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Sample	pH	Soil organic matter (g/kg) **	Total N (g/kg) **	Alkaline hydrolyzable N (g/kg) **	Available P (mg/kg) **	AmmoniumN (mg/kg) **	Nitrate N (mg/kg) **	Available K (mg/kg) *	Conductivity (μs/cm) **
XaRHL	8.82±0.01	22.24±0.25	1.03±0.02	41.53±1.20	10.34±0.15	4.60±0.39	4.66±0.10	233.80±2.86	127.73±0.55
XaRLL	8.75±0.03	15.43±0.48	1.38±0.03	64.99±2.40	22.31±0.54	6.84±0.52	18.95±0.32	246.78±3.02	88.53±0.31

Sample	pH	Soil organic matter (g/kg) **	Total N (g/kg) **	Alkaline hydrolyzable N (g/kg) **	Available P (mg/kg) **	AmmoniumN (mg/kg) **	Nitrate N (mg/kg) **	Available K (mg/kg) *	Conductivity (μs/cm) **
XaRHL	8.82±0.01	22.24±0.25	1.03±0.02	41.53±1.20	10.34±0.15	4.60±0.39	4.66±0.10	233.80±2.86	127.73±0.55
XaRLL	8.75±0.03	15.43±0.48	1.38±0.03	64.99±2.40	22.31±0.54	6.84±0.52	18.95±0.32	246.78±3.02	88.53±0.31

Sample	Urease ((μg/g)/24 h) **	Catalase ((μmol/g)/h) **	Alkaline phosphatase ((mg/g)/24 h) **	Dehydrogenase ((μL/g)/6 h) *	Sucrase ((mg/g)/24 h) **
XaRHL	185.65±12.32	2 288.08±12.89	0.84±0.03	0.006 2±0.000 1	8.68±0.49
XaRLL	1 641.12±66.64	2 661.86±28.92	1.65±0.03	0.007 5±0.000 3	13.77±0.18

Sample	Urease ((μg/g)/24 h) **	Catalase ((μmol/g)/h) **	Alkaline phosphatase ((mg/g)/24 h) **	Dehydrogenase ((μL/g)/6 h) *	Sucrase ((mg/g)/24 h) **
XaRHL	185.65±12.32	2 288.08±12.89	0.84±0.03	0.006 2±0.000 1	8.68±0.49
XaRLL	1 641.12±66.64	2 661.86±28.92	1.65±0.03	0.007 5±0.000 3	13.77±0.18

Sample	Microbe	Chao1	Faith_pd	Observed_features	Shannon	Simpson
XaRHL	Bacteria	2 998.87±186.76	161.84±10.59	2 977.33±185.71	10.15±0.21	0.996 0±0.00
	Fungi	579.00±197.61	87.37±21.86	579.00±197.61	4.58±1.06	0.850 0±0.08
XaRLL	Bacteria	3 014.49±315.97	172.50±19.66	2 991.33±313.29	10.46±0.08	0.998 0±0.00
	Fungi	763.67±76.88	109.66±9.31	763.67±76.88	6.25±0.96	0.923 0±0.08

Sample	Microbe	Chao1	Faith_pd	Observed_features	Shannon	Simpson
XaRHL	Bacteria	2 998.87±186.76	161.84±10.59	2 977.33±185.71	10.15±0.21	0.996 0±0.00
	Fungi	579.00±197.61	87.37±21.86	579.00±197.61	4.58±1.06	0.850 0±0.08
XaRLL	Bacteria	3 014.49±315.97	172.50±19.66	2 991.33±313.29	10.46±0.08	0.998 0±0.00
	Fungi	763.67±76.88	109.66±9.31	763.67±76.88	6.25±0.96	0.923 0±0.08

三峡水库消落区苍耳根际微生物群落结构解析

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周兰芳 ¹^,²^,³ , 吴胜军 ²^,^* , 马茂华 ² , 邹航 ¹ , 黄金夏 ¹ , 杨军 ²^,³

微生物学报 | 研究报告 2025,65(2): 582-596

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微生物学报 | 研究报告 2025, 65(2): 582-596

三峡水库消落区苍耳根际微生物群落结构解析

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周兰芳¹^,²^,³, 吴胜军²^,^*, 马茂华², 邹航¹, 黄金夏¹, 杨军²^,³

作者信息

1 重庆交通大学河海学院，重庆

2 中国科学院重庆绿色智能技术研究院水库水环境重点实验室，重庆

3 中国科学院大学重庆学院，重庆

Microbial community structure in the rhizosphere of Xanthium sibiricum in the water-level-fluctuation zone of the Three Gorges Reservoir

Lanfang ZHOU¹^,²^,³, Shengjun WU²^,^*, Maohua MA², Hang ZOU¹, Jinxia HUANG¹, Jun YANG²^,³

