Microbial Diversity in Rhizosphere Soil of Pseudostellaria heterophylla and Its Correlation with Main Soil Physicochemical Factor
-
摘要:
目的 探究不同区域栽培太子参产生连作效应的潜在原因。 方法 采用高通量测序对柘荣县5个样地的太子参根际土壤进行微生物群落结构及多样性分析。 结果 土壤真菌的有效序列共744 331条,聚类后获得1314个真菌分类单元;获得细菌1032029条有效序列,聚类后获得10310个细菌分类单元。5个样地的共有OTU数明显多于每个样地特有的OTU数,表明不同样地的太子参根际土壤微生物群落趋势大致相同。α多样性分析表明,不同样地根际土壤微生物群落组成及丰度之间有明显差异,Ⅳ、Ⅱ、Ⅰ样地的物种丰度较高,而Ⅴ、Ⅲ样地的物种丰度较低。典范对应分析表明土壤化学性质和根系分泌物酚酸类物质影响太子参根际土壤微生物的群落分布。5个样地中都鉴定到镰刀菌属等致病真菌,且致病菌的含量与根际土壤其他微生物物种数呈反比,暗示镰刀菌影响太子参根际土壤其他微生物物种数。 结论 太子参连作效应及其病害的成因很大程度上归因于其根系分泌物参与调控的根际微生物菌群结构和多样性演变,导致病原菌增多。 Abstract:Objective Chemical and microbial factors in field soils related to disease occurrence of continuously cropped Pesudostellaria heterophylla were investigated. Method Microbial community structure and diversity of rhizosphere soil from 5 continuous cropping P. heterophylla fields in Zherong County, Fujian were analyzed on data obtained by high-throughput sequencing technology. Correlation analysis of the microbial distribution was conducted based on the chemical properties of rhizosphere soil and phenolic acids. Result A clustering analysis on 744,331 fungal and 1,032,029 bacterial effective sequences resulted in 1,314 operational taxonomic units for the fungi and 10,310 for the bacteria. At each of the 5 locations, the number of common OTUs was significantly greater than that of unique ones. On the other hand, the alpha diversity analysis showed significant differences in microbial composition and abundance among the habitats as the species richness at Habitat Ⅳ, Ⅱ, and Ⅰ were higher than those at Habitat Ⅴ and Ⅲ. The canonical correspondence analysis indicated the soil chemical properties and phenolic acids in the root exudates significantly affected the microbial distribution. Fusarium and other pathogenic fungi were found at all 5 sites with the count of the pathogenic fungi inversely proportional to that of other microbial species, i.e., a possible correlation between Fusarium and the rhizosphere community. Conclusion It appeared that the continuous cropping that frequent fungal diseases on the plants might be significantly associated with the evolution of rhizosphere microbial structure and diversity mediated by the root exudate from P. heterophylla itself. -
图 4 太子参根际土壤优势微生物群落组成
A:“门”水平真菌组成;B:“门”水平细菌组成;C:“属”水平真菌热图;D:“属”水平细菌热图。
Figure 4. Compositions of dominant microbial communities in rhizosphere soil of P. heterophylla habitats
Note: A: fungi community at phylum level; B: bacteria community at phylum level; C: fungi heatmap at genus level; D: bacteria heatmap at genus level.
图 6 太子参根际土壤微生物群落分布、化学性质和酚酸物质的典范对应分析
OM:有机质;TN:总氮;TP:总磷;TK:总钾;NH4+:铵态氮;AP:有效磷;AK:速效钾;Fe:有效态铁;Mn:有效态猛;Zn:有效态锌;Cu:有效态铜;VaA:香草酸;SyA:丁香酸;Van:香兰素;pHA:对羟基苯甲酸;pCA:对香豆酸;FeA:阿魏酸;BeA:安息香酸;SaA:水杨酸;CiA:肉桂酸。
Figure 6. Canonical correspondence analysis on community distribution, chemical properties and phenolic acids in rhizosphere soil of P. heterophylla habitats
Note: OM: organic matter; TN: total nitrogen; TP: total phosphorus; TK: total potassium; NH4+: ammonium nitrogen; AP: available phosphorus; AK: available potassium; Fe: iron element; Mn: manganese element; Zn: zinc element; Cu: copper element; VaA: vanillic acid; SyA: syringic acid; Van: vanillin; pHA: p-Hydroxybenzoic acid: pCA: p-Coumaric acid; FeA: ferulic acid; BeA: benzoic acid; SaA: salicylic acid; CiA: cinnamic acid.
