Effect of arbuscular mycorrhizal fungi and rhizobium inoculation on soil fungal community structure and function in the rhizosphere of Medicago sativa
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摘要:
在我国，优良豆科牧草紫花苜蓿(Medicago sativa)一直处于供不应求的状态，接种AM (arbuscular mycorrhizae)真菌和根瘤菌可以促进土壤氮、磷循环以及提高植株抗逆性，从而提高苜蓿产量。为探究接种AM真菌和根瘤菌对苜蓿根际真菌群落结构和功能的影响，采集苜蓿6个不同处理组(不接种、单接种摩西球囊霉、单接种根内球囊霉、单接种根瘤菌、双接种根内球囊霉和根瘤菌、双接种摩西球囊霉和根瘤菌)根际和非根际土壤样品，基于真菌ITS区高通量测序技术，分析比较不同接种处理组苜蓿根际、非根际土壤中真菌多样性和群落分布的规律，并采用FUNGuild软件对不同处理组间菌群功能进行预测。结果表明：6个不同处理组中，在门水平上土壤真菌群落主要由子囊菌门、担子菌门构成，且以子囊菌门为主(19.29%～61.18%)。在属水平上发现与不接种相比，接种AM真菌和根瘤菌显著提高了有益菌的丰度，如单接种摩西球囊霉GMR处理组中优势属为支顶孢属(Acremonium)、单接种根瘤菌KR处理组中优势属为葡萄穗霉属(Stachybotrys)和毛束霉属(Trichurus)，双接种根内球囊霉和根瘤菌GIKR处理组中优势属为木霉属(Trichoderma)，双接种摩西球囊霉和根瘤菌GMKR处理组中优势属为曲霉菌属(Aspergillus)。与不接种相比，接种AM真菌和根瘤菌提高了真菌群落的操作分类单元数(OTUs)、Shannon-Wiener指数、菌种丰富度Chao1指数。不同接种处理间主要以腐生营养型为主，接种AM真菌和根瘤菌均会增加共生营养型的相对丰度，降低病原营养型的相对丰度。综上所述，接种AM真菌和根瘤菌后，有利于土壤氮磷的循环，增加了土壤真菌群落的丰富度与多样性，在致病菌存在的同时增加了生防菌，以及与固氮、溶磷有关菌的丰度，增加了共生营养型的相对丰度，降低了病原营养型的相对丰度，故接种 根瘤菌丰富了真菌群落组成，优化了原有的微生物生态环境，从而有利于苜蓿植株品质及产量的提高。
Abstract:In China, the yield of Medicago sativa, an excellent legume forage, has been in short supply. Inoculation with AM fungi and rhizobium can promote soil nitrogen and phosphorus cycling and increase stress resistance to increase M. sativa yield. In order to investigate the effects of AM fungi and Rhizobium on the structure and function of rhizosphere fungal community, in this experiment, the rhizosphere and non-rhizosphere soil samples of six different treatment groups (Without inoculation,only inoculation with Glomus mosseae, only inoculation with Glomus intraradices, only inoculation Sinorhizobium meliloti, double inoculation with Glomus intraradices and Sinorhizobium meliloti, double inoculation with Glomus mosseae and Sinorhizobium meliloti) were collected, based on the high-throughput sequencing technology of the fungal ITS region, the laws of fungal diversity and community distribution in the rhizosphere and non-rhizosphere soils of M. sativa in different inoculation treatment groups were analyzed and compared, the FUNGuild software was used to predict the flora function between different treatment groups. The results show, in 6 different treatment groups, the soil fungal community was mainly composed of Ascomycota and Basidiomycota, with Ascomycota as the main species (19.29%～61.18%). It was found at the genus level that inoculation with AM fungi and rhizobium significantly increased the abundance of beneficial fungi compared with no inoculation. For example, the dominant genus in the inoculation with Glomus mosseae GMR treatment group was Acremonium, and the dominant genus in the only inoculation Sinorhizobium meliloti KR treatment group were Stachybotrys and Trichurus, the dominant genus in the double inoculation with Glomus intraradices and Sinorhizobium meliloti GIKR treatment group was Trichoderma, and the dominant genus in the double inoculation with Glomus mosseae and Sinorhizobium meliloti GMKR treatment group was Aspergillus. And inoculation with AM fungi and rhizobium significantly increased the OTUs, Shannon index, and Chao1 index of the fungal community. Among different inoculation treatments, saprophytic types were dominant type. Inoculation of AM fungi and rhizobia increased the relative abundance of symbiotic types and decreased the relative abundance of pathogenic types. To sum up, the inoculation of AM fungi and rhizobia is conducive to the circulation of soil nitrogen and phosphorus, increases the richness and diversity of soil fungal community, increases the abundance of biocontrol bacteria, nitrogen fixation and phosphorus solubilizing bacteria, increases the relative abundance of symbiotic type, and reduces the relative abundance of pathogenic type. Therefore, inoculation with AM fungi and rhizobium enriched fungal community composition and optimized the original microbial ecological environment, which was beneficial to the improvement of M. sativa quality and yield.