Affiliations

1 School of River and Ocean Engineering, Chongqing Jiaotong University, Chongqing, China

2 Key Laboratory of Reservoir Aquatic Environment, Chongqing Institute of Green and Intelligent Technology, Chinese Academy of Sciences, Chongqing, China

3 Chongqing School, University of Chinese Academy of Sciences, Chongqing, China

出版时间: 2025-02-04 doi: 10.13343/j.cnki.wsxb.20240580

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摘要

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【目的】 揭示三峡水库消落区典型优势植物苍耳(Xanthium sibiricum)根际微生物群落结构特征，阐明其与土壤质量的关系。 【方法】 在三峡库区腹心所在地云阳县典型消落区采集不同水淹胁迫强度下的苍耳根际土壤，随后进行高通量测序、微生物多样性分析、群落结构解析和冗余分析。 【结果】 在水淹胁迫强(XaRLL)和低(XaRHL)区域中，变形菌门(Proteobacteria)是苍耳根际细菌群落中共同的最优势细菌门，最优势的真菌门则分别为子囊菌门(Ascomycota)和担子菌门(Basidiomycota)。无论是细菌还是真菌，线性判别分析(linear discriminant analysis effect size, LEfSe)结果显示，XaRLL的关键生物标志物总是多于XaRHL。功能预测分析发现，与有氧呼吸相关的PWY-3781是XaRLL和XaRHL二者共同富集的优势代谢途径。整体而言，苍耳根际微生物群落对土壤的理化性质和酶活的变化反应强烈。 【结论】 本研究为理解水库消落区植物与其根际微生物的关系，以及它们对强烈水淹胁迫逆境的适应性提供了理论基础。

关键词

三峡水库 / 消落区 / 苍耳 / 根际微生物

Abstract

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[Objective] To understand the microbial community structure and its relationship with soil quality in the rhizosphere of the dominant plant Xanthium sibiricum in the water-level-fluctuation zone (WLFZ) of the Three Gorges Reservoir. [Methods] We collected the rhizosphere soil samples of X. sibiricum exposed to different flooding stress conditions in a typical WLFZ in Yunyang County, the heart of the Three Gorges Reservoir area. High-throughput sequencing was carried out to analyze the microbial diversity and community structure, and the redundancy analysis was then conducted. [Results] Proteobacteria was the dominant bacterial phylum in the rhizosphere bacteria of X. sibiricum under strong flooding stress (XaRLL) and weak flooding stress (XaRHL), while Ascomycota and Basidiomycota were the dominant fungal phyla in the two types of soil, respectively. Regardless of bacteria or fungi, the linear discriminant analysis effect size (LEfSe) showed that XaRLL always had more key biomarkers than XaRHL. Functional prediction revealed that PWY-3781 associated with aerobic respiration was a dominant metabolic pathway enriched by microorganisms from both XaRLL and XaRHL. Overall, the bacteria and fungi in the rhizosphere of X. sibiricum had strong responses to changes in soil physicochemical properties and enzyme activity. [Conclusion] The results provide a theoretical basis for understanding the relationship between plants and their rhizosphere microorganisms in the WLFZ, as well as their adaptability to strong flooding stress.

Key words

Three Gorges Reservoir / water-level-fluctuation zone / Xanthium sibiricum / rhizosphere microorganism

引用本文

周兰芳, 吴胜军, 马茂华, 邹航, 黄金夏, 杨军. 三峡水库消落区苍耳根际微生物群落结构解析. 微生物学报, 2025 , 65 (2) : 582 -596 . DOI: 10.13343/j.cnki.wsxb.20240580

Lanfang ZHOU, Shengjun WU, Maohua MA, Hang ZOU, Jinxia HUANG, Jun YANG. Microbial community structure in the rhizosphere of Xanthium sibiricum in the water-level-fluctuation zone of the Three Gorges Reservoir[J]. Acta Microbiologica Sinica, 2025 , 65 (2) : 582 -596 . DOI: 10.13343/j.cnki.wsxb.20240580

正文

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苍耳(Xanthium sibiricum)属于一年生草本植物，隶属于菊科苍耳属。苍耳在我国分布广泛，常见于路边、草地和坡地，是三峡水库消落区的优势植物之一^[1-2]。三峡水库消落区是三峡工程建成后形成的生态敏感脆弱带，反季节性、周期性的水淹胁迫对消落区的植被及土壤造成了深远的影响^[3-4]。水库消落区的苍耳等优势植物可能通过重组其根际微生物群落结构来帮助植物适应这种强烈的水淹胁迫。研究证实，在自然的环境下，植物的繁殖生长和生物量的提升在很大程度上与植物-根际微生物群落的互作有关^[5-6]。研究三峡水库消落区较早出现的一批演替优势物种——苍耳与其根际微生物之间的相互作用，对理解水库消落区植被的适应性及生态恢复具有重要意义。然而，截至目前，尚无研究解析不同水淹胁迫下三峡水库消落区苍耳根际微生物群落结构组成特征及其关键影响因子。