表 1 太子参根际土壤样品信息
Table 1. Information on soils sampled from rhizosphere of P. heterophylla habitats
序号
Number采集地点
Collection places经纬度
Latitude-longitude采样条件
Sampling conditionsⅠ 柘荣县英山乡油麻厝村 Youmacuo Village, Yingshan Township, Zherong County N27°16′42″ E119°49′2″ 一年生 annual Ⅱ 柘荣县东源乡鸳鸯头村 Yuanyangtou Village, Dongyuan Township, Zherong County N27°10′17″ E119°56′42″ 一年生 annual Ⅲ 柘荣县东源乡绸岭村 Chouling Village, Dongyuan Township, Zherong County N27°11′20″ E119°55′3″ 一年生 annual Ⅳ 柘荣县东源乡东源村 Dongyuan Village, Dongyuan Township, Zherong County N27°13′1″ E119°53′40″ 一年生 annual Ⅴ 柘荣县乍洋乡束刘厝村 Shuliucuo Village, Zhayang Township, Zherong County N27°11′54″ E119°59′43″ 一年生 annual 表 2 样品α多样性指数
Table 2. α diversity indices on 5 rhizosphere soil samples at P. heterophylla habitats
微生物 Microbe 样本 Samples 物种丰富度指数 Observed_species 香农指数 Shannon 辛普森指数 Simpson Chao1指数 Chao1 细菌 Bacteria Ⅰ 5016.67ab 10.51a 1.00a 6403.94ab Ⅱ 5458.00a 10.94a 1.00a 6804.53a Ⅲ 3019.67c 9.32b 0.99a 3907.68c Ⅳ 5502.00a 10.68a 1.00a 7007.58a Ⅴ 4407.33b 10.19ab 0.99a 5717.60b 真菌 Fungi Ⅰ 470.00a 6.09a 0.97a 565.75a Ⅱ 417.67a 5.74a 0.95a 512.36a Ⅲ 348.33a 4.70a 0.82a 426.65a Ⅳ 469.00a 6.18a 0.97a 501.44a Ⅴ 410.67a 4.68a 0.81a 506.34a 注:同列数据后字母分别表示差异显著(P<0.05). 表5~8同.
Note: Data with letters on same column indicate significant difference (P<0.05). Same for Table 5-8.表 3 不同样地太子参根际土壤真菌群落中前20种优势分类单元及其相对丰度
Table 3. Top 20 dominant taxa and their relative abundance of fungi community in rhizosphere soil of P. heterophylla habitats
属名 Genus 相对丰度 Relative abundance /% Ⅰ Ⅱ Ⅲ Ⅳ Ⅴ 子囊菌门_未分类 Ascomycota_unclassified 12.98 a 14.06 a 3.13 a 12.37 a 24.03 a 未鉴定 unidentified 14.85 a 13.25 ab 4.96 b 17.16 a 8.03 ab 被孢霉菌属 Mortierella 20.14 a 9.03 b 6.78 b 11.65 b 6.13 b 粪壳菌纲_未分类 Sordariomycetes_unclassified 3.44 a 6.54 a 15.03 a 1.72 a 26.65 a 腐质霉属 Humicola 7.14 a 6.06 a 4.60 a 3.93 a 7.25 a 赤霉属 Gibberella 1.74 a 2.99 b 2.96 b 9.45 a 1.90 b 瘦脐菇属 Rickenella 0.01 a 0.00 a 17.27 a 1.18 a 0.00 a 捷别尔达孢属 Teberdinia 3.51 ab 0.95 b 0.07 b 7.59 a 0.16 b 双担菌属 Geminibasidium 0.01 b 0.00 b 10.59 a 0.15 b 0.00 b 粪壳菌目_未分类 Sordariales_unclassified 1.98 bc 3.93 a 0.06 d 3.05 ab 1.00 cd 隐球菌属 Cryptococcus 3.52 a 1.00 b 2.27 ab 0.59 b 1.16 b 正青霉属 Eupenicillium 2.69 a 1.67 a 1.42 a 0.84 a 1.70 a 稻曲菌属 Villosiclava 2.74 a 2.53 a 0.07 b 2.22 a 0.11 b 粉褶蕈属 Entoloma 0.19 a 7.24 a 0.02 a 0.03 a 0.10 a 柄孢壳属 Podospora 3.75 a 1.76 ab 0.87 ab 0.22 b 0.31 b 真菌_未分类 Fungi_unclassified 0.02 a 1.53 a 1.66 a 1.94 a 0.06 a 肉座菌属 Hypocrea 0.31 a 0.34 a 0.36 a 1.29 a 2.44 a 毛壳菌属 Chaetomium 0.22 b 0.33 b 0.90 b 2.18 a 0.23 a 镰刀菌属 Fusarium 0.57 a 0.69 a 0.06 a 1.66 a 0.40 a 田头菇属 Agrocybe 0.51 a 0.21 a 0.11 a 0.23 a 0.25 a 注:同行数据后字母分别表示差异显著(P<0.05).表4同。
