养分转化，从而协同提升植物生产力和系统稳定性。Zhang等[37]证实，AMF可选择性招募菌根辅助细菌，通过促进AMF菌丝定殖间接增强植物对磷素的获取，且联合接种的促生效应显著优于单一接种。此外，Wang等[38]的研究表明，PGPR可诱导植物根系类黄酮的积累，而类黄酮已被证明能够促进AMF孢子萌发及菌丝在根系中的定殖。因此，PGPR可能通过调控根系分泌物组成间接促进AMF的定殖过程。本研究还发现，PGPR和AMF联合接种显著改变了柴胡根系的形态特征，表现为比根长降低；该结果与de Vries等[39]的研究一致，反映了在有益微生物参与下，植物根系由“资源获取型”向“资源保守型”策略转变的过程，也暗示了在联合接种情况下植物的养分限制得到了很大程度的缓解。由于本研究的PGPR为各种功能细菌的混合菌剂，到底是哪类或哪种细菌和AMF具有显著的协同增效作用仍有待进一步阐明。
综上所述，本研究对柴胡根际PGPR和AMF进行了同步分离与鉴定，获得了25种PGPR和2种AMF纯培养菌株，并筛选出4株具有较强促生潜力的PGPR菌株。盆栽试验进一步证实，PGPR和AMF联合接种能够协同促进柴胡生长和养分吸收。本研究为利用微生物技术改善柴胡生长、推动其生态化栽培提供了重要的菌种资源和理论依据。未来研究应进一步从温室条件拓展至田间尺度，开发适宜田间应用的PGPR-AMF复合菌剂，并结合基因组学、转录组学和代谢组学等多组学手段，深入解析微生物调控柴胡生长的分子机制，为柴胡的可持续生产提供更加坚实的理论基础和技术支撑。
基金
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  • 国家自然科学基金(W2512021)
  • 国家自然科学基金(32471674)
  • 甘肃省自然科学基金(23JRRA1029)
  • 舟曲县科技计划
文献
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2026年第66卷第4期
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doi: 10.13343/j.cnki.wsxb.20251017
  • 接收时间：2025-12-31
  • 首发时间：2026-04-14
  • 出版时间：2026-04-04
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  • 收稿日期：2025-12-31
  • 录用日期：2026-02-09
基金
National Natural Science Foundation of China(W2512021)
国家自然科学基金(W2512021)
National Natural Science Foundation of China(32471674)
国家自然科学基金(32471674)
Natural Science Foundation of Gansu Province(23JRRA1029)
甘肃省自然科学基金(23JRRA1029)
Science and Technology Program of Zhouqu County
舟曲县科技计划
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    兰州大学 生态学院，甘肃 兰州
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2种不同金属材料的力学参数
科
Family		 属数
Number of
genus		 种数
Number of
species		 占总种数比例
Percentage of
total species (%)		 属
Genus		 种数
Number of
species		 占总种数比例
Percentage of total
species (%)
鹅膏菌科Amanitaceae 2 11 5.26 鹅膏菌属 Amanita 10 4.78
小菇科 Mycenaceae 2 12 5.74 丝盖伞属 Inocybe 5 2.39
多孔菌科 Polyporaceae 8 14 6.70 蜡蘑属 Laccaria 5 2.39
红菇科 Russulaceae 3 23 11.00 小皮伞属 Marasmius 6 2.87
小菇属 Mycena 11 5.26
光柄菇属 Pluteus 5 2.39
红菇属 Russula 17 8.13
栓菌属 Trametes 5 2.39
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