植物根际是微生物群落富集区域，是大量土壤微生物与植物互作最为紧密的区域，含有大约10¹¹个微生物细胞^[6-8]。通过这些根际微生物，植物能够改善自身的生存状况，促进生长发育，适应逆境胁迫并防治病虫害^[9-11]。植物根际微生物受环境条件的影响。研究发现，攸县油茶根际真菌的丰度与总磷和土壤月平均温度呈正相关关系，但与总钾、硝态氮以及总氮和总磷的比例呈负相关关系^[12]。谭雪等^[13]研究发现，三峡水库消落区的3种植物(旱柳、狗牙根和牛鞭草)的土壤细菌群落易受到土壤pH、土壤硝态氮及全氮质量分数的影响。Xu等^[14]研究显示，四合木(Tetraena mongolica)、霸王(Sarcozygium xanthoxylon)、白刺(Nitraria tangutorum)根际微生物群落组成受到土壤环境变量的显著影响，其中总磷对细菌群落组成影响最大，而有效钾对真菌群落组成影响最大。对于峨眉拟单性木兰(Magnolia sinica)根际土壤真菌群落，土壤pH、有效钾、总氮、总磷、总钾被发现具有显著影响^[15]。冗余分析发现，土壤环境因子对马铃薯根际细菌和真菌群落影响显著，其中影响较大的是土壤全氮、有机碳和pH^[16]。蒙特卡罗置换检验显示，土壤有效磷、总磷、有效钾和有机质是影响樟子松根际细菌的主要环境变量^[17]。

本研究以重庆市三峡库区腹心所在地云阳县典型消落区作为研究对象，将消落区划分为水淹胁迫强区和低区，并从中采集根际土壤，分析土壤养分及理化性质、5种土壤酶活对三峡水库消落区苍耳的根际细菌群落和真菌群落组成的影响；对比分析水淹胁迫强与低区根际微生物群落组成、α多样性、关键生物标志物及代谢途径的差异；进一步通过冗余分析，揭示各种环境因子与苍耳根际细菌和真菌群落之间的关系。本研究旨在为理解极度敏感脆弱的三峡水库消落区生态系统中植物-微生物的互作及植被恢复提供理论基础。

1 材料与方法

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1.1 苍耳根际土壤采集

云阳县地处重庆市东北部、三峡库区腹心，年均气温、年降水量和年均日照时数分别为18.7 ℃、1 145 mm和1 528 h。在云阳县消落带，典型的优势植物为苍耳。根际土壤样品采自于重庆市云阳县桔园村附近消落区(30.947°N，108.692°E)。布设的采样区域位于165 m以上及以下，分别对应水淹胁迫强度低和高的区域。由于该区域的苍耳普遍比较幼小，因此对于这两类区域的采样点均在15个以上。将采集的根际土壤按照水淹胁迫强度分装成12份，即用于根际土壤理化性质分析的样品6份和用于根际土壤微生物高通量测序分析的样品6份。对于取自水淹胁迫强区域的苍耳根际土壤(XaRLL)，3份样品分别标记为XaRLL-1、XaRLL-2和XaRLL-3；对于取自水淹胁迫低区域的苍耳根际土壤(XaRHL)，3份样品分别标记为XaRHL-1、XaRHL-2和XaRHL-3。

1.2 苍耳土壤理化性质和酶活测定

将采集的苍耳根际土壤样品在实验室阴凉处摊开晾干，共测定9个理化性质指标(电导率、pH、有效磷、有效钾、铵态氮、碱解氮、硝态氮、全氮、土壤有机碳)和5个土壤酶活(过氧化氢酶、脲酶、碱性磷酸酶、脱氢酶与蔗糖酶)。土壤pH和有机质的测定参考NY/T 1121.2—2006标准^[18]。土壤全氮和碱解氮的测定分别依据LY/T 1228—2015^[19]和LY/T 1229—1999^[20]。土壤有效磷的测定依据NY/T 1121.7—2014^[21]中的钼锑抗比色法。土壤有效钾含量的测定则参考的是鲍士旦的《土壤农化分析》(第3版)^[22]。土壤电导率参考HJ 802—2016方法^[23]测定。以氯化钾为浸提液，采用紫外分光光度法分析硝态氮和铵态氮的含量。除了脱氢酶，过氧化氢酶、脲酶、碱性磷酸酶与蔗糖酶酶活的分析主要采用购买的试剂盒，按照相应说明完成检测。采用氯化三苯基四氮唑比色法分析土壤脱氢酶活。检测的各理化性质与酶活值为3个平行样品的平均值。