Note: Data with letters on same row indicate significant difference (P<0.05). Same for Table 4.表 4 不同样地太子参根际土壤细菌群落中前20种优势分类单元及其相对丰度
Table 4. Top 20 dominant taxa and their relative abundance of bacteria community in rhizosphere soil of P. heterophylla habitats
属名 Genus 相对丰度 Relative abundance/% Ⅰ Ⅱ Ⅲ Ⅳ Ⅴ 酸杆菌 Gp1 8.28a 9.13a 5.91a 8.98a 5.51a 厚壁菌门_未分类 Firmicutes_unclassified 2.26b 3.19b 6.20a 4.60ab 4.40ab 黄单胞菌科_未分类 Xanthomonadaceae_unclassified 3.25a 1.33a 3.73a 2.14a 8.35a 绿弯菌门_未分类 Chloroflexi_unclassified 3.79a 3.02a 3.76a 3.28a 2.75a β-变形菌纲_未分类 Betaproteobacteria_unclassified 3.58a 3.36ab 1.94b 3.33ab 2.61ab 罗河杆菌属 Rhodanobacter 2.48ab 0.20b 7.51a 0.34b 3.67ab γ-变形菌纲_未分类 Gammaproteobacteria_unclassified 3.22a 1.83b 2.63b 3.41a 2.57ab 细菌_未分类 Bacteria_unclassified 2.20a 2.73a 2.83a 2.90a 2.49a 芽单孢菌属 Gemmatimonas 3.13a 3.76a 1.70a 2.06a 2.34a 放线菌目_未分类 Actinomycetales_unclassified 2.03ab 1.09c 5.72a 1.47ab 2.20a 酸杆菌 Gp3 2.50a 2.77a 1.66a 3.28a 2.12a 红螺菌科_未分类 Rhodospirillaceae_unclassified 1.32b 2.02ab 3.62a 2.51ab 1.55b 根瘤菌目_未分类 Rhizobiales_unclassified 2.09a 2.34a 2.00a 2.31a 2.21a 红螺菌目_未分类 Rhodospirillales_unclassified 1.76b 1.50b 3.84a 2.16b 1.46b 纤线杆菌属 Ktedonobacter 2.51a 1.72ab 1.33b 1.71ab 2.57a 酸杆菌 Gp2 1.75ab 2.14b 1.01c 3.29a 1.34ab 酸杆菌门_酸杆菌_未分类 Acidobacteria_Gp1_unclassified 2.21a 1.84ab 2.01ab 1.42bc 1.23c 勒氏菌属 Gaiella 1.36a 1.23a 2.06a 1.96a 1.81a 放线菌纲_未分类 Actinobacteria_unclassified 2.44a 1.72ab 1.31b 1.96ab 0.95b 未命名 WPS-2_genera_incertae_sedis 1.51a 1.16a 1.68a 1.16a 1.12a 表 5 不同样地太子参根际土壤化学性质
Table 5. Chemical properties of rhizosphere soil from P. heterophylla habitats
序号
Number酸碱度
pH有机质
Organic matter/
(g·kg−1)全氮
Total N/
(g·kg−1)全磷
Total P/
(g·kg−1)全钾
Total K/
(g·kg−1)铵态氮
Ammonium N/
(mg·kg−1)有效磷
Available P/
(mg·kg−1)速效钾
Available K/
(mg·kg−1)Ⅰ 4.82c 32.56c 1.58b 1.54e 1.05b 14.67b 45.90ab 90.00b Ⅱ 5.31a 43.83a 2.37a 2.06c 1.08b 14.00b 51.30ab 62.00c Ⅲ 4.64d 23.30d 1.36c 2.59a 0.67c 14.00b 52.20b 195.00a Ⅳ 4.95b 38.07b 1.28c 1.61d 1.10b 18.67a 44.00b 107.00b Ⅴ 4.82c 39.84b 2.27a 2.38b 2.19a 14.67b 54.50a 106.50b 表 6 不同样地太子参根际土壤中有效态微量元素的含量
Table 6. Content of available trace elements in rhizosphere soil of P. heterophylla habitats
序号
Number有效态锰
Available Mn/(mg·kg−1)有效态铜
Available Cu/(mg·kg−1)有效态锌
