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梵净山地处热带、亚热带生物区系向温带生物区系的生态交错区,由于独特的地理位置、复杂的生物和生态演化过程、多样化的植物类群特征、众多地方特有和濒危动植物种类等特点[1-2],引起国内外不同领域学者的广泛关注,相关研究涉及到土壤理化性质、物种种类及多样性、珍稀濒危物种种群结构特征[2-5]等多个方面。关于地下微生物群落的研究则集中在丛枝菌根真菌[6-7],对真菌群落结构及功能类群特征尚无系统报道。真菌特别是某些功能类群在促进有机质分解和养分循环、增强植物对矿物质营养元素的吸收、提高植物对生物和非生物胁迫的耐受性、维持生态系统的稳定性和生产力等方面发挥着重要作用[8-10]。此外,真菌群落组成不仅受土壤理化性质、气候等非生物因素影响[11-12],其对植物种类、群落组成和多样性等[13-14]生物因素也较为敏感,而植被类型可通过影响非生物和生物因素作用于真菌群落。尤为特殊的是,不同植被类型还会塑造特定的真菌功能类群:Tedersoo等[11]通过对全世界范围内不同植被类型的土壤真菌群落进行分析后发现,外生菌根真菌在温带落叶林中较为丰富,热带潮湿雨林和热带干旱森林中植物病原菌更为多样;Liu等[15]发现在中国东部森林地区,温带森林以外生菌根真菌为主,热带和亚热带森林中具有丰富的丛枝菌根真菌和植物病原菌;Sheng等[16]研究了神农架山地森林生态系统中的土壤真菌功能类群,发现外生菌根真菌在落叶阔叶林、针阔混交林和针叶林中尤为丰富,植物病原菌-木腐生菌主要出现在常绿阔叶林,植物病原菌和寄生性真菌在山顶灌丛中具有较高相对丰度。菌在山顶灌丛中具有较高相对丰度。
梵净山拥有的典型植物区系过渡性及地球上同纬度保存完好的森林生态系统,为丰富且独特真菌种类的形成创造良好生存条件。此外,由于水热条件等差异造就了梵净山多样且分异明显的植被类型,为探讨植被类型与真菌群落间的关系提供了天然理想实验室。本研究以梵净山国家级自然保护区典型且具有代表性的常绿阔叶林、常绿落叶阔叶混交林及位于山顶附近的亚高山矮林3种森林植被类型为研究对象,分析真菌群落结构和功能类群特征与植被类型间的关系,探讨土壤性质、细根性状和叶性状对真菌群落的影响。研究结果可为后续探讨亚热带地区不同植被类型树种的环境适应机制及地上和地下部分的养分交换策略提供来自于地下真菌群落方面的数据支撑。
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48个根系样本共得到5 103个ASVs,属于9门、193科和385个属(不包括unidentified fungi)。子囊菌门(Ascomycota)和担子菌门(Basidiomycota)为优势门,占全部ASVs数的89.93%。小蔓毛壳科(Herpotrichiellaceae)、球囊霉科(Glomeraceae)和柔膜菌科(Helotiaceae)为优势科,晶杯菌科(Hyaloscyphaceae)、Sebacinales_Group_A、被孢霉科(Mortierellaceae)、皮盘菌科(Dermateaceae)、红菇科(Russulaceae)、小菇科(Mycenaceae)、丛赤壳科(Nectriaceae)、水盘菌科(Vibrisseaceae)、锤舌菌科(Leotiaceae)、肉座菌科(Hypocreaceae)和革菌科(Thelephoraceae)相对丰度超过1%。常绿阔叶林具有最多ASVs数(2 287个),子囊菌门为优势门,球囊霉科为优势科;亚高山矮林共得到ASVs数为1 636个,子囊菌门为优势门,柔膜菌科和晶杯菌科为优势科;常绿落叶阔叶混交林ASVs数最少,为1 608个,子囊菌门和担子菌门为优势门,球囊霉科和小蔓毛壳科为优势科。LEfSe分析(图1A)显示,常绿阔叶林显著差异类群为球囊菌门(Glomeromycota)、球囊菌纲(Glomeromycetes)、粪壳菌纲(Sordariomycetes)、Pezizomycotina_Incertae_sedis纲、Pezizomycotina_Incertae_sedis目、粪壳菌目(Sordariales)、鸡油菌目(Cantharellales)、球囊霉目(Glomerales)、球囊霉科、丛赤壳科、Pezizomycotina_Incertae_sedis科和发菌科(Trichocomaceae),常绿落叶阔叶混交林中显著差异类群为革菌目(Thelephorales)、红菇目(Russulales)、革菌科和红菇科,锤舌菌纲(Leotiomycetes)、柔膜菌目(Helotiales)、蜡壳耳目(Sebacinales)、Sebacinales_Group_B科、柔膜菌科和锤舌菌科是亚高山矮林的显著差异类群。
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3种植被类型树种根系真菌在“极可能”和“很可能”水平上进行功能分组的ASVs数为2 231个,共被分为病理营养型(pathotroph)、共生营养型(symbiotroph)、腐生营养型(saprotroph)、病理-共生营养型(pathotroph-symbiotroph)和病理-腐生营养型(pathotroph-saprotroph)5个营养型。其中,41.86%的ASVs被归为腐生营养型,其次是共生营养型(28.73%)和病理营养型(15.37%),其余2个营养型所占比例较低。在功能亚类水平上,undefined saprotroph相对丰度最高,其次是丛枝菌根(arbuscular mycorrhizal)真菌、外生菌根(ectomycorrhizal)真菌、植物病原菌(plant pathogen)、内生真菌(endophyte)、寄生性真菌(fungal parasite)和杜鹃花类菌根(ericoid mycorrhizal)真菌,其余功能亚类相对丰度较低。此外,丛枝菌根真菌仅出现在常绿阔叶林和常绿落叶阔叶混交林,且在常绿阔叶林(7.77%)的相对丰度高于常绿落叶阔叶混交林(7.30%);外生菌根真菌在常绿落叶阔叶混交林(10.20%)的相对丰度明显高于常绿阔叶林(3.28%)和亚高山矮林(0.98%),内生真菌和杜鹃花类菌根真菌只出现在亚高山矮林。LEfSe分析(图1B)还发现,丛枝菌根真菌是常绿阔叶林的显著差异功能类群,外生菌根真菌是常绿落叶阔叶混交林的显著差异功能类群,亚高山矮林的显著差异功能类群为杜鹃花类菌根真菌。
