羊肚菌连作对土壤理化性质、微生物群落变化的影响及其关联机制

doi:10.13560/j.cnki.biotech.bull.1985.2025-1093

摘要/Abstract

摘要：

目的羊肚菌连续种植引发的连作障碍问题会导致土壤理化和微生态环境恶化、病虫害加剧、生产效益降低等问题，解析羊肚菌连作过程土壤理化性质及微生物群落的变化及其相关性，为破解羊肚菌连作障碍难题和制定高效防控技术提供理论支持。方法以未种植羊肚菌、连作0-3年羊肚菌的根际土壤为研究对象。测定各种植年限羊肚菌出菇后土壤理化环境因子，基于16S rRNA和ITS序列测定土壤微生物组成，解析其多样性及群落结构。结果羊肚菌种植过程中显著提高了硝态氮、交换性钙、有效磷、电导率和pH值，有效铜和有效铁显著下降；与未种植羊肚菌相比，连作显著降低了土壤细菌和真菌的多样性和丰富度；其中变形菌门和酸杆菌门在羊肚菌种植后相对丰度增加，放线菌门和拟杆菌门则降低；子囊菌门、担子菌门和被孢霉门随羊肚菌连作下降，毛霉门相对丰度增加。16种差异菌属产生变化，其中酸杆菌Gp4、青霉菌属和四枝孢霉属显著增加，维希尼克氏酵母属、被孢霉属、树粉孢属、篮状菌属和毛枝霉属显著降低。羊肚菌连作通过增强细菌关联网络并削弱真菌关联网络，重塑了根际土壤微生物群落的共现格局；而且土壤pH、交换性钙、有效锌和电导率与差异菌属存在显著相关性。结论羊肚菌连作显著改变土壤理化因子和土壤微生物群落结构及共现性特征，且土壤微生物群落与土壤理化性质存在一定关联，它们之间的关联性随种植年限的不同而产生差异，两者相关性的恶化可能是羊肚菌连作障碍发生的主要因素。

关键词: 羊肚菌, 连作障碍, 土壤理化性质, 微生物群落, 土壤-微生物关联

Abstract:

Objective Continuous cropping of Morchella leads to a series of problems, including deterioration of soil physicochemical properties and micro-ecological environments, intensified pest and disease outbreaks, and reduced production efficiency. Elucidating the changes in soil physicochemical properties and microbial communities during continuous Morchella cultivation and their correlations, offers a theoretical basis for developing strategies to overcome continuous cropping obstacles. Methods To investigate the impact of continuous cropping, we collected rhizosphere soils from plots with no Morchella (control) and 0–3 consecutive years of Morchella cultivation. We assessed soil physicochemical properties post-fruiting and characterized microbial composition, diversity, and communities structure through 16S rRNA and ITS sequencing. Results Morchella cultivation significantly elevated levels of nitrate nitrogen, exchangeable calcium, available phosphorus, electrical conductivity, and pH, while in parallel, it markedly depleted available copper and iron. Compared to non-cultivated plots, the diversity and abundance of soil bacteria and fungi were significantly reduced in continuous-cropping plots. Specifically, the relative abundance of the Proteobacteria and Acidobacteria phyla increased after Morchella cultivation, whereas that of the Actinobacteria and Bacteroidetes phyla decreased. Ascomycota, Basidiomycota, and Mortierellomycota declined with continuous Morchella cultivation, while the relative abundance of Mucoromycota increased. Changes were observed in 16 different bacterial genera, with significant increases in Acidobacteria_Gp4, Penicillium, and Tetramonas, and significant decreases in Vishniacozyma, Mortierella, Oidiodendron, Talaromyces and Trichocladium. Continuous cropping of Morchella reshaped the co-occurrence patterns of rhizosphere soil microbial communities by strengthening bacterial association networks while weakening fungal association networks. Furthermore, soil pH, exchangeable calcium, available zinc, and electrical conductivity exhibited significant correlations with the differential microbial genera. Conclusion Continuous Morchella cultivation significantly alters soil physicochemical factors, microbial communities structure and its co-occurrence patterns. Moreover, a significant correlation was observed between the soil microbial communities and soil physicochemical properties, and this correlation varied with the duration of cropping. The subsequent deterioration of this interplay may be a primary factor driving the occurrence of continuous cropping obstacles in Morchella.

