浑善达克沙地生物土壤结皮及其下层土壤中固氮细菌群落结构和多样性
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摘要
【背景】荒漠化是一个重大环境问题,生物土壤结皮(Biological soil crusts,BSCs)可遏制荒漠化,其中的固氮微生物对BSCs的形成和发育有重要作用,但目前BSCs中固氮细菌群落结构和多样性尚不十分清楚。【目的】阐明浑善达克沙地中不同类型生物土壤结皮及其下层土壤固氮细菌的群落结构、多样性及其影响因素。【方法】利用稀释热法和碱解扩散法检测土壤的有机质(Organic matter,OM)和速效氮(Available nitrogen,AN)含量;利用高通量测序对nifH基因进行测序,基于nifH序列比较分析固氮细菌群落结构和多样性;利用典范对应分析(Canonical correlation analysis,CCA)分析群落、样品和土壤理化参数的相关性。【结果】固氮细菌优势菌门除在苔藓结皮(HSM)中为Cyanobacteria和Proteobacteria外,在其他类型BSCs中均只为Cyanobacteria;苔藓结皮下层土壤(HSMs)(下层土壤中只有HSMs检测到了nifH)优势菌门为Proteobacteria,优势菌纲为Alphaproteobacteria和Betaproteobacteria;优势菌属差异较大,藻结皮(HSA)中Unclassified_f_Nostocaceae占90.99%;地衣结皮(HSL)中Scytonema和Unclassified_f_Nostocaceae分别占45.85%和44.14%;HSM中Unclassified_f_Nostocaceae、Scytonema、Nostoc、Skermanella、Unclassified_o_Nostocales分别占29.21%、22.57%、15.34%、14.74%和10.60%;HSMs中Skermanella、Azohydromonas、Unclassified_p_Proteobacteria、Unclassified_c_Alphaproteobacteria分别占33.80%、25.66%、18.20%和10.62%;固氮细菌多样性随结皮的发育逐渐提高;OM和AN对结皮的发育起促进作用。【结论】藻结皮、地衣结皮和苔藓结皮及其紧邻下层土壤中的固氮细菌群落结构和多样性差异明显,且固氮细菌类群和多样性指数随BSCs发育阶段的提高而增加。本研究为认识和利用生物土壤结皮相关固氮细菌提供了基础依据。
[Background] Desertification is a major environmental problem, biological soil crusts(BSCs) can inhibit desertification, and the diazotrophs play an important role in the formation and development of BSCs. However, the community structure and diversity of diazotrophs in BSCs are not clear enough. [Objective] To clarify the community structure and diversity of diazotrophs in different types of biological soil crusts(BSCs) and soils under them and the effectors of soil environmental factors. [Methods] Using dilution heat method and alkali solution diffusion to measure the content of organic matter(OM) and the available nitrogen(AN) of soil, respectively;nifH gene was sequenced via the high-throughput sequencing platform, and the community structure, diversity and the variance analysis of diazotrophs were analyzed by bioinformatics analysis; Canonical Correlation Analysis(CCA) were used to analyze the correlation among the community structure, samples and the physico-chemical parameters of soil. [Results] Dominant phylum were Proteobacteria and Cyanobacteria in moss crusts(HSM), Cyanobacteria in the other types of BSCs; the class of Alphaproteobacteria and Betaproteobacteria accounted for the most fraction in the soil underneath moss crusts(nifH was only detected in soil underneath moss crusts, HSMs);significant differences existed in community structure at genus level, algae crusts(HSA) was predominated by unclassified_f_Nostocaceae(90.99%), Lichen crusts(HSL) by Scytonema(45.85%)and unclassified_f_Nostocaceae(44.14%), HSM by unclassified_f_Nostocaceae(29.21%), Scytonema(22.57%), Nostoc(15.34%), Skermanella(14.74%), unclassified_o_Nostocales(10.60%), and HSMs by Skermanella(33.80%), Azohydromonas(25.66%), unclassified_p_Proteobacteria(18.20%) and unclassified_c_Alphaproteobacteria(10.62%). [Conclusion] The community structure and diversity of diazotrophs in Algae, Lichen and Moss crusts and soil underneath crusts were markedly different, and the community species and the diversity increased with the development of the BSCs. This study provides a basis for appreciation and utilization of the diazotrophs in the BSCs.