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混合效应模型结果表明,植被类型能显著影响真菌丰富度和Shannon指数(P < 0.05),对Simpson指数和Pielou指数无显著影响(P > 0.05),常绿落叶林中真菌丰富度显著高于常绿落叶阔叶混交林和亚高山矮林,Shannon指数显著高于亚高山矮林(图2)。
图 2 不同植被类型根系真菌多样性指数
Figure 2. The diversity indices of root-associated fungi in different vegetation types
NMDS分析结果显示真菌群落组成在植被类型间存在差异(图3A),PERMANOVA揭示了相同的结果(F = 2.856,P = 0.001)。此外,Bray-Curtis相异性指数(图3B)在3种植被类型间存在极显著差异(P < 0.001),且在常绿落叶阔叶混交林中最高,常绿阔叶林次之,亚高山矮林最低,表明真菌群落组成在亚高山矮林样本间具有最高相似性,其次是常绿阔叶林,常绿落叶阔叶混交林的相似性最低,NMDS分析也印证了该结果。
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以植被类型为固定因子,树种为随机因子构建的混合效应模型发现,植被类型可显著影响土壤pH、土壤全碳、土壤全氮、土壤全磷、比根长、根组织密度、平均根直径、根比表面积、根碳含量、根氮含量、叶面积和叶碳含量(P<0.05)。此外,常绿阔叶林和常绿落叶阔叶混交林土壤pH显著高于亚高山矮林,亚高山矮林土壤具有最高的全碳和全氮含量,常绿阔叶林具有最高的土壤全磷含量;比根长、根比表面积和根碳含量在亚高山矮林显著高于常绿阔叶林和常绿落叶阔叶混交林,常绿阔叶林具有最高的根组织密度、平均根直径和根氮含量;叶面积在常绿阔叶林和常绿落叶阔叶混交林显著高于亚高山矮林,叶碳含量在亚高山矮林显著高于常绿阔叶林和常绿落叶阔叶混交林(表1)。
表 1 不同植被类型土壤理化性质、细根性状和叶性状
Table 1. Soil properties, fine root traits and leaf traits in different vegetation types
指标
Index常绿阔叶林
EBF常绿落叶阔叶混交林
EDBMF亚高山矮林
SDF土壤 pH
Soil pH6.067 ± 0.101 a 5.858 ± 0.156 a 5.006 ± 0.106 b 土壤全碳
Soil total carbon/(g·kg−1)7.057 ± 0.312 b 12.315 ± 1.677 ab 17.598 ± 2.946 a 土壤全氮
Soil total nitrogen/(g·kg−1)0.560 ± 0.022 b 0.875 ± 0.091 ab 1.127 ± 0.166 a 土壤全磷
Soil total phosphorus/(g·kg−1)6.067 ± 0.015a 5.858 ± 0.054 b 5.006 ± 0.053 ab 根生物量
Root biomass/g0.321 ± 0.063 a 0.239 ± 0.057 a 0.246 ± 0.019 a 根干物质含量
Root dry matter content/(g·g−1)0.255 ± 0.041 a 0.274 ± 0.030 a 0.277 ± 0.009 a 比根长
Specific root length/(cm·g−1)29.242 ± 3.209 b 44.158 ± 4.993 b 71.392 ± 10.885 a 根组织密度
Root tissue density/(g·cm−3)1.115 ± 0.249 a 1.014 ± 0.120 a 0.468 ± 0.052 b 平均根直径
Mean root diameter/cm0.237 ± 0.008 a 0.196 ± 0.014 b 0.220 ± 0.009 ab 根比表面积
Specific surface area/(cm2·g−1)217.188 ± 25.446 b 253.534 ± 28.078 b 479.641 ± 29.043 a 根碳含量
Root carbon content/(g·kg−1)46.115 ± 0.423 b 47.918 ± 0.756 b 50.371 ± 0.217 a 根氮含量
Root nitrogen content/(g·kg−1)2.357 ± 0.108 a 2.134 ± 0.128 ab 1.833 ± 0.050 b 根磷含量
Root phosphorus content/(g·kg−1)0.554 ± 0.015 a 0.673 ± 0.073 a 0.730 ± 0.049 a 叶生物量
Leaf biomass/g1.741 ± 0.222 a 1.609 ± 0.198 a 1.536 ± 0.172 a 叶面积