Available Zn/(mg·kg−1)有效态铁
Available Fe/(mg·kg−1)Ⅰ 18.45d 4.18a 9.35d 151.94a Ⅱ 19.33c 1.23d 13.19a 64.08c Ⅲ 16.80e 0.76e 5.19e 43.85e Ⅳ 23.14a 2.53c 11.80b 98.50b Ⅴ 21.22b 3.29b 11.61c 49.46d 表 7 不同样地太子参根系分泌物酚酸的含量
Table 7. Content of phenolic acids in root exudates of P. heterophylla at different locations
序号
Number香草酸
Vanillic acid/(mg·kg−1)丁香酸
Syringic acid/(mg·kg−1)对羟基苯甲酸
p-Hydroxybenzoic acid/(mg·kg−1)对香豆酸
p-Coumaric acid/(mg·kg−1)Ⅰ 0.84b 2.99b 1.73c 3.83c Ⅱ 1.38a 4.05a 1.86a 6.07a Ⅲ 0.78c 1.93e 1.76b 1.83e Ⅳ 0.87b 2.85c 1.53d 3.28d Ⅴ 0.71d 2.19d 1.28e 4.77b 表 8 不同样地太子参根系分泌物酚酸的含量
Table 8. Content of phenolic acids in root exudates of P. heterophylla at different locations
序号
Number香兰素
Vanillin/(mg·kg−1)阿魏酸
Ferulic acid/(mg·kg−1)安息香酸
Benzoic Acid/(mg·kg−1)水杨酸
Salicylic acid/(mg·kg−1)肉桂酸
Cinnamic acid/(mg·kg−1)Ⅰ 0.68d 1.45b 3.83a 1.51ab 0.13c Ⅱ 1.02a 2.03a 2.58c 1.40b 0.15a Ⅲ 0.65e 0.90d 2.45c 1.37bc 0.12d Ⅳ 0.72b 1.35c 3.31b 1.24c 0.13b Ⅴ 0.70c 1.46b 3.37b 1.57a 0.15d -
[1] 黄泽豪. 太子参 [J]. 生命世界, 2015(10):38−39.HUANG Z H. Pseudostellaria heterophylla [J]. LIFE WORLD, 2015(10): 38−39.(in Chinese) [2] 黄冬寿, 王树贵. 福建“柘荣太子参”栽培环境的道地性研究 [J]. 中国野生植物资源, 2010, 29(2):12−14. doi: 10.3969/j.issn.1006-9690.2010.02.004HUANG D S, WANG S G. Research on planting environment of genuine Pseudostellaria heterophylla from Zherong, Fujian Province [J]. Chinese Wild Plant Resources, 2010, 29(2): 12−14.(in Chinese) doi: 10.3969/j.issn.1006-9690.2010.02.004 [3] 曾令杰, 林茂兹, 李振方, 等. 连作对太子参光合作用及药用品质的影响 [J]. 作物学报, 2012, 38(8):1522−1528.ZENG L J, LIN M Z, LI Z F, et al. Effects of continuous cropping on photosynthesis and medicinal quality of Pseudostellariae heterophylla [J]. Acta Agronomica Sinica, 2012, 38(8): 1522−1528.(in Chinese) [4] 张重义, 林文雄. 药用植物的化感自毒作用与连作障碍 [J]. 中国生态农业学报, 2009, 17(1):189−196. doi: 10.3724/SP.J.1011.2009.00189ZHANG Z Y, LIN W X. Continuous cropping obstacle and allelopathic autotoxicity of medicinal plants [J]. Chinese Journal of Eco-Agriculture, 2009, 17(1): 189−196.(in Chinese) doi: 10.3724/SP.J.1011.2009.00189 [5] 陈建祥, 王飞, 左群, 等. 贵州省施秉县太子参生产存在问题与高产栽培关键技术 [J]. 农技服务, 2011, 28(2):231−232. doi: 10.3969/j.issn.1004-8421.2011.02.065CHEN J X, WANG F, ZUO Q, et al. Shibing County, Guizhou Province Radix pseudostellariae produces have problem and high-yield culture key technology [J]. Agricultural Technology Service, 2011, 28(2): 231−232.(in Chinese) doi: 10.3969/j.issn.1004-8421.2011.02.065 [6] 林文雄, 熊君, 周军建, 等. 化感植物根际生物学特性研究现状与展望 [J]. 