1.3 苍耳根际微生物基因组提取及高通量测序

利用CTAB方法提取DNA，从而获取苍耳根际土壤微生物基因组，检测DNA纯度与浓度。使用特异性引物341F (5′-CCTAYGGGRBGCASCAG-3′)和806R (5′-GGACTACNNGGGTATCTAAT-3′)扩增16S rRNA基因V3-V4可变区；同时采用特异性引物ITS1F (5′-CTTGGTCATTTAGA GGAAGTAA-3′)和ITS2R (5′-GCTGCGTTCTT CATCGATGC-3′)扩增ITS区。PCR反应体系：Husion^® High-Fidelity PCR Master Mix (New England Biolabs公司) 5 µL，上、下游引物各2 µmol/L，模板DNA约10 ng。PCR反应条件：98 ℃预变性1 min；98 ℃变性10 s，50 ℃退火30 s，72 ℃延伸30 s，30个循环；最后72 ℃终延伸5 min。扩增完成后，对水淹胁迫强和低区各组的3个平行样品进行Illumina NovaSeq测序。将测序得到的原始数据通过QIIME 2等软件进行加工处理，包括修剪、去噪、拼接、嵌合体去除、质控、数据库比对、物种分类信息表生成等，以便解析微生物群落结构。利用Chao1、Observed-features、Simpson、Faith-pd和Shannon指数分析苍耳根际微生物群落的α多样性^[24]。从门、目和属水平上分析苍耳根际细菌和真菌的群落结构组成特征。利用冗余分析探索以上土壤理化性质和土壤酶活对苍耳根际微生物群落的影响。苍耳根际微生物群落中的关键生物标志物通过线性判别分析(linear discriminant analysis effect size, LEfSe)方法(LDA score≥4.0)识别。苍耳根际微生物群落的代谢途径预测及分析采用PICRUSt2^[25]和MetaCyc通路数据库^[26]完成。

2 结果与讨论

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2.1 苍耳根际土壤理化性质与酶活分析

通过理化性质分析(表1)，处于水淹胁迫强(XaRLL)和低(XaRHL)区的苍耳根际土壤均表现为弱碱性，且二者在pH方面无显著性差异(P>0.05)。除了pH，其余理化参数均存在显著差异，其中有机碳、全氮、碱解氮、碱性有效磷、铵态氮、硝态氮、电导率在XaRLL和XaRHL间呈极显著差异(P<0.01)。对于苍耳根际土壤酶活(表2)，其情况与理化性质相似，XaRLL和XaRHL的脲酶、过氧化氢酶、碱性磷酸酶、脱氢酶和蔗糖酶均存在显著差异，其中除了脱氢酶外，均呈极显著差异(P<0.01)。这些结果表明周期性的水淹胁迫已引起了苍耳根际土壤理化性质和酶活的改变，进而可能影响其根际微生物群落结构的组成及特征。

2.2 苍耳根际微生物群落组成

在处于水淹胁迫强(XaRLL)和低(XaRHL)的苍耳根际细菌群落中，变形菌门(Proteobacteria)在门水平上相对丰度均是最高的，分别占据55.4%和39.4% (图1A)。Proteobacteria为原核生物中最大的分支之一，已知革兰氏阴性菌的绝大多数是Proteobacteria^[27]。许多已知的人类、植物和动物致病菌属于Proteobacteria。以前的研究也发现，在石斛^[7]、拟南芥^[28]、Chinese leek^[29]、Setaria viridis、Flaveria bidentis、maize^[30]等植物的根际细菌群落中，Proteobacteria是优势菌门。通过对557组16S rRNA基因扩增子进行分析，Ling等证实Proteobacteria在植物根际富集^[31]。然而，位于Proteobacteria之后的优势菌门存在差异。XaRLL中处于最优势菌门之后的菌门分别是厚壁菌门(Firmicutes) (10.4%)、酸杆菌门(Acidobacteria)(10.0%)和拟杆菌门(Bacteroidetes) (7.2%)，而XaRHL中紧随最优势菌门之后的优势菌门发生了变化，分别是Bacteroidetes (17.2%)、Firmicutes (16.4%)和放线菌门(Actinobacteria) (9.1%)。值得注意的是，最优势菌门Proteobacteria在水淹胁迫强的苍耳根际土壤中的占比相较于水淹胁迫低的苍耳根际土壤中高了16.0%，这可能是由于更多的Proteobacteria有助于寄主植物苍耳适应强烈的水淹胁迫逆境压力。在目水平上，XaRLL和XaRHL细菌群落结构在优势菌目的相对丰度上存在较大差异(图1B)。在XaRHL上的最优势菌目为伯克霍尔德氏菌目(Burkholderiales) (10.4%)和腐败螺旋菌目(Saprospirales) (9.1%)在XaRLL中分别只占6.3%和4.2%。在属水平上，XaRLL和XaRHL最优势的菌属均是乳杆菌属(Lactobacillus)(图1C)，但是这种优势在XaRLL中并不明显，Kaistobacter (7.0%)、硫杆菌属(Thiobacillus)(6.3%)和Lactobacillus (8.5%)的占比接近。在XaRHL中占优势的是黄色土壤杆菌属(Flavisolibacter)(7.0%)、节杆菌属(Arthrobacter)(4.1%)和沙壤土杆菌属(Ramlibacter)(3.6%)在XaRLL中几乎消失。在XaRLL中占据优势的Thiobacillus (6.3%)和红长命菌属(Rubrivivax)(3.4%)在XaRHL中几乎消失，占比接近0。