Key words: Morchella, continuous cropping, soil physicochemical properties, microbial communities, soil-microbe interactions

廖艳婷, 王灿琴, 韦娇君, 赵承刚, 黄世旅, 罗阳兰, 阎勇. 羊肚菌连作对土壤理化性质、微生物群落变化的影响及其关联机制[J]. 生物技术通报, 2026, 42(5): 134-146.

LIAO Yan-ting, WANG Can-qin, WEI Jiao-jun, ZHAO Cheng-gang, HUANG Shi-lyu, LUO Yang-lan, YAN Yong. Changes in Soil Physicochemical Properties and Microbial Communities under Continuous Cropping of Morchella and Their Associated Mechanisms[J]. Biotechnology Bulletin, 2026, 42(5): 134-146.

图/表 8

图1 羊肚菌连作3年土壤理化性质变化CK：未种植；NH4+-N：铵态氮；NO3--N：硝态氮；AK：速效钾；Exc.Ca：交换性钙；Exc.Mg：交换性镁；Exc.Na：交换性钠；AP：有效磷；AZn：有效锌；ACu：有效铜；AFe：有效铁；AMn：有效锰；AB：有效硼；EC：电导率；OM：有机质。不同小写字母代表组间差异显著（P<0.05）

Fig. 1 Changes in soil physicochemical properties after three years of continuous Morchella croppingCK: Unplanted; NH4+-N: ammonium nitrogen; NO3--N: nitrate nitrogen; AK: available potassium; Exc.Ca:exchangeable calcium; Exc.Mg:exchangeable magnesium; Exc.Na: exchangeable sodium; AP: available phosphorus; AZn: available zinc; ACu: effective copper; AFe: available iron; AMn: available manganese; AB: effective boron; EC: electrical conductivity; OM: organic matter. Different lowercase letters indicate significant differences between groups (P<0.05)

表1 羊肚菌连作0-3年土壤理化性质变化率

Table 1 Rate of change in soil physicochemical properties from 0 to 3 years of continuous Morchella cropping

土壤理化性质变化率 Rate of change in soil physicochemical properties （%）	连作年限 Continuous cropping period（Year）
	0	1	2	3
铵态氮 NH₄⁺-N	-77.76	-71.62	-15.23	15.38
硝态氮 $N O 3 -$ -N	-34.04	-7.61	-3.03	385.75
速效钾 Available potassium	23.34	-24.14	-3.33	19.93
交换性钙 Exchangeable calcium	40.3	13.72	24.63	23.34
交换性镁 Exchangeable magnesium	16.31	-26.81	13.33	-14.42
交换性钠 Exchangeable sodium	-20.25	-28.32	46.48	28.73
有效磷 Available phosphorus	-56.3	46.54	-19.14	15.88
有效锌 Available zinc	-47.46	-21.2	-44.95	-16.36
有效铜 Available copper	-7.84	-22.64	-22.92	-18.4
有效铁 Available iron	-45.95	-31.85	-59.63	-58.28
有效锰 Available manganese	142.2	-13.89	7.96	-47.94
有效硼 Effective boron	192.26	15.55	25.04	17.25
电导率 Electrical conductivity	200.56	123	235.57	207.95
有机质 Organic matter	24.63	-5.49	5.03	9.17
pH	17.02	12.73	15.89	11.94

表1 羊肚菌连作0-3年土壤理化性质变化率

Table 1 Rate of change in soil physicochemical properties from 0 to 3 years of continuous Morchella cropping

土壤理化性质变化率 Rate of change in soil physicochemical properties （%）	连作年限 Continuous cropping period（Year）
	0	1	2	3
铵态氮 NH₄⁺-N	-77.76	-71.62	-15.23	15.38
硝态氮 $N O 3 -$ -N	-34.04	-7.61	-3.03	385.75
速效钾 Available potassium	23.34	-24.14	-3.33	19.93
交换性钙 Exchangeable calcium	40.3	13.72	24.63	23.34
交换性镁 Exchangeable magnesium	16.31	-26.81	13.33	-14.42
交换性钠 Exchangeable sodium	-20.25	-28.32	46.48	28.73
有效磷 Available phosphorus	-56.3	46.54	-19.14	15.88
有效锌 Available zinc	-47.46	-21.2	-44.95	-16.36
有效铜 Available copper	-7.84	-22.64	-22.92	-18.4
有效铁 Available iron	-45.95	-31.85	-59.63	-58.28
有效锰 Available manganese	142.2	-13.89	7.96	-47.94
有效硼 Effective boron	192.26	15.55	25.04	17.25
电导率 Electrical conductivity	200.56	123	235.57	207.95
有机质 Organic matter	24.63	-5.49	5.03	9.17
pH	17.02	12.73	15.89	11.94