中国生态农业学报, 2007, 15(4):1−8.LIN W X, XIONG J, ZHOU J J, et al. Research status and its perspective on the properties of rhizosphere biology mediated by allelopathic plants [J]. Chinese Journal of Eco-Agriculture, 2007, 15(4): 1−8.(in Chinese) [7] 吴宗伟, 王明道, 刘新育, 等. 重茬地黄土壤酚酸的动态积累及其对地黄生长的影响 [J]. 生态学杂志, 2009, 28(4):660−664.WU Z W, WANG M D, LIU X Y, et al. Phenolic compounds accumulation in continuously cropped Rehmannia glutinosa soil and their effects on R. glutinosa growth [J]. Chinese Journal of Ecology, 2009, 28(4): 660−664.(in Chinese) [8] MCCULLY M. Rhizosphere allelopathy [J]. Allelopathy Journal, 2007, 19(1): 75−84. [9] 陈军, 黄珊瑜, 刘冰, 等. 不同菌肥处理对太子参根际微生物群落的影响 [J]. 福建农业学报, 2015, 30(12):1171−177. doi: 10.3969/j.issn.1008-0384.2015.12.008CHEN J, HUANG S Y, LIU B, et al. Effects of microbial fertilizers on microbial community structure in Radix pseudostellariae Rhizosphere [J]. Fujian Journal of Agricultural Sciences, 2015, 30(12): 1171−177.(in Chinese) doi: 10.3969/j.issn.1008-0384.2015.12.008 [10] 邹立思, 马阳, 侯娅, 等. 基于磷脂脂肪酸(PLFA)技术的不同产地太子参根际土壤微生物群落结构多样性分析 [J]. 中药材, 2018, 41(5):1054−1060.ZOU L S, MA Y, HOU Y, et al. Analysis of diversity of microbial community in rhizosphere soil of Pseudostellaria heterophylla in different habitats based on phospholipid fatty acid(PLFA) [J]. Journal of Chinese Medicinal Materials, 2018, 41(5): 1054−1060.(in Chinese) [11] 袁济端. 福建柘荣太子参基地土壤肥力状况评价 [J]. 福建热作科技, 2007, 32(3):3−7. doi: 10.3969/j.issn.1006-2327.2007.03.002YUAN J D. An evaluation of the fertility of the soil for Pseudostellaria heterophylla falsestarwort at Zherong, fujain [J]. Fujian Science & Technology of Tropical Crops, 2007, 32(3): 3−7.(in Chinese) doi: 10.3969/j.issn.1006-2327.2007.03.002 [12] 陈雅涵, 谢宗强, 薛丽萍. 碳氮元素分析仪测试土壤与植物样品的流程优化 [J]. 现代化工, 2016, 36(4):185−187, 189.CHEN Y H, XIE Z Q, XUE L P. Process optimization for measurements of soil and plant samples with carbon/nitrogen element analyzer [J]. Modern Chemical Industry, 2016, 36(4): 185−187, 189.(in Chinese) [13] 鲍士旦. 土壤农化分析[M]. 3版. 北京: 中国农业出版社, 2000: 36-148. [14] 曾豪杰, 刘宇, 祝建国, 等. 电感耦合等离子体发射光谱法测定土壤中铁、锌、铜、锰元素含量 [J]. 中兽医医药杂志, 2016, 35(5):45−47.ZENG H J, LIU Y, ZHU J G, et al. Determination of Fe, Zn, Cu and Mn in Dawashan soils by inductively coupled plasma-atomic emission spectrometry [J]. Journal of Traditional Chinese Veterinary Medicine, 2016, 35(5): 45−47.(in Chinese) [15] 吴丹, 赵立, 庞文生, 等. 太子参根际土壤酚酸类自毒物质的分析测定 [J]. 