保护地土壤中AM真菌群落结构与功能的初步研究
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摘要
作为重要环境与功能菌物,丛枝菌根(arbuscular mycorrhizas, AM)真菌通过与植物根系建立具有特定形态结构和功能的互惠共生体,对促进农、林、牧业生产、维持和提高生物多样性、生态系统生产力、环境安全性和可持续发展做出了巨大贡献。在自然条件下,AM真菌是以其独特的群落结构特征和功能,直接和间接发挥着各种生理、生化和生态作用。保护地栽培中,AM真菌修复连作退化土壤和生物防治土传病害潜力巨大。土壤中的生物往往是以一定群落结构形式而存在和发挥作用的。因此,从群落结构的角度研究其群落结构特征与功能会更有实际意义。然而,对保护地AM真菌群落结构及其生理生态功能的了解目前还十分有限。本研究采用传统形态学方法和分子技术,探索北京大兴、山东昌乐和山东莱阳保护地不同连作年限(5,10,15和20年)土壤和非保护地自然农田土壤中AM真菌群落结构特征,以非保护地自然土壤中AM真菌群落结构特征为依据,设计不同的AM真菌群落组合,初步探索不同AM真菌群落组合对保护地植物健康、土壤质量、保护地生态系统的生产力的贡献,以其为进一步研发专一的AM真菌群落结构制剂及其在保护地无公害蔬菜生产中的应用提供理论依据和技术支持。主要研究结果如下：
1.我国北方保护地土壤具有独特的AM真菌群落结构特征,其水平结构特征表现在各样地之间AM真菌群落构成差异明显：球囊霉属(Glomus)是各样地的优势属。幼套球囊霉(G. etunicatum)是北京大兴和山东昌乐样地连作5年的土壤中共同优势种。近明球囊霉(G. claroideum)是山东昌乐和莱阳样地连作5年和10年的土壤中共同优势种,浅窝无梗囊霉(Acaulospora. lacunosa)是北京大兴样地连作10年的土壤中的优势种。3个样地连作15和20年的土壤中的优势种并不明显。3个样地4个连作年限共有的菌种为近明球囊霉(G. claroideum); 3个样地连作5年之间相似性系数在0.67到0.84,而3个样地连作20年之间的相似性系数在0.29到0.33之间。垂直结构特征表现为：随着土壤深度的增加,3个样地(北京大兴、山东昌乐和莱阳)的AM真菌最大孢子密度、种丰度和多样性指数均出现在0-10 cm土层,且随土壤剖面深度增加呈现递减的趋势；时间结构特征表现为：6月和10月孢子密度、丰度、Shannon指数最高,8月和12月孢子密度最低。
2.PCR-DGGE图谱分析表明,山东昌乐保护地不同连作年限的西瓜土壤AM真菌群落结构组成及变化明显。共获得28条DNA序列,保护地土壤中AM真菌种的丰度和多样性指数较低,随着连作年限的增加,种的丰度减少。Shannon指数随种植年限增加呈现先下降后趋于缓和的变化。典范对应分析(CCA)结果显示CCA的第1排序轴和第2排序轴的物种与环境关系的累积率达67.6%,说明土壤性质对AM真菌群落组成具有重要的影响。土壤速效磷含量和种植年限是影响AM真菌群落组成的重要因素。
3.接种AM真菌群落盆栽试验结果表明,AM真菌群落显著增加了2-5 mm,1-2 mm, 1-0.5 mm,0.5-0.25 mm土壤水稳性团聚体结构数量,降低了土壤电导率；提高了细菌和放线菌的数量,而真菌的数量显著降低,提高了脲酶、蔗糖酶和多酚氧化酶的活性；AM真菌群落能明显改善黄瓜的生长,增加株高、干重、最大叶面积,提高了黄瓜叶片的叶绿素相对含量、可溶性糖含量、根系活力,增强了对逆境的抵抗能力。这种效应显著高于单接种AM真菌处理和对照。与其他各AM真菌群落相比,以AM真菌群落组合F(Glomus etunicatum+G. mosseae+Gigaspora margarita+Acaulospora lcunosa)与G(Gaggregatum+G.etunicatum+G.mosseae+G.versiforme+Gi.margarita+A.lacunosa)对黄瓜生长和土壤质量的贡献更大。
作者在国内外首次系统研究了保护地连作土壤中AM真菌群落结构与功能,证实了AM真菌群落具有强大的生物修复退化土壤的功能。为今后进一步研发AM真菌群落组合制剂奠定了基础。
As an important environmental and functional fungi, arbuscular mycorrhizal (AM) fungi widely distribute in terrestrial ecosystems. They make great contributions in sustainable agriculture and forestry, and play a vital role in maintaining and enhancing biological diversity, and ecosystem productivity. Under natural conditions, AM fungi play various of physiological, biochemical and ecological roles directly and indirectly according to its unique community structure and function. AM fungi have huge potential for both soil bioremediation and biological control of soil borne diseases. Soil is a heterogeneous system including many microorganism strains, so it is more significant to study the composition effectiveness of multiple strains from the perspective of community. However, we know little about AM fungal community structure and Physiological functions in commercial greenhouse at present. In this paper, the community structure of arbuscular mycorrhizal fungi in commercial greenhouse soil at three locations in China (Daxing in Beijing, Changle and Laiyang in Shandong) was investigated using both morphological identification and denaturing gradient gel electrophoresis. Based on AM fungal community structure analysis in farmland soil, we designed different AM fungal community composition. The contribution to plant health, the soil quality, the ecosystems productivity in protected cultivation field caused by AM fungal community was examined.as few studies were carried out to determine the effects of AM fungal community on plant growth and soil quality. It will be helpful for providing an important theoretical basis and reference value for the environmental restoration and exploitation and application of AM fungal agents with high efficiency in the soil of commercial greenhouse. The main results showed that:
1. Horizontal structure showed that: Glomus was a dominant genus in the sampling sites, G.etunicatum was the dominant species in 5-year replanting soil of Daxing and Changle, G.claroideum was the dominant species in 5-year and 10-year replanting soil of Changle and Laiyang, A.lacunosa was the dominant species in 10-year replanting soil of Daxing, the dominant species in 15-year and 20-year replanting soil was not obvious. G.claroideum was present in replanting soil at all sites. The Sorenson's similarity coefficients of AM fungal community composition ranged from 0.67 to 0.84 in 5-year replanting soil, and 0.29 to 0.33 in 10-year replanting soil. Vertical structure showed that: the spore abundance, species richness and Shannon index were highest in the topsoils (0-10 cm), and decreased with soil depth in all soils investigated. Time structure showed that: the spore density, species richness and Shannon index were greatest in June and October. Species richness and Shannon index were lower in August and December.
2. PCR-DGGE analytical results suggested that Shannon indice was first declined and then tending to mitigate. Species richness and Shannon index were lower in the soil of commercial greenhouse, species richness decreased with the increasing of replanting years The canonical correspondence analysis (CCA) showed that the two first axes, indicators of closely relationship between the soil properties and AM fungal community, explained 67.6% of the total variance. Available P and replanting years were main factors contributing to the variance of the AM fungal community composition.
3 The greenhouse experiments showed that water-stable aggregate fractions in 2-5,1-2,1-0.5 and 0.5-0.25 mm were generally improved in the tested degraded soil when treated with AM fungal communities, and soil salt ion concentration and soil EC were decreased as well. No significant changes were found in soil pH. Inoculation with AM fungal communities increased the number of bacteria and actinomycetes, but reduced the number of fungi. AM fungal communities apparently increased plant growth and dry mass, the soluble sugar content, Chlorophyll content and root activity of cucumber plants when compared to non-mycorrhizal or single-inoculation treatments. Among all inoculation treatments, AM fungal community F (Glomus etunicatum + G. mosseae + Gigaspora margarita + Acaulospora lacunose) and AM fungal community G (G. aggregatum + G. etunicatum + G. mosseae + G. versiforme + Gi. margarita+A. lacunose) showed greater contribution to the improvement of soil quality and plant growth, indicating that inoculation with AM fungal community had more beneficial effects than with single or double species.