Leaf area/cm2278.285 ± 24.778 a 299.781 ± 26.502 a 167.156 ± 16.237 b 比叶面积
Specific leaf area/(cm2·g−1)169.114 ± 14.169 a 172.458 ± 19.323 a 138.494 ± 24.582 a 叶干物质含量
Leaf dry matter content/(g·g−1)0.386 ± 0.012 a 0.439 ± 0.017 a 0.370 ± 0.028 a 叶碳含量
Leaf carbon content/(g·kg−1)47.592 ± 0.715 b 47.937 ± 0.837 b 50.751 ± 0.456 a 叶氮含量
Leaf nitrogen content/(g·kg−1)1.902 ± 0.126 a 2.052 ± 0.110 a 1.722 ± 0.131 a 叶磷含量
Leaf phosphorus content/(g·kg−1)0.547 ± 0.039 a 0.701 ± 0.063 a 0.615 ± 0.077 a 注:数据为平均值 ± 标准误,同行后不同小写字母表示在P < 0.05水平上差异显著。
Notes: Data are means ± standard error. Different lowercase letters in the same row indicate significant difference at P < 0.05 level.ABT分析用来揭示植被类型、树种、土壤理化性质、细根性状和叶性状对真菌丰富度和多样性指数影响的相对重要性(图4)。结果表明:根碳含量是影响真菌丰富度的首要因子,相对重要性可达到24.47%(图4A);影响Shannon指数、Simpson指数和Pielou指数的首要因子均为叶干物质含量,相对重要性分别为14.63%、18.08%和15.55%(图4B、图4C、图4D)。Pearson相关性分析结果指出,根碳含量与真菌丰富度(r = −0.626,P < 0.001)、Shannon指数(r = −0.407,P = 0.004)呈显著负相关,叶干物质含量与Shannon指数(r = −0.293,P = 0.043)、Simpson指数(r = −0.591,P = 0.002)、Pielou指数(r = −0.531,P = 0.0259)呈显著负相关。
图 4 植被类型、树种、土壤性质、细根性状和叶性状对根系真菌丰富度(A)、Shannon指数(B)、Simpson指数(C)和Pielou指数(D)影响的相对重要性分析
Figure 4. Relative influences of vegetation type, tree species, soil properties, fine root traits and leaf traits on root-associated fungal richness (A), Shannon index (B), Simpson index (C) and Pielou index (D)
对包括植被类型和树种在内的22个解释变量进行前向选择后剩余7个,分别是植被类型、土壤pH、土壤全磷、根干物质含量、根碳含量、叶干物质含量和叶碳含量,与真菌群落构建RDA模型(图5A)。结果发现,植被类型、土壤pH、根碳含量和叶碳含量是影响真菌群落的显著因子(P < 0.05)。方差分解分析进一步揭示植被类型、土壤性质(土壤pH、土壤全磷)、细根性状(根干物质含量、根碳含量)和叶性状(叶干物质含量、叶碳含量)对真菌群落的影响(图5B),发现植被类型(26.96%)是影响真菌群落的首要因子,其次是细根性状(20.10%)和土壤性质(13.53%),叶性状(3.92%)影响最小。
梵净山典型植被类型的根系真菌群落结构及功能类群特征研究
Community Structure of Root-associated Fungi and Functional Characteristics in Typical Vegetation Types of Fanjingshan
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摘要:
目的 解析世界自然遗产地梵净山地区典型植被类型对真菌群落结构及功能的影响,为深入探讨可见的地上植物群落和未知的地下真菌群落结构间的关联机制提供数据支撑。 方法 以梵净山常绿阔叶林、常绿落叶阔叶混交林和亚高山矮林3种植被类型为研究对象,高通量测序技术探讨根系真菌群落结构在植被类型间的差异,FUNGuild对真菌进行功能预测,并分析真菌群落与土壤性质、细根性状和叶性状间的关系。 结果 子囊菌门和担子菌门为优势真菌门,undefined saprotroph、丛枝菌根真菌、外生菌根真菌、植物病原菌、内生真菌、寄生性真菌和杜鹃花类菌根真菌为主要功能类群。常绿阔叶林、常绿落叶阔叶混交林和亚高山矮林的显著差异功能类群分别是丛枝菌根真菌、外生菌根真菌和杜鹃花类菌根真菌。真菌群落组成在植被类型间存在显著差异,常绿阔叶林真菌多样性和Shannon指数显著高于常绿落叶阔叶混交林和亚高山矮林,而Bray-Curtis相异性指数在亚高山矮林显著低于常绿阔叶林和常绿落叶阔叶混交林。ABT分析表明,根碳含量是影响真菌丰富度的首要因子,叶干物质含量是影响真菌Shannon指数、Simpson指数和Pielou指数的首要因子。冗余分析揭示植被类型、土壤pH、根碳含量和叶碳含量显著影响真菌群落;方差分解分析表明植被类型对真菌群落的影响最大,其次是细根性状和土壤性质,叶性状影响最小。差分解分析表明植被类型对真菌群落的影响最大,其次是细根性状和土壤性质,叶性状影响最小。 结论 植被类型能显著影响真菌群落结构,真菌功能类群与植被类型间存在某种特定联系,植物性状对根系真菌群落的构建具有重要意义。 Abstract:Objective To reveal the effects of vegetation types on root-associated fungal community structure and function in the Fanjingshan World Heritage property, supportive data was provided to predict the relationships between visible vegetation types and invisible community structure in terrestrial ecosystem. Method Three representative vegetation types, including an evergreen broadleaved forest (EBF), an evergreen and deciduous broad-leaved mixed forest (EDBMF) and a subalpine dwarf shrub (SDF), were investigated in the Fanjingshan. Illumina MiSeq sequencing and the FUNGuild annotation tool were used to obtain and analyse the characteristics of the root-associated fungal community structure and function in different vegetation types, and the contributions of soil properties, fine root traits and leaf traits to variations in composition of root-associated fungal community were also analysed. Result Ascomycota and Basidiomycota were the two most abundant phyla, and undefined saprotroph, arbuscular mycorrhizal fungi, ectomycorrhizal fungi, plant pathogen, endophyte, fungal parasite and ericoid mycorrhizal fungi were the main guilds. Arbuscular mycorrhizal fungi were notably enriched in EBF, ectomycorrhizal fungi were more significantly abundant in EDBMF, and ericoid mycorrhizal fungi were significantly overrepresented in SDF. The significant difference in the composition of root-associated fungi among vegetation types was confirmed by non-metric multidimensional scaling analysis and permutational multivariate analysis of variance. A higher alpha diversity was detected in EBF, and the Bray-Curtis dissimilarity index was significantly lower in SDF than in EBF and EDBMF. Aggregated boosted tree analysis indicated that root carbon content was the primary factor influencing observed richness, while leaf dry matter content was the dominant factors associated with the changes in Shannon index, Simpson index and Pielou index of root-associated fungi. Redundancy analysis showed that changes in vegetation type, soil pH, root carbon content and leaf carbon content could significantly affect the fungal community composition. The variance partitioning analysis further revealed that vegetation type had the greatest impact on the composition of root-associated fungi, followed by fine root traits and soil properties, and leaf traits had the least impact. Conclusion Vegetation type can significantly affect root-associated fungal community structure. There is a specific relationship between vegetation type and functional taxa, and plant traits affect the construction of fungal communities. -