中国民族民间医药, 2017, 26(24):32−34.WU D, ZHAO L, PANG W S, et al. Analysis and determination of phenolic acid self-toxic substances inrhizosphere soil of Radix pseudostellariae [J]. Chinese Journal of Ethnomedicine and Ethnopharmacy, 2017, 26(24): 32−34.(in Chinese) [16] 吴红淼, 林文雄. 药用植物连作障碍研究评述和发展透视 [J]. 中国生态农业学报(中英文), 2020, 28(6):775−793.WU H M, LIN W X. A commentary and development perspective on the consecutive monoculture problems of medicinal plants [J]. Chinese Journal of Eco-Agriculture, 2020, 28(6): 775−793.(in Chinese) [17] HU J, PANG W S, CHEN J L, et al. Hypoglycemic effect of polysaccharides with different molecular weight of Pseudostellaria heterophylla [J]. BMC Complementary and Alternative Medicine, 2013, 13(1): 1−9. doi: 10.1186/1472-6882-13-1 [18] 赵庆芳, 周紫鹃, 王树红, 等. 7种植物对黄芪根腐病病原菌的抑制作用研究 [J]. 西北师范大学学报(自然科学版), 2009, 45(5):92−95.ZHAO Q F, ZHOU Z J, WANG S H, et al. Study on the inhibition effects of 7 kinds of plant to the growth of Astragalus membranaceus root rot fungi [J]. Journal of Northwest Normal University (Natural Science), 2009, 45(5): 92−95.(in Chinese) [19] 薛彩云, 严雪瑞, 林天行, 等. 五味子茎基腐病发生初报 [J]. 植物保护, 2007(4):96−99. doi: 10.3969/j.issn.0529-1542.2007.04.027XUE C Y, YAN X R, LIN T X, et al. A preliminary report on occurrence of Schisandra berry stalk rot [J]. Plant Protection, 2007(4): 96−99.(in Chinese) doi: 10.3969/j.issn.0529-1542.2007.04.027 [20] 朱春雨, 刘西莉, 董槿, 等. 麻黄根腐病病原物的分离及鉴定 [J]. 植物病理学报, 2003(3):193−197. doi: 10.3321/j.issn:0412-0914.2003.03.001ZHU C Y, LIU X L, DONG J, et al. Isolation and identification of the pathogens causing root rot disease of Chinese Ephedra [J]. Acta Phytopathologica Sinica, 2003(3): 193−197.(in Chinese) doi: 10.3321/j.issn:0412-0914.2003.03.001 [21] CHEN J, WU L K, XIAO Z G, et al. Assessment of the diversity of Pseudomonas spp. and Fusarium spp. in Radix pseudostellariae rhizosphere under monoculture by combining dgge and quantitative pcr [J]. Frontiers in Microbiology, 2017, 8: 1748. doi: 10.3389/fmicb.2017.01748 [22] WU L K, CHEN J, WU H M, et al. Insights into the regulation of rhizosphere bacterial communities by application of bio-organic fertilizer in Pseudostellaria heterophylla monoculture regime [J]. Frontiers in Microbiology, 2016, 7: 1788. [23] DE FERRIGO D, MONDIN M, LADURNER E, et al. Effect of seed biopriming with Trichoderma harzianum strain INAT11 on Fusarium ear rot and Gibberella ear rot diseases [J]. Biological Control, 2020, 147: 104286. doi: 10.1016/j.biocontrol.2020.104286 [24] 胡贤锋, 王健, 李明, 等. 水稻稻曲病菌侵染行为的研究现状及展望 [J]. 