We studyed the community structure and function of arbuscular mycorrhizal fungi in the soil of commercial greenhouse at home and abroad for the first time, proved that AM fungal community had a function of soil bioremediation. This paper provided an important theoretical basis and reference value for the exploitation and application of AM fungal agents in the soil of commercial greenhouse.
1.我国北方保护地土壤具有独特的AM真菌群落结构特征,其水平结构特征表现在各样地之间AM真菌群落构成差异明显：球囊霉属(Glomus)是各样地的优势属。幼套球囊霉(G. etunicatum)是北京大兴和山东昌乐样地连作5年的土壤中共同优势种。近明球囊霉(G. claroideum)是山东昌乐和莱阳样地连作5年和10年的土壤中共同优势种,浅窝无梗囊霉(Acaulospora. lacunosa)是北京大兴样地连作10年的土壤中的优势种。3个样地连作15和20年的土壤中的优势种并不明显。3个样地4个连作年限共有的菌种为近明球囊霉(G. claroideum); 3个样地连作5年之间相似性系数在0.67到0.84,而3个样地连作20年之间的相似性系数在0.29到0.33之间。垂直结构特征表现为：随着土壤深度的增加,3个样地(北京大兴、山东昌乐和莱阳)的AM真菌最大孢子密度、种丰度和多样性指数均出现在0-10 cm土层,且随土壤剖面深度增加呈现递减的趋势；时间结构特征表现为：6月和10月孢子密度、丰度、Shannon指数最高,8月和12月孢子密度最低。
2.PCR-DGGE图谱分析表明,山东昌乐保护地不同连作年限的西瓜土壤AM真菌群落结构组成及变化明显。共获得28条DNA序列,保护地土壤中AM真菌种的丰度和多样性指数较低,随着连作年限的增加,种的丰度减少。Shannon指数随种植年限增加呈现先下降后趋于缓和的变化。典范对应分析(CCA)结果显示CCA的第1排序轴和第2排序轴的物种与环境关系的累积率达67.6%,说明土壤性质对AM真菌群落组成具有重要的影响。土壤速效磷含量和种植年限是影响AM真菌群落组成的重要因素。
3.接种AM真菌群落盆栽试验结果表明,AM真菌群落显著增加了2-5 mm,1-2 mm, 1-0.5 mm,0.5-0.25 mm土壤水稳性团聚体结构数量,降低了土壤电导率；提高了细菌和放线菌的数量,而真菌的数量显著降低,提高了脲酶、蔗糖酶和多酚氧化酶的活性；AM真菌群落能明显改善黄瓜的生长,增加株高、干重、最大叶面积,提高了黄瓜叶片的叶绿素相对含量、可溶性糖含量、根系活力,增强了对逆境的抵抗能力。这种效应显著高于单接种AM真菌处理和对照。与其他各AM真菌群落相比,以AM真菌群落组合F(Glomus etunicatum+G. mosseae+Gigaspora margarita+Acaulospora lcunosa)与G(Gaggregatum+G.etunicatum+G.mosseae+G.versiforme+Gi.margarita+A.lacunosa)对黄瓜生长和土壤质量的贡献更大。
作者在国内外首次系统研究了保护地连作土壤中AM真菌群落结构与功能,证实了AM真菌群落具有强大的生物修复退化土壤的功能。为今后进一步研发AM真菌群落组合制剂奠定了基础。
As an important environmental and functional fungi, arbuscular mycorrhizal (AM) fungi widely distribute in terrestrial ecosystems. They make great contributions in sustainable agriculture and forestry, and play a vital role in maintaining and enhancing biological diversity, and ecosystem productivity. Under natural conditions, AM fungi play various of physiological, biochemical and ecological roles directly and indirectly according to its unique community structure and function. AM fungi have huge potential for both soil bioremediation and biological control of soil borne diseases. Soil is a heterogeneous system including many microorganism strains, so it is more significant to study the composition effectiveness of multiple strains from the perspective of community. However, we know little about AM fungal community structure and Physiological functions in commercial greenhouse at present. In this paper, the community structure of arbuscular mycorrhizal fungi in commercial greenhouse soil at three locations in China (Daxing in Beijing, Changle and Laiyang in Shandong) was investigated using both morphological identification and denaturing gradient gel electrophoresis. Based on AM fungal community structure analysis in farmland soil, we designed different AM fungal community composition. The contribution to plant health, the soil quality, the ecosystems productivity in protected cultivation field caused by AM fungal community was examined.as few studies were carried out to determine the effects of AM fungal community on plant growth and soil quality. It will be helpful for providing an important theoretical basis and reference value for the environmental restoration and exploitation and application of AM fungal agents with high efficiency in the soil of commercial greenhouse. The main results showed that:
1. Horizontal structure showed that: Glomus was a dominant genus in the sampling sites, G.etunicatum was the dominant species in 5-year replanting soil of Daxing and Changle, G.claroideum was the dominant species in 5-year and 10-year replanting soil of Changle and Laiyang, A.lacunosa was the dominant species in 10-year replanting soil of Daxing, the dominant species in 15-year and 20-year replanting soil was not obvious. G.claroideum was present in replanting soil at all sites. The Sorenson's similarity coefficients of AM fungal community composition ranged from 0.67 to 0.84 in 5-year replanting soil, and 0.29 to 0.33 in 10-year replanting soil. Vertical structure showed that: the spore abundance, species richness and Shannon index were highest in the topsoils (0-10 cm), and decreased with soil depth in all soils investigated. Time structure showed that: the spore density, species richness and Shannon index were greatest in June and October. Species richness and Shannon index were lower in August and December.
2. PCR-DGGE analytical results suggested that Shannon indice was first declined and then tending to mitigate. Species richness and Shannon index were lower in the soil of commercial greenhouse, species richness decreased with the increasing of replanting years The canonical correspondence analysis (CCA) showed that the two first axes, indicators of closely relationship between the soil properties and AM fungal community, explained 67.6% of the total variance. Available P and replanting years were main factors contributing to the variance of the AM fungal community composition.
3 The greenhouse experiments showed that water-stable aggregate fractions in 2-5,1-2,1-0.5 and 0.5-0.25 mm were generally improved in the tested degraded soil when treated with AM fungal communities, and soil salt ion concentration and soil EC were decreased as well. No significant changes were found in soil pH. Inoculation with AM fungal communities increased the number of bacteria and actinomycetes, but reduced the number of fungi. AM fungal communities apparently increased plant growth and dry mass, the soluble sugar content, Chlorophyll content and root activity of cucumber plants when compared to non-mycorrhizal or single-inoculation treatments. Among all inoculation treatments, AM fungal community F (Glomus etunicatum + G. mosseae + Gigaspora margarita + Acaulospora lacunose) and AM fungal community G (G. aggregatum + G. etunicatum + G. mosseae + G. versiforme + Gi. margarita+A. lacunose) showed greater contribution to the improvement of soil quality and plant growth, indicating that inoculation with AM fungal community had more beneficial effects than with single or double species.
We studyed the community structure and function of arbuscular mycorrhizal fungi in the soil of commercial greenhouse at home and abroad for the first time, proved that AM fungal community had a function of soil bioremediation. This paper provided an important theoretical basis and reference value for the exploitation and application of AM fungal agents in the soil of commercial greenhouse.
引文
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