氮添加对黄土丘陵区油松人工林根际土壤微生物群落结构的影响
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摘要
中国南方及中部为高氮沉降区(～35 kg·hm-2·a-1),氮沉降量向西北依次递减(～7.55 kg·hm-2·a-1).黄土高原区历经几十年的退耕还林还草而面貌一新,但该区域人工林土壤微生物群落结构对氮素添加响应的研究还鲜有报道.本研究以黄土丘陵区不同林龄油松人工林为研究对象,应用Illumina Hi Seq测序技术对细菌16S r DNA和真菌ITS进行序列测定与分析,探讨根际土壤细菌和真菌群落结构组成对土壤氮添加(施用量为200 kg N·hm-2·a-1纯氮)的响应,旨在探究我国西北黄土丘陵区油松人工林根际土壤微生物多样性及群落结构对定量氮添加的响应.结果表明:氮添加显著增加了25年龄林地细菌和真菌Shannon多样性,显著增加了40年龄林地细菌丰度指数.氮添加显著增加了40年龄林地拟杆菌门的相对丰度和25年龄林地酸杆菌门及接合菌门的相对丰度,但显著降低了40年龄林地奇古菌门的相对丰度.非度量多维尺度分析结果显示,氮添加对土壤细菌群落结构组成的影响程度大于真菌,对25年龄林地微生物群落结构组成的影响程度大于40年龄林地.表明不同林龄油松人工林根际土壤微生物的多样性及群落结构对土壤氮添加响应具有差异性.相对于真菌群落结构组成,细菌群落结构组成对氮添加更敏感;相对于40年龄林地,25年龄林地根际土壤微生物的群落结构组成对氮添加更敏感.因此,黄土高原区人工林地生态系统演替到一定阶段(40年左右),比幼龄林地生态系统更能承受外界较大的氮添加扰动.
Atmospheric nitrogen deposition in China displays a pronounced southeast-northwest gradient,with a range of ～ 35 kg·hm-2·a-1in southern and central China and ～ 7.55 kg·hm-2·a-1in the northwest. With the "Grain for Green"project,the environment of Loess Plateau has been greatly improved in the past decades. Little is known about how nitrogen deposition impacts the soil microbial communities in plantations in this region. In order to investigate the responses of diversity and structure of soil microbial community in Pinus tabuliformis plantations to nitrogen addition on Loess Plateau,China,we used high-throughput sequencing of 16 S ribosomal RNA genes of bacteria and ITS genes of fungi to investigate the effects of nitrogen addition( 200 kg N·hm-2·a-1) on the bacterial and fungal community over a 40-year chronosequence of coniferous P. tabuliformis plantation forests. After nitrogen addition,the Shannon diversity of both bacteria and fungi significantly increased in the 25-year-old stand,while the abundance-based coverage estimator of bacteria increased significantly in the 40-year-old stand. Nitrogen addition enhanced the relative abundance of Bacteroidetes in the 40-year-old stand and that of Acidobacteria/Zygomycota in the 25-year-old stand. In contrast,the relative abundance of Thaumarchaeota was reduced by nitrogen addition in the 40-year old stand. The effects of nitrogen addition were stronger on bacterial than on fungal community structure as indicated by the results from non-metric multidimensional scaling analysis.Moreover,the effects of nitrogen addition were stronger in the 25-year-old stand than in the 40-yearold stand. These results indicated that the effects of nitrogen addition on microbial community structure diminished over time in temperate forests on Loess Plateau,China. The stronger effects of nitrogen addition was found on the community structure of bacteria than that of fungi,and on the microbial community structure in the 25-year-old stand than in the 40-year-old stand. Our findings suggested that forest ecosystem at a certain stage( ～ 40-year old) would be less sensitive to disturbance such as nitrogen addition than young forests.
Atmospheric nitrogen deposition in China displays a pronounced southeast-northwest gradient,with a range of ～ 35 kg·hm-2·a-1in southern and central China and ～ 7.55 kg·hm-2·a-1in the northwest. With the "Grain for Green"project,the environment of Loess Plateau has been greatly improved in the past decades. Little is known about how nitrogen deposition impacts the soil microbial communities in plantations in this region. In order to investigate the responses of diversity and structure of soil microbial community in Pinus tabuliformis plantations to nitrogen addition on Loess Plateau,China,we used high-throughput sequencing of 16 S ribosomal RNA genes of bacteria and ITS genes of fungi to investigate the effects of nitrogen addition( 200 kg N·hm-2·a-1) on the bacterial and fungal community over a 40-year chronosequence of coniferous P. tabuliformis plantation forests. After nitrogen addition,the Shannon diversity of both bacteria and fungi significantly increased in the 25-year-old stand,while the abundance-based coverage estimator of bacteria increased significantly in the 40-year-old stand. Nitrogen addition enhanced the relative abundance of Bacteroidetes in the 40-year-old stand and that of Acidobacteria/Zygomycota in the 25-year-old stand. In contrast,the relative abundance of Thaumarchaeota was reduced by nitrogen addition in the 40-year old stand. The effects of nitrogen addition were stronger on bacterial than on fungal community structure as indicated by the results from non-metric multidimensional scaling analysis.Moreover,the effects of nitrogen addition were stronger in the 25-year-old stand than in the 40-yearold stand. These results indicated that the effects of nitrogen addition on microbial community structure diminished over time in temperate forests on Loess Plateau,China. The stronger effects of nitrogen addition was found on the community structure of bacteria than that of fungi,and on the microbial community structure in the 25-year-old stand than in the 40-year-old stand. Our findings suggested that forest ecosystem at a certain stage( ～ 40-year old) would be less sensitive to disturbance such as nitrogen addition than young forests.
引文
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[8]Deng W-Y(邓文悦).Effects of Long-term Fertilization on Soil Organic Matter and Soil Microbial Functional Diversity in Jiangxi Paddy Soil.Master Thesis.Xi’an:Northwest University,2017(in Chinese)
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[11]Zi H-B(字洪标),Dai D(代迪),Hu L(胡雷),et al.Responses of soil microbial functional diversity to phosphorus addition in an alpine meadow on Northwestern Plateau of Sichuan Province.Chinese Journal of Soil Science(土壤通报),2017,48(3):647-655(in Chinese)
[12]Zeng Q-P(曾清苹),He B-H(何丙辉).Effect of nitrogen deposition on soil microbial community structure determined with the PLFA method under the Masson pine forest from Mt.Jinyun,Chongqing.Environmental Science(环境科学),2016,37(9):3590-3597(in Chinese)
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[14]Hong P-Z(洪丕征),Liu S-R(刘世荣),Yu H-L(于浩龙),et al.Effects of simulated nitrogen deposition on soil microbial biomass and community structure in a young plantation of Castanopsis hystrix.Journal of Shandong University(Natural Science)(山东大学学报:理学版),2016,51(5):18-28(in Chinese)
[15]Xu K(许可),Wang C-M(王春梅),Zhang Y(张艺),et al.Effect of simulated atmospheric nitrogen deposition on soil microbial community structure in a temperate forest.Chinese Journal of Ecology(生态学杂志),2016,35(10):2676-2683(in Chinese)
[16]Wang C,Feng Z,Xiang Z,et al.The effects of N and Padditions on microbial N transformations and biomass on saline-alkaline grassland of Loess Plateau of Northern China.Geoderma,2014,213:419-425
[17]Zhou HJ,Rompaey AV,Wang JA.Detecting the impact of the“Grain for Green”program on the mean annual vegetation cover in the Shaanxi Province,China using SPOT-VGT NDVI data.Land Use Policy,2009,26:954-960
[18]Wang Z-H(王志辉),Zhang Y(张颖),Liu X-J(刘学军),et al.Dry and wet nitrogen deposition in agricultural soils in the Loess area.Acta Ecologica Sinica(生态学报),2008,28(7):3295-3301(in Chinese)
[19]Chen LL,Mu XM,Yuan ZY,et al.Soil nutrients and water affect the age-related fine root biomass but not production in two plantation forests on the Loess Plateau,China.Journal of Arid Environments,2016,135:173-180
[20]Tu L-H(涂利华),Hu T-X(胡庭兴),Zhang J(张健),et al.Effects of simulated nitrogen deposition on the fine root characteristics and soil respiration in a Pleioblastus amarus plantation in Rainy Area of West China.Chinese Journal of Applied Ecology(应用生态学报),2010,21(10):2472-2478(in Chinese)
[21]Chen L-L(陈莉莉).The Belowground Productivity and Ecological Stoichiometry in Pinus tabuliformis and Robinia pseudoacacia Plantations on the Loess Plateau.PhD Thesis.Yangling,Shaanxi:Research Center of Soil and Water Conservation and Ecological Environment,Chinese Academy of Sciences and Ministry of Education,2017(in Chinese)
[22]Kidd PS,Prieto-Fernandez A,Monterroso C,et al.Rhizosphere microbial community and hexachlorocyclohexane degradative potential in contrasting plant species.Plant and Soil,2008,302:233-247
[23]O’Donnell AG,Davies JT.Plants and fertilisers as drivers of change in microbial community structure and function in soils.Plant and Soil,2001,232:135-145
[24]Sui X(隋心),Zhang R-T(张荣涛),Yang L-B(杨立宾),et al.Effect of simulation nitrogen depositions on bacterial diversity of Deyeuxia angustifolia in wetland of Sanjing Plain.Pratacultural Science(草业科学),2016,33(4):589-598(in Chinese)
[25]Jiang S-L(姜思领).Response to Nitrogen Deposition of Winter Soil Microorganism on in Lesser Khingan Mountains Forest.Master Thesis Harbin:Heilongjiang University(黑龙江大学),2016(in Chinese)
[26]Hao Y-Q(郝亚群),Xie L(谢麟),Chen Y-M(陈岳民),et al.Effects of nitrogen deposition on diversity and composition of soil bacterial community in a subtropical Cunninghamia lanceolata plantation.Chinese Journal of Applied Ecology(应用生态学报),2018,29(1):53-58(in Chinese)
[27]Chen Z-H(陈志豪),Liang X(梁雪),Li Y-C(李永春),et al.Effects of different fertilization regimes on soil fungal communities under Phyllostachys violascens stand.Chinese Journal of Applied Ecology(应用生态学报),2017,28(4):1168-1176(in Chinese)
[28]Wang C,Liu D,Bai E.Decreasing soil microbial diversity is associated with decreasing microbial biomass under nitrogen addition.Soil Biology&Biochemistry,2018,120:126-133
[29]Li CH,Yan K,Tang LS,et al.Change in deep soil microbial communities due to long-term fertilization.Soil Biology&Biochemistry,2014,75:264-272
[30]Nemergut DR,Townsend AR,Sattin SR,et al.The effects of chronic nitrogen fertilization on alpine tundra soil microbial communities:Implications for carbon and nitrogen cycling.Environmental Microbiology,2008,10:3093-3105
[31]Yu H-L(于海玲).Effect of Nitrogen Application on Composition Characters of Soil Microbial Communities.PhD Thesis.Changchun:Jilin Agricultural University,2017(in Chinese)
[32]Yating HE,Yuchun QI,Dong Y,et al.Advances in the influence of external nitrogen input on soil microbiological characteristics of grassland ecosystem.Advances in Earth Science,2010,25:877-885
[33]Xin L(辛亮).Effects of Long-term Fertilization on Soil Nitrification and N2O Emission in the Semiarid Loess Plateau.Master Thesis.Yangling,Shaanxi:Northwest A&F University,2012(in Chinese)
[34]Zhang H,Sekiguchi Y,Hanada S,et al.Gemmatimonas aurantiaca gen.nov.,sp.nov.,a gram-negative,aerobic,polyphosphate-accumulating micro-organism,the first cultured representative of the new bacterial phylum Gemmatimonadetes phyl.nov.International Journal of Systematic&Evolutionary Microbiology,2003,53:1155-1163
[35]Deforest JL,Zak DR,Pregitzer KS,et al.Atmospheric nitrate deposition and the microbial degradation of cellobiose and vanillin in a northern hardwood forest.Soil Biology&Biochemistry,2004,36:965-971
[2]King GM.Urban microbiomes and urban ecology:How do microbes in the built environment affect human sustainability in cities?Journal of Microbiology,2014,52:721-728
[3]Li Y-M(李延茂),Hu J-C(胡江春),Wang S-L(汪思龙),et al.Function and application of soil microorganisms in forest ecosystem.Chinese Journal of Applied Ecology(应用生态学报),2004,15(10):1943-1946(in Chinese)
[4]Fang Y,Yoh M,Koba K,et al.Nitrogen deposition and forest nitrogen cycling along an urban-rural transect in southern China.Global Change Biology,2011,17:872-885
[5]Zhang C-J(张翠景),He J-Z(贺纪正),Shen J-P(沈菊培).Global change field manipulative experiments and their applications in soil microbial ecology.Chinese Journal of Applied Ecology(应用生态学报),2016,27(5):1663-1673(in Chinese)
[6]Ramirez KS,Fierer N.Consistent effects of nitrogen fertilization on soil bacterial communities in contrasting systems.Ecology,2010,91:3463-3470
[7]Treseder KK.Nitrogen additions and microbial biomass:A meta-analysis of ecosystem studies.Ecology Letters,2008,11:1111-1120
[8]Deng W-Y(邓文悦).Effects of Long-term Fertilization on Soil Organic Matter and Soil Microbial Functional Diversity in Jiangxi Paddy Soil.Master Thesis.Xi’an:Northwest University,2017(in Chinese)
[9]Zhang L-C(张立超),He C-F(何昌福),Zhang W-M(张文明),et al.Effects of fertilizer treatments on soil organic matter content and microbial flora of potato.Journal of Gansu Agricultural University(甘肃农业大学学报),2017,52(2):27-33(in Chinese)
[10]Zhang F-G(张风革),Huo Y-Q(霍云倩),Sun Y(孙艺),et al.Effect of consecutive biofertilizer application on aboveground biomass and management of soil microflora in grassland.Journal of Nanjing Agricultural University(南京农业大学学报),2018,41(2):382-388(in Chinese)
[11]Zi H-B(字洪标),Dai D(代迪),Hu L(胡雷),et al.Responses of soil microbial functional diversity to phosphorus addition in an alpine meadow on Northwestern Plateau of Sichuan Province.Chinese Journal of Soil Science(土壤通报),2017,48(3):647-655(in Chinese)
[12]Zeng Q-P(曾清苹),He B-H(何丙辉).Effect of nitrogen deposition on soil microbial community structure determined with the PLFA method under the Masson pine forest from Mt.Jinyun,Chongqing.Environmental Science(环境科学),2016,37(9):3590-3597(in Chinese)
[13]Yuan Y-H(袁颖红),Fan H-B(樊后保),Li H-X(李辉信),et al.Effects of simulated nitrogen deposition on soil microorganism in a Chinese fir plantation.Scientia Silvae Sinicae(林业科学),2012,48(9):8-14(in Chinese)
[14]Hong P-Z(洪丕征),Liu S-R(刘世荣),Yu H-L(于浩龙),et al.Effects of simulated nitrogen deposition on soil microbial biomass and community structure in a young plantation of Castanopsis hystrix.Journal of Shandong University(Natural Science)(山东大学学报:理学版),2016,51(5):18-28(in Chinese)
[15]Xu K(许可),Wang C-M(王春梅),Zhang Y(张艺),et al.Effect of simulated atmospheric nitrogen deposition on soil microbial community structure in a temperate forest.Chinese Journal of Ecology(生态学杂志),2016,35(10):2676-2683(in Chinese)
[16]Wang C,Feng Z,Xiang Z,et al.The effects of N and Padditions on microbial N transformations and biomass on saline-alkaline grassland of Loess Plateau of Northern China.Geoderma,2014,213:419-425
[17]Zhou HJ,Rompaey AV,Wang JA.Detecting the impact of the“Grain for Green”program on the mean annual vegetation cover in the Shaanxi Province,China using SPOT-VGT NDVI data.Land Use Policy,2009,26:954-960
[18]Wang Z-H(王志辉),Zhang Y(张颖),Liu X-J(刘学军),et al.Dry and wet nitrogen deposition in agricultural soils in the Loess area.Acta Ecologica Sinica(生态学报),2008,28(7):3295-3301(in Chinese)
[19]Chen LL,Mu XM,Yuan ZY,et al.Soil nutrients and water affect the age-related fine root biomass but not production in two plantation forests on the Loess Plateau,China.Journal of Arid Environments,2016,135:173-180
[20]Tu L-H(涂利华),Hu T-X(胡庭兴),Zhang J(张健),et al.Effects of simulated nitrogen deposition on the fine root characteristics and soil respiration in a Pleioblastus amarus plantation in Rainy Area of West China.Chinese Journal of Applied Ecology(应用生态学报),2010,21(10):2472-2478(in Chinese)
[21]Chen L-L(陈莉莉).The Belowground Productivity and Ecological Stoichiometry in Pinus tabuliformis and Robinia pseudoacacia Plantations on the Loess Plateau.PhD Thesis.Yangling,Shaanxi:Research Center of Soil and Water Conservation and Ecological Environment,Chinese Academy of Sciences and Ministry of Education,2017(in Chinese)
[22]Kidd PS,Prieto-Fernandez A,Monterroso C,et al.Rhizosphere microbial community and hexachlorocyclohexane degradative potential in contrasting plant species.Plant and Soil,2008,302:233-247
[23]O’Donnell AG,Davies JT.Plants and fertilisers as drivers of change in microbial community structure and function in soils.Plant and Soil,2001,232:135-145
[24]Sui X(隋心),Zhang R-T(张荣涛),Yang L-B(杨立宾),et al.Effect of simulation nitrogen depositions on bacterial diversity of Deyeuxia angustifolia in wetland of Sanjing Plain.Pratacultural Science(草业科学),2016,33(4):589-598(in Chinese)
[25]Jiang S-L(姜思领).Response to Nitrogen Deposition of Winter Soil Microorganism on in Lesser Khingan Mountains Forest.Master Thesis Harbin:Heilongjiang University(黑龙江大学),2016(in Chinese)
[26]Hao Y-Q(郝亚群),Xie L(谢麟),Chen Y-M(陈岳民),et al.Effects of nitrogen deposition on diversity and composition of soil bacterial community in a subtropical Cunninghamia lanceolata plantation.Chinese Journal of Applied Ecology(应用生态学报),2018,29(1):53-58(in Chinese)
[27]Chen Z-H(陈志豪),Liang X(梁雪),Li Y-C(李永春),et al.Effects of different fertilization regimes on soil fungal communities under Phyllostachys violascens stand.Chinese Journal of Applied Ecology(应用生态学报),2017,28(4):1168-1176(in Chinese)
[28]Wang C,Liu D,Bai E.Decreasing soil microbial diversity is associated with decreasing microbial biomass under nitrogen addition.Soil Biology&Biochemistry,2018,120:126-133
[29]Li CH,Yan K,Tang LS,et al.Change in deep soil microbial communities due to long-term fertilization.Soil Biology&Biochemistry,2014,75:264-272
[30]Nemergut DR,Townsend AR,Sattin SR,et al.The effects of chronic nitrogen fertilization on alpine tundra soil microbial communities:Implications for carbon and nitrogen cycling.Environmental Microbiology,2008,10:3093-3105
[31]Yu H-L(于海玲).Effect of Nitrogen Application on Composition Characters of Soil Microbial Communities.PhD Thesis.Changchun:Jilin Agricultural University,2017(in Chinese)
[32]Yating HE,Yuchun QI,Dong Y,et al.Advances in the influence of external nitrogen input on soil microbiological characteristics of grassland ecosystem.Advances in Earth Science,2010,25:877-885
[33]Xin L(辛亮).Effects of Long-term Fertilization on Soil Nitrification and N2O Emission in the Semiarid Loess Plateau.Master Thesis.Yangling,Shaanxi:Northwest A&F University,2012(in Chinese)
[34]Zhang H,Sekiguchi Y,Hanada S,et al.Gemmatimonas aurantiaca gen.nov.,sp.nov.,a gram-negative,aerobic,polyphosphate-accumulating micro-organism,the first cultured representative of the new bacterial phylum Gemmatimonadetes phyl.nov.International Journal of Systematic&Evolutionary Microbiology,2003,53:1155-1163
[35]Deforest JL,Zak DR,Pregitzer KS,et al.Atmospheric nitrate deposition and the microbial degradation of cellobiose and vanillin in a northern hardwood forest.Soil Biology&Biochemistry,2004,36:965-971
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