在XaRLL和XaRHL真菌群落中，苍耳根际土壤中最优势的菌门不同(图2A)，分别是子囊菌门(Ascomycota) (79.5%)和担子菌门（Basidiomycota) (54.5%)。这二者的总和在XaRLL (88.6%)和XaRHL (86.9%)真菌群落中是接近的。Ascomycota被报道与植物细胞壁多糖的降解有关，而Basidiomycota常与木质素的分解有关^[32]。葛应兰和孙廷^[16]的研究也发现，马铃薯根际土壤中子囊菌门(Ascomycota)和担子菌门(Basidiomycota)是占据优势的真菌门。Gqozo等^[33]的研究也通过高通量测序分发现，Ascomycota和Basidiomycota是小麦根际土壤真菌群落中呈现优势的微生物。此外，Ascomycota和Basidiomycota也是三七^[34]、苹果^[35]中常见的优势真菌菌门。Manici等^[32]比较分析了土壤中Ascomycota和Basidiomycota酶组成的异同，发现它们均含有很多相同的植物多糖降解酶，但Ascomycota更加富含磷和硫代谢酶。除这2个菌门外，在XaRLL中占据较大优势的是壶菌门(Chytridiomycota) (7.1%)，而在XaRHL中占据较大优势的是丝孢菌门(Mortierellomycota )(11.4%)。在目水平上，XaRLL和XaRHL真菌群落组成差异较大(图2B)。在XaRHL中拥有绝对优势的是伞菌目(Agaricales) (54.2%)，在XaRLL中仅占4.3%。在XaRLL中占据明显优势的粪壳菌目(Sordariales) (26.2%)和肉座菌目(Hypocreales) (16.2%)，在XaRHL中仅占很小的比例(<5.0%)。在属水平上，XaRLL和XaRHL真菌群落也展现出明显不同(图2C)。锥盖伞属(Conocybe)在XaRLL中仅占1.8%，但在XaRHL中占据绝对优势(54.5%)。腐质霉属(Humicola)在XaRLL中占比18.5%，但在XaRHL中占比接近于0。Zopfiella (6.5%)、Dokmaia (4.7%)和赤霉菌属(Gibberella)(5.1%)在XaRLL中均占有一定比例，但在XaRHL中接近消失。被孢霉属(Mortierella)在XaRHL中占比12.7%，但在XaRLL中仅占2.1%。