Key words:
- vegetation type
- / FUNGuild
- / root-associated fungi
- / functional taxa
- / fine root traits
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表 1 不同植被类型土壤理化性质、细根性状和叶性状
Table 1. Soil properties, fine root traits and leaf traits in different vegetation types
指标
Index常绿阔叶林
EBF常绿落叶阔叶混交林
EDBMF亚高山矮林
SDF土壤 pH
Soil pH6.067 ± 0.101 a 5.858 ± 0.156 a 5.006 ± 0.106 b 土壤全碳
Soil total carbon/(g·kg−1)7.057 ± 0.312 b 12.315 ± 1.677 ab 17.598 ± 2.946 a 土壤全氮
Soil total nitrogen/(g·kg−1)0.560 ± 0.022 b 0.875 ± 0.091 ab 1.127 ± 0.166 a 土壤全磷
Soil total phosphorus/(g·kg−1)6.067 ± 0.015a 5.858 ± 0.054 b 5.006 ± 0.053 ab 根生物量
Root biomass/g0.321 ± 0.063 a 0.239 ± 0.057 a 0.246 ± 0.019 a 根干物质含量
Root dry matter content/(g·g−1)0.255 ± 0.041 a 0.274 ± 0.030 a 0.277 ± 0.009 a 比根长
Specific root length/(cm·g−1)29.242 ± 3.209 b 44.158 ± 4.993 b 71.392 ± 10.885 a 根组织密度
Root tissue density/(g·cm−3)1.115 ± 0.249 a 1.014 ± 0.120 a 0.468 ± 0.052 b 平均根直径
Mean root diameter/cm0.237 ± 0.008 a 0.196 ± 0.014 b 0.220 ± 0.009 ab 根比表面积
Specific surface area/(cm2·g−1)217.188 ± 25.446 b 253.534 ± 28.078 b 479.641 ± 29.043 a 根碳含量
Root carbon content/(g·kg−1)46.115 ± 0.423 b 47.918 ± 0.756 b 50.371 ± 0.217 a 根氮含量
Root nitrogen content/(g·kg−1)2.357 ± 0.108 a 2.134 ± 0.128 ab 1.833 ± 0.050 b 根磷含量
Root phosphorus content/(g·kg−1)0.554 ± 0.015 a 0.673 ± 0.073 a 0.730 ± 0.049 a 叶生物量
Leaf biomass/g1.741 ± 0.222 a 1.609 ± 0.198 a 1.536 ± 0.172 a 叶面积
Leaf area/cm2278.285 ± 24.778 a 299.781 ± 26.502 a 167.156 ± 16.237 b 比叶面积
Specific leaf area/(cm2·g−1)169.114 ± 14.169 a 172.458 ± 19.323 a 138.494 ± 24.582 a 叶干物质含量
Leaf dry matter content/(g·g−1)0.386 ± 0.012 a 0.439 ± 0.017 a 0.370 ± 0.028 a 叶碳含量
Leaf carbon content/(g·kg−1)47.592 ± 0.715 b 47.937 ± 0.837 b 50.751 ± 0.456 a 叶氮含量
Leaf nitrogen content/(g·kg−1)1.902 ± 0.126 a 2.052 ± 0.110 a 1.722 ± 0.131 a 叶磷含量
Leaf phosphorus content/(g·kg−1)0.547 ± 0.039 a 0.701 ± 0.063 a 0.615 ± 0.077 a 注:数据为平均值 ± 标准误,同行后不同小写字母表示在P < 0.05水平上差异显著。
Notes: Data are means ± standard error. Different lowercase letters in the same row indicate significant difference at P < 0.05 level. -
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