图2 羊肚菌连作3年土壤根际微生物群落多样性变化A：细菌Chao1指数和Shannon指数；B：真菌Chao1指数和Shannon指数；C：细菌主坐标分析图；D：真菌主坐标分析图

Fig. 2 Changes in the diversity of rhizosphere soil microbial communities after three years of continuous Morchella croppingA: The Chao1 index and Shannon index of bacteria. B: The Chao1 index and Shannon index of fungi. C: The principal coordinate analysis (PCoA) of bacteria. D: The principal coordinate analysis (PCoA) of fungi

图3 羊肚菌连作3年土壤根际微生物物种丰度柱状堆叠图A：细菌在门水平上的物种分布；B：细菌属水平上的物种分布；C：真菌在门水平上的物种分布；D：真菌在属水平上的物种分布

Fig. 3 Column stacking of species abundance of soil rhizosphere microorganisms in 3 years of continuous cropping of MorchellaA: Species distribution of bacteria at the phylum level. B: Species distribution of bacteria at the genus level.C: Species distribution of fungi at the phylum level. D: Species distribution of fungi at the genus level

图4 羊肚菌连作条件下根际土壤细菌和真菌差异菌属

Fig. 4 Differential bacterial and fungal genera in rhizosphere soil under continuous Morchella cropping

图5 羊肚菌连作条件下根际土壤微生物群落共现性网络图A：细菌；B：真菌

Fig. 5 Co-occurrence network diagram of rhizosphere soil microbial communities under continuous Morchella croppingA: Bacteria. B: Fungi

表2 不同处理对土壤微生物群落网络拓扑指数的影响

Table 2 Effects of different treatments on topological indices of soil microbial community networks

物种分类 Taxonomy of species	处理 Treatment	节点数 Number of nodes	边数 Number of edges	平均度 Average degree	网络密度 Network density	模块化 Modularity
细菌 Bacteria	CK	241	2 654	22.025	0.092	0.518
	0Y	311	4 883	31.402	0.101	0.523
	1Y	359	6 091	33.933	0.095	0.510
	2Y	284	3 678	25.901	0.092	0.491
	3Y	408	9 592	47.02	0.116	0.476
真菌 Fungi	CK	226	1 845	16.327	0.073	0.540
	0Y	90	364	8.089	0.091	0.586
	1Y	134	785	11.716	0.088	0.582
	2Y	155	877	11.316	0.073	0.595
	3Y	104	495	9.519	0.092	0.602

图6 羊肚菌连作土壤微生物群落与理化环境因子的相关性分析A：羊肚菌连作土壤差异菌属与环境因子的配对比较矩阵图。方格颜色深浅度表示皮尔森相关系数大小，连线颜色深浅和粗细表示微生物矩阵与环境因子的Mantel检验相关性，颜色越深、线条越粗表示相关性越强（*P<0.05；**P<0.01；***P<0.001）。spec01：鞘氨醇黄单胞菌属、土地杆菌、砂单胞菌属、青霉菌属和四枝孢霉属；spec02：马赛菌属、毛枝霉属、树粉孢属和篮状菌属；spec03：酸杆菌GP4、酸杆菌GP7和腐质霉属；spec04：Gaiella、溶杆菌属、维希尼克氏酵母属和被孢霉属。B：细菌冗余分析图；C：真菌冗余分析图

Fig. 6 Correlation analysis between soil microbial communities and physicochemical environmental factors under continuous Morchella croppingA: Matrix plot of pairwise comparisons between differential genera and environmental factors in soils under continuous Morchella cropping. The color intensity of the squares indicates the magnitude of the Pearson correlation coefficient, while the shade and thickness of the lines indicate the strength of the Mantel test correlation between the microbial matrix and environmental factors. Darker colors and thicker lines denote stronger correlations(*P<0.05;**P<0.01;***P<0.001). spec01: Sphingomonas、Pedobacter、Arenimonas、Penicillium and Tetracladium; spec02: Massilia, Trichocladium, Oidiodendron and Talaromyces; spec03: Acidobacteria_Gp4、Acidobacteria_Gp7 and Humicola; spec04: Gaiella, Lysobacter, Vishniacozyma and Mortierella.B: Bacterial redundancy analysis plot. C: Fungal redundancy analysis plot

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