LEfSe分析是一种区分不同样品间微生物差异的常用方法^[36-38]。本研究通过LEfSe分析(LDA score≥4.0)调查了XaRLL和XaRHL的微生物群落间呈显著差异的生物标志物(图3)。在细菌群落中，对于XaRHL，呈显著差异的生物标志物含2个门(Actinobacteria和Bacteroidetes)，2个纲(Actinobacteria和Saprospirae)，2个目(Actinomycetales和Saprospirales)，3个科(Oxalobacteraceae、Micrococcaceae和Chitinophagaceae)以及3个属(Ramlibacter、Arthrobacter和Flavisolibacter)；对于XaRLL，呈显著差异的生物标志物包括2个门(Proteobacteria和Nitrospirae)，2个纲(Deltaproteobacteria和Nitrospira)，3个目(Hydrogenophilales、Nitrospirales和Desulfuromonadales)，3个科(Hydrogenophilaceae、Hyphomicrobiaceae和Geobacteraceae)以及5个属(Thiobacillus、Rubrivivax、Geobacter、Rhodoplanes和Rhodobacter)。这表明水淹胁迫强度大的区域中苍耳根际细菌特异的生物标志物种类更多，这与α多样性分析的结果一致。本研究也发现在真菌群落中，XaRLL中特异性的生物标志物比起XaRHL中的多，XaRLL中特异的生物标记物包括19个(1个门，Ascomycota；3个纲，粪壳菌纲(Sordariomycetes)、座囊菌纲(Dothideomycetes)和黑粉菌纲(Ustilaginomycetes)；4个目，Sordariales、Hypocreales、黑粉菌目(Ustilaginales)和小丛壳科(Glomerellales)；5个科，毛壳科(Chaetomiaceae)、葡萄穗霉科(Stachybotryaceae)、黑粉菌科(Ustilaginaceae)、毛刷囊菌科(Trichocomaceae)和小囊菌科(Plectosphaerellaceae)；6个属，腐质霉属(Humicola)、Striaticonidium、踝节菌属(Talaromyces)、黑粉菌属(Ustilago)、赭霉属(Ochroconis)和寡营养丛孢霉属(Plectosphaerella)，而XaRHL中特异的生物标记物只有10个(2个门，Mortierellomycota和Basidiomycota；2个纲，被孢霉纲(Mortierellomycetes)和伞菌纲(Agaricomycetes)；2个目，被孢霉目(Mortierellales)和Agaricales；2个科，被孢霉科(Mortierellaceae)和粪锈伞科(Bolbitiaceae)；2个属，被孢霉属(Mortierella)和锥盖伞属(Conocybe)。更多的细菌或真菌特异生物标志物间接反映了细菌多样性和真菌多样性，这些多样性可能有助于苍耳适应三峡水库消落区如此强烈的逆境胁迫。以前的研究也采用LEfSe分析了其他植物根际土壤的关键生物标记物；例如，Liu等^[39]采用LEfSe分析识别关键生物标记物，并发现山核桃种植园中根际土壤关键的生物标记物为Actinobacteria和鸡油菌目(Cantharellales)，这与本研究的发现存在差异。

2.3 苍耳根际微生物的α多样性与代谢功能

本研究采用了5个指标参数来表征苍耳根际微生物的α多样性(表3)，它们分别是表征微生物群落物种种数和丰富度的Chao1指数和Observed_features，反映微生物群落均匀度的Simpson指数^[40-41]，衡量微生物多样性的Faith_pd指数^[42]，以及评估群落多样性的Shannon指数^[43]。XaRLL和XaRHL的Chao1指数值分别为3 014.49±315.97和2 998.87±186.76；同样地，它们的Observed-features值分别为 2 991.33±313.29和2 977.33±185.71。这表明，在水淹胁迫强度高的区域，细菌的丰度相较于水淹胁迫强度低的区域更高。此外，XaRLL的Faith_pd值、Shannon值和Simpson值均大于XaRHL。这进一步表明，在水淹胁迫强度高的区域，细菌群落的多样性和均匀性相较于水淹胁迫强度低的区域更高。XaRLL中独有的细菌OTUs数量多于XaRHL (5 948 vs. 5 723)，二者共有的细菌OTUs数量为1 085个(图4A)。

通过PICRUSt2^[25]和MetaCyc通路数据库^[26]的代谢功能分析，发现PWY-3781 [aerobic respiration I (cytochrome c)]的相对丰度最高。在XaRLL中，其占比约为1.5%；而在XaRHL中，占比约为1.4% (图5A)。另外，这一相对丰度值与我们之前在长江上游乌东德水库消落区观察到的正常生长状态的蓟罂粟根际细菌群落中的相对丰度值几乎相同(约为1.5%)^[44]。其次是PWY-7111 [pyruvate fermentation to isobutanol (engineered)]、PWY-5101 [l-isoleucine biosynthesis II]、ILEUSYN-PWY [l-isoleucine biosynthesis I (from threonine)]、VALSYN-PWY [l-valine biosynthesis]等。XaRLL和XaRHL二者细菌的群落代谢功能组成模式极为相似，top 20代谢功能的总丰度在整个群落中分别仅占据15.7%和15.1%。这表明在水淹胁迫强度高的区域，细菌群落top 20代谢功能的总丰度相较于水淹胁迫强度低的区域更高。

苍耳根际真菌群落的情况与细菌群落整体趋势相似(表3)。XaRLL的Chao1指数值为763.67±76.88，而XaRHL的为579.00±197.61，二者相差184.67，这表明水淹胁迫的增加可能使苍耳对根际真菌群落进行了重新组装，从而提高了其根际真菌的丰富度。与Chao1功能类似的Observed_features指数的分析也进一步证明了这一点。此外，水淹胁迫强度大的XaRLL的真菌多样性也有一定的提升，正如Shannon指数和Simpson指数显示的，与XaRHL相比，分别提升了1.67和0.073 0。XaRLL和XaRHL共有的真菌OTUs数量为484个，独有的OTUs数量分别为1 078和786 (图4B)。

此外，本研究还分析了XaRLL和XaRHL真菌群落的代谢功能(图5B)，发现二者的最优势代谢途径同为PWY-3781，其次为PWY-7279 [aerobic respiration II (cytochrome c) (yeast)]，这与乌东德水库消落区蓟罂粟根际真菌群落的情况完全相同^[44]。这些发现表明，在水库消落区极端逆境下，植物根际真菌倾向于富集与有氧呼吸相关的代谢途径，以此作为适应生存的策略。在中国云南高原湖泊的相关研究中，与前体代谢产物的生成相关的代谢途径中，PWY-3781和PWY-7279的相对丰度也被报道高于其他代谢途径^[45]。此外，PWY-3781被发现是Brachystegia boehmii和Brachystegia spiciformis根际土壤中最重要的代谢途径^[46]。至于剩余的18条top代谢途径，在二者中的相对丰度也极为接近。此外，top 20代谢途径的总丰度恰好与细菌群落的情况截然相反，即XaRLL中top 20途径的总丰度低于XaRHL。

2.4 苍耳根际微生物群落变化的环境驱动因子

植物根际微生物群落结构受多种环境因子的共同影响^[38,47-48]。相应地，根际土壤中的微生物也会调控土壤中营养元素的转化和循环^[49]。本研究采用冗余分析方法，探究了9种理化因子(电导率、pH、有效磷、有效钾、铵态氮、碱解氮、硝态氮、全氮、土壤有机碳)和5种土壤酶活(过氧化氢酶、脲酶、碱性磷酸酶、脱氢酶与蔗糖酶)对苍耳根际细菌和真菌群落的影响(图6)。研究结果显示，第一主轴和第二主轴共同解释了苍耳根际细菌群落总变化中的66.40% (图6A)；在各种环境因子中，pH (P<0.05)、有效磷(P<0.05)、碱解氮(P<0.05)、全氮(P<0.05)和铵态氮(P<0.05)均显著影响苍耳根际细菌群落。Chen等^[49]在对香樟(Cinnamomum camphora)土壤中的细菌群落分析中也发现，pH、有效磷、碱解氮对细菌群落结构具有显著性的影响。多项研究已经证实，pH是调控土壤细菌和真菌群落结果的关键因子^[37-39,50]。此外，脲酶活性、脱氢酶活性和蔗糖酶活性也能显著影响苍耳根际的细菌群落。Shi等研究报道了土壤酶在土壤的生化过程中发挥重要的作用，与土壤微生物群落之间存在密切的关系^[51]。对于真菌群落，冗余分析结果显示，第一主轴和第二主轴共同解释了苍耳根际真菌群落总变化中的72.40% (图6B)，比对细菌群落的解释比例更高。这可能是由于苍耳根际真菌群落更易受到消落区土壤环境因子的影响。除了电导率、铵态氮和土壤有机碳外，其余所有测试的变量均能显著影响苍耳根际真菌群落结构(P<0.05)。这与处于不同的生长阶段(幼期、中期、过熟期和衰退期)的胡杨(Populus euphratica)根际真菌群落的情况有所不同，后者主要受电导率的影响，其次是总可溶性盐和有效钾^[52]。在土壤酶活性方面，脲酶活性(P<0.05)、过氧化氢酶活性(P<0.05)和脱氢酶活性(P<0.05)显著影响了苍耳根际真菌群落，而碱性磷酸酶活性和蔗糖酶活性则未表现出显著影响。这些结果表明，苍耳真菌群落结构对土壤理化性质及酶活的变化是反应强烈的。Li等^[53]的研究也证实了土壤酶活与土壤微生物群落结构之间的相关性。

3 结论

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本研究选取了重庆市云阳县典型三峡水库消落区作为研究区域，针对该区域常见优势植物——苍耳的根际土壤微生物进，行了高通量测序分析。研究结果显示，在水淹胁迫强的XaRLL中，细菌和真菌的α多样性均高于水淹胁迫低的XaRHL。微生物群落的韦恩图分析及关键生物标志物识别结果也间接表明，XaRLL中微生物多样性略高于XaRHL。在细菌群落中，Proteobacteria是最为优势的菌门，且XaRLL中Proteobacteria的相对丰度高于XaRHL；而在真菌群落中，最为优势的菌门则有所不同，分别为Ascomycota和Basidiomycota。此外，这2个真菌菌门的相对丰度总和在XaRLL和XaRHL中是相近的。三峡水库消落区苍耳根际微生物群落易受其环境因子的影响，冗余分析结果显示，第一主轴和第二主轴分别解释了细菌群落和真菌群落变化中的66.40%和72.40%。本研究成果有望为理解水库消落区极端生境下优势植物如何通过组装根际微生物群落结构来适应逆境环境提供理论基础。

基金

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三峡后续规划研究项目(5000002021BF40001)

国家自然科学基金(42371071)

中国科学院先导专项A(XDA23040303)

参考文献

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文献

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doi: 10.13343/j.cnki.wsxb.20240580

接收时间：2024-09-19

首发时间：2026-02-05

出版时间：2025-02-04

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收稿日期：2024-09-19

录用日期：2024-10-30

基金

Scientific Research Project from Chongqing Water Resources Bureau(5000002021BF40001)

三峡后续规划研究项目(5000002021BF40001)

National Natural Science Foundation of China(42371071)

国家自然科学基金(42371071)

Leading Special Project A of Chinese Academy of Sciences(XDA23040303)

中国科学院先导专项A(XDA23040303)

作者信息

1 重庆交通大学河海学院，重庆

2 中国科学院重庆绿色智能技术研究院水库水环境重点实验室，重庆

3 中国科学院大学重庆学院，重庆

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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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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

Sample	pH	Soil organic matter (g/kg) **	Total N (g/kg) **	Alkaline hydrolyzable N (g/kg) **	Available P (mg/kg) **	AmmoniumN (mg/kg) **	Nitrate N (mg/kg) **	Available K (mg/kg) *	Conductivity (μs/cm) **
XaRHL	8.82±0.01	22.24±0.25	1.03±0.02	41.53±1.20	10.34±0.15	4.60±0.39	4.66±0.10	233.80±2.86	127.73±0.55
XaRLL	8.75±0.03	15.43±0.48	1.38±0.03	64.99±2.40	22.31±0.54	6.84±0.52	18.95±0.32	246.78±3.02	88.53±0.31

Sample

Soil organic matter (g/kg)

Total N (g/kg)

Alkaline hydrolyzable N (g/kg)

Available P (mg/kg)

AmmoniumN (mg/kg)

Nitrate N (mg/kg)

Available K (mg/kg)

Conductivity (μs/cm)

XaRHL

8.82±0.01

22.24±0.25

1.03±0.02

41.53±1.20

10.34±0.15

4.60±0.39

4.66±0.10

233.80±2.86

127.73±0.55

XaRLL

8.75±0.03

15.43±0.48

1.38±0.03

64.99±2.40

22.31±0.54

6.84±0.52

18.95±0.32

246.78±3.02

88.53±0.31

Sample	Urease ((μg/g)/24 h) **	Catalase ((μmol/g)/h) **	Alkaline phosphatase ((mg/g)/24 h) **	Dehydrogenase ((μL/g)/6 h) *	Sucrase ((mg/g)/24 h) **
XaRHL	185.65±12.32	2 288.08±12.89	0.84±0.03	0.006 2±0.000 1	8.68±0.49
XaRLL	1 641.12±66.64	2 661.86±28.92	1.65±0.03	0.007 5±0.000 3	13.77±0.18

Sample

Urease ((μg/g)/24 h)

Catalase ((μmol/g)/h)

Alkaline phosphatase ((mg/g)/24 h)

Dehydrogenase ((μL/g)/6 h)

Sucrase ((mg/g)/24 h)

XaRHL

185.65±12.32

2 288.08±12.89

0.84±0.03

0.006 2±0.000 1

8.68±0.49

XaRLL

1 641.12±66.64

2 661.86±28.92

1.65±0.03

0.007 5±0.000 3

13.77±0.18

Sample	Microbe	Chao1	Faith_pd	Observed_features	Shannon	Simpson
XaRHL	Bacteria	2 998.87±186.76	161.84±10.59	2 977.33±185.71	10.15±0.21	0.996 0±0.00
	Fungi	579.00±197.61	87.37±21.86	579.00±197.61	4.58±1.06	0.850 0±0.08
XaRLL	Bacteria	3 014.49±315.97	172.50±19.66	2 991.33±313.29	10.46±0.08	0.998 0±0.00
	Fungi	763.67±76.88	109.66±9.31	763.67±76.88	6.25±0.96	0.923 0±0.08

Sample

Microbe

Chao1

Faith_pd

Observed_features

Shannon

Simpson

XaRHL

Bacteria

2 998.87±186.76

161.84±10.59

2 977.33±185.71

10.15±0.21

0.996 0±0.00

Fungi

579.00±197.61

87.37±21.86

579.00±197.61

4.58±1.06

0.850 0±0.08

XaRLL

Bacteria

3 014.49±315.97

172.50±19.66

2 991.33±313.29

10.46±0.08

0.998 0±0.00

Fungi

763.67±76.88

109.66±9.31

763.67±76.88

6.25±0.96

0.923 0±0.08