河南农业科学, 2020, 49(7):1−7.HU X F, WANG J, LI M, et al. Progress and perspect of infection behavior of Ustilaginoidea virens [J]. Journal of Henan Agricultural Sciences, 2020, 49(7): 1−7.(in Chinese) [25] 王光华, 刘俊杰, 于镇华, 等. 土壤酸杆菌门细菌生态学研究进展 [J]. 生物技术通报, 2016, 32(2):14−20.WANG G H, LIU J J, YU Z H, et al. Research progress of Acidobacteria ecology in soils [J]. Biotechnology Bulletin, 2016, 32(2): 14−20.(in Chinese) [26] 徐志霞, 王璇, 李慧敏, 等. 不同林龄木麻黄林地土壤细菌及与土壤因子的相关性分析 [J]. 基因组学与应用生物学, 2018, 37(2):780−788.XU Z X, WANG X, LI H M, et al. Correlative analysis of soil factors and soil bacteria in Casuarina equisetifolia woodlands at different stand ages [J]. Genomics and Applied Biology, 2018, 37(2): 780−788.(in Chinese) [27] 葛艺, 徐绍辉, 徐艳. 根际微生物组构建的影响因素研究进展 [J]. 浙江农业学报, 2019, 31(12):2120−2130. doi: 10.3969/j.issn.1004-1524.2019.12.21GE Y, XU S H, XU Y. Review on influencing factors of rhizosphere microbiome assemblage [J]. Acta Agriculturae Zhejiangensis, 2019, 31(12): 2120−2130.(in Chinese) doi: 10.3969/j.issn.1004-1524.2019.12.21 [28] LIU J G, LI X G, JIA Z J, et al. Effect of benzoic acid on soil microbial communities associated with soilborne peanut diseases [J]. Applied Soil Ecology, 2017, 110: 34−42. doi: 10.1016/j.apsoil.2016.11.001 [29] WU H M, QIN X J, WANG J Y, et al. Rhizosphere responses to environmental conditions in Radix pseudostellariae under continuous monoculture regimes [J]. Agriculture, Ecosystems & Environment, 2019, 270/271: 19−31. [30] 高慧芳, 彭晓倩, 李赣新, 等. 太子参根际土壤中群体感应产生菌与淬灭菌筛选、鉴定及对太子参生长的影响 [J]. 中国科技论文在线, 2016, 9(2):1−8.GAO H F, PENG X Q, LI G X, et al. Screening and identification of quorum sensing and quorum quenching bacteria of Pseudostellaria heterophylla in rhizosphere soil and their effects on the growth of Pseudostellaria heterophylla [J]. Sciencepaper Online, 2016, 9(2): 1−8.(in Chinese) [31] 高慧芳, 彭晓倩, 郭泽望, 等. 太子参连作过程根系分泌物中酚酸类物质的测定及其对黏质沙雷氏菌生长的影响 [J]. 中国科技论文在线, 2017, 1(3):1−7.GAO H F, PENG X Q, GUO Z W, et al. Assay of phenolic acid from root exudates during continuous cropping process of Pseudostellaria heterophylla and their effects on the growth of Serratia marcescens from rhizosphere soil [J]. Sciencepaper Online, 2017, 1(3): 1−7.(in Chinese) [32] ZHANG L Y, GUO Z W, GAO H F, et al. Interaction of Pseudostellaria heterophylla with quorum sensing and quorum quenching bacteria mediated by root exudates in a consecutive monoculture system [J]. Journal of Microbiology and Biotechnology, 2016, 26(12): 2159−2170. doi: 10.4014/jmb.1607.07073