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图 1 不同接种处理下苜蓿土壤根际真菌门水平的相对丰度弦图
a, Ascomycota 子囊菌门; b, Mortierellomycota 被孢霉门; c, Basidiomycota 担子菌门; d, Mucoromycota 毛霉菌门; e, Glomeromycota 球囊菌门; f, Aphelidiomycota; g, Chytridiomycota 壶菌门; h, Rozellomycota 罗兹菌门; i, Calcarisporiellomycota; j, Blastocladiomycota 芽枝霉门; k, others其他.
Figure 1. Relative abundance chord diagram of rhizosphere fungal division levels of alfalfa under different inoculation treatments
图 2 不同接种处理下苜蓿土壤真菌群落基于属水平上的聚类分析
Figure 2. Cluster analysis of alfalfa soil fungal community based on genus level under different inoculation treatments
图 3 不同接种处理苜蓿土壤真菌功能多样性热图
Figure 3. Heat map of fungal functional diversity in alfalfa soil under different inoculation treatments
表 1 苜蓿各试验组样本土壤的理化因子
Table 1. Value for soil physical and chemical factors of each testing group sample
样品分组
Group namepH 有机质含量
OM content/
(g·kg−1)全氮含量
TN content/
(g·kg−1)全磷含量
TP content/
(g·kg−1)有效磷含量
AP content/
(mg·kg−1)碱解氮含量
AN content/
(g·kg−1)过氧化氢酶活性
CAT activity/
(U·mL−1)中性磷酸酶活性NP activity/
(U·L−1)CKTR 8.19 ± 0.00abcd 29.10 ± 0.39de 0.73 ± 0.14a 1.47 ± 0.06a 31.85 ± 1.35ab 57.93 ± 2.33cd 0.38 ± 0.06ab 0.45 ± 0.02f CKTB 8.12 ± 0.01d 47.35 ± 0.40a 0.67 ± 0.03bc 1.34 ± 0.02bc 23.80 ± 2.57cd 63.67 ± 3.24bc 0.34 ± 0.04cd 0.25 ± 0.05i GIR 8.28 ± 0.01a 22.10 ± 0.25g 0.57 ± 0.05de 1.15 ± 0.09de 18.33 ± 3.66e 59.00 ± 3.16cd 0.33 ± 0.03cd 0.47 ± 0.01e GIB 8.22 ± 0.01abc 20.93 ± 0.45g 0.51 ± 0.32f 1.03 ± 0.02f 27.82 ± 5.89bc 58.83 ± 2.56cd 0.22 ± 0.01f 0.24 ± 0.03g GMR 8.22 ± 0.00abc 32.73 ± 0.36bc 0.52 ± 0.24f 1.05 ± 0.08f 22.64 ± 3.67de 58.50 ± 2.79cd 0.37 ± 0.02b 0.55 ± 0.01c GMB 8.26 ± 0.00ab 21.76 ± 0.55g 0.56 ± 0.09ef 1.12 ± 0.05ef 23.65 ± 4.44cd 73.50 ± 3.87a 0.27 ± 0.02e 0.22 ± 0.02k KR 8.16 ± 0.01bcd 35.96 ± 0.53b 0.59 ± 0.06de 1.19 ± 0.03de 32.00 ± 5.35ab 66.00 ± 5.21b 0.40 ± 0.03a 0.58 ± 0.03b KB 8.17 ± 0.00bcd 25.94 ± 0.46ef 0.71 ± 0.01ab 1.41 ± 0.04ab 31.71 ± 6.65ab 56.00 ± 4.23d 0.32 ± 0.01d 0.40 ± 0.01g GIKR 8.14 ± 0.00cd 26.77 ± 0.37ef 0.70 ± 0.06ab 1.39 ± 0.08ab 31.13 ± 4.34ab 58.17 ± 4.67cd 0.34 ± 0.04cd 0.60 ± 0.02ab GIKB 8.17 ± 0.01bcd 23.38 ± 0.42fg 0.72 ± 0.01a 1.44 ± 0.07a 35.74 ± 5.42a 65.33 ± 5.33b 0.34 ± 0.01c 0.31 ± 0.04h GMKR 8.25 ± 0.01ab 31.76 ± 0.40cd 0.66 ± 0.06bc 1.32 ± 0.09bc 21.21 ± 6.67de 68.33 ± 3.47ab 0.34 ± 0.03cd 0.60 ± 0.01a GMKB 8.23 ± 0.00abc 25.70 ± 0.44ef 0.62 ± 0.01cd 1.24 ± 0.07cd 21.35 ± 3.84de 68.30 ± 3.18bc 0.34 ± 0.01cd 0.50 ± 0.03d CKTR：不接种处理组根际土壤；CKTB：不接种处理组非根际土壤；GIR：单接种根内球囊霉处理组根际土壤；GIB：单接种根内球囊霉处理组非根际土壤；GMR：单接种摩西球囊霉处理组根际土壤；GMB：单接种摩西球囊霉处理组非根际土壤；KR：单接种根瘤菌处理组根际土壤；KB：单接种根瘤菌处理组非根际土壤；GIKR：双接种根内球囊霉和根瘤菌处理组根际土壤；GIKB：双接种根内球囊霉和根瘤菌处理组非根际土壤；GMKR：双接种摩西球囊霉和根瘤菌处理组根际土壤；GMKB：双接种摩西球囊霉和根瘤菌处理组非根际土壤；下同。OM：有机质；TN：全氮；TP：全磷；AP：有效磷；AN：碱解氮；CAT：过氧化氢酶；NP：中性磷酸酶；同列不同小写字母表示不同样品分组之间差异显著(P < 0.05)；下同。
CKTR: Rhizosphere soil of non-inoculation treatment group; CKTB: Non-rhizosphere soil in non-inoculation treatment group; GIR: Rhizosphere soil of only inoculation withGlomus intraradices treatment group; GIB: Non-rhizosphere soil of only inoculation withGlomus intraradices treatment group; GMR: Rhizosphere soil of only inoculationGlomus mosseae treatment group; GMB: Non-rhizosphere soil of only inoculation withGlomus mosseaetreatment group; KR: Rhizosphere soil of only inoculation Sinorhizobium melilotitreatment group; KB: Non-rhizosphere soil of only inoculation withSinorhizobium melilotitreatment group; GIKR: Rhizosphere soil of double inoculation with Glomus intraradicesand Sinorhizobium meliloti treatment group; GIKB: Non-rhizosphere soil of double inoculation with Glomus intraradicesand Sinorhizobium meliloti treatment group; GMKR: Rhizosphere soil of double inoculation with Glomus mosseaeand Sinorhizobium meliloti treatment group; GMKB: Non-rhizosphere soil of double inoculation with Glomus mosseaeand Sinorhizobium melilotitreatment group; this is applicable for the following figures and tables as well. OM: organic matter; TN: total nitrogen; TP: total phosphorus; AP: available phosphorus; AN: alkali nitrogen; CAT: Catalase; NP: Neutral phosphatase. Different lowercase letters within the same column indicate significant differences between different sample groups at the 0.05 level; this is applicable for the following figures as well.
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表 2 不同接种处理组苜蓿根际、非根际土壤真菌丰度和多样性指数
Table 2. Fungal abundance and diversity index of alfalfa rhizosphere and non-rhizosphere soil
样品分组
Group nameOTUs数
OTUsShannon-Wiener指数
Shannon-Wiener indexChao1 指数
Chao1 index覆盖率
Coverage/%CKTR 908.01 ± 55.91c 5.76 ± 0.09a 1101.42 ± 16.78c 0.993 0 ± 0.000 1a CKTB 991.11 ± 53.21c 6.51 ± 0.33a 1212.90 ± 16.45bc 0.991 0 ± 0.000 3a GIR 1101.08 ± 78.32bc 6.31 ± 0.12a 1376.79 ± 12.34bc 0.991 0 ± 0.000 1a GIB 1041.35 ± 45.31c 5.95 ± 0.16a 1368.43 ± 14.23bc 0.990 0 ± 0.000 1a GMR 1077.24 ± 65.16c 6.35 ± 0.19a 1361.18 ± 15.32bc 0.990 0 ± 0.000 2a GMB 2633.17 ± 123.51a 7.27 ± 0.35a 4967.61 ± 33.62a 0.996 0 ± 0.000 1a KR 2083.26 ± 136.80ab 7.57 ± 0.21a 3272.07 ± 16.34ab 0.982 0 ± 0.000 3a KB 1025.35 ± 64.78c 5.76 ± 0.16a 1291.18 ± 18.95bc 0.991 0 ± 0.000 1a GIKR 1254.26 ± 63.21bc 6.70 ± 0.31a 1613.12 ± 23.16bc 0.989 0 ± 0.000 3a GIKB 1090.32 ± 79.32bc 6.67 ± 0.15a 1420.70 ± 25.14bc 0.990 0 ± 0.000 1a GMKB 1070.17 ± 78.32bc 6.43 ± 0.18a 1346.61 ± 31.21bc 0.990 0 ± 0.000 1a GMKR 1099.11 ± 64.21bc 6.31 ± 0.32a 1409.56 ± 22.78bc 0.990 0 ± 0.000 2a
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