[Background] Desertification is a major environmental problem, biological soil crusts(BSCs) can inhibit desertification, and the diazotrophs play an important role in the formation and development of BSCs. However, the community structure and diversity of diazotrophs in BSCs are not clear enough. [Objective] To clarify the community structure and diversity of diazotrophs in different types of biological soil crusts(BSCs) and soils under them and the effectors of soil environmental factors. [Methods] Using dilution heat method and alkali solution diffusion to measure the content of organic matter(OM) and the available nitrogen(AN) of soil, respectively;nifH gene was sequenced via the high-throughput sequencing platform, and the community structure, diversity and the variance analysis of diazotrophs were analyzed by bioinformatics analysis; Canonical Correlation Analysis(CCA) were used to analyze the correlation among the community structure, samples and the physico-chemical parameters of soil. [Results] Dominant phylum were Proteobacteria and Cyanobacteria in moss crusts(HSM), Cyanobacteria in the other types of BSCs; the class of Alphaproteobacteria and Betaproteobacteria accounted for the most fraction in the soil underneath moss crusts(nifH was only detected in soil underneath moss crusts, HSMs);significant differences existed in community structure at genus level, algae crusts(HSA) was predominated by unclassified_f_Nostocaceae(90.99%), Lichen crusts(HSL) by Scytonema(45.85%)and unclassified_f_Nostocaceae(44.14%), HSM by unclassified_f_Nostocaceae(29.21%), Scytonema(22.57%), Nostoc(15.34%), Skermanella(14.74%), unclassified_o_Nostocales(10.60%), and HSMs by Skermanella(33.80%), Azohydromonas(25.66%), unclassified_p_Proteobacteria(18.20%) and unclassified_c_Alphaproteobacteria(10.62%). [Conclusion] The community structure and diversity of diazotrophs in Algae, Lichen and Moss crusts and soil underneath crusts were markedly different, and the community species and the diversity increased with the development of the BSCs. This study provides a basis for appreciation and utilization of the diazotrophs in the BSCs.
引文
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[16]Rajeev L,da Rocha UN,Klitgord N,et al.Dynamic cyanobacterial response to hydration and dehydration in a desert biological soil crust[J].The ISME Journal,2013,7(11):2178-2191
[17]Tahon G,Tytgat B,Stragier P,et al.Analysis of cbb L,nif H,and puf LM in soils from the S?r Rondane Mountains,Antarctica,reveals a large diversity of autotrophic and phototrophic bacteria[J].Microbial Ecology,2016,71(1):131-149
[18]K?berl M,Erlacher A,Ramadan EM,et al.Comparisons of diazotrophic communities in native and agricultural desert ecosystems reveal plants as important drivers in diversity[J].FEMS Microbiology Ecology,2016,92(2):fiv166
[19]Zhang BC,Kong WD,Wu N,et al.Bacterial diversity and community along the succession of biological soil crusts in the Gurbantunggut Desert,Northern China[J].Journal of Basic Microbiology,2016,56(6):670-679
[20]Yachi S,Loreau M.Biodiversity and ecosystem productivity in a fluctuating environment:the insurance hypothesis[J].Proceedings of the National Academy of Sciences of the United States of America,1999,96(4):1463-1468
[2]Zhang W,Yang L,Wang ZJ.Advance and development trend of biological nitrogen fixation research[J].Journal of Yunnan Agricultural University(Natural Science),2015,30(5):810-821(in Chinese)张武,杨琳,王紫娟.生物固氮的研究进展及发展趋势[J].云南农业大学学报:自然科学,2015,30(5):810-821
[3]Belnap J,Weber B,Büdel B.Biological Soil Crusts as an Organizing Principle in Drylands[M].Switzerland:Springer International Publishing,2016:3-13
[4]Li XR,Zhou HY,Wang XP,et al.Ecological restoration and recovery in arid desert regions of China:a review for 60-year research progresses of Shapotou desert research and experiment station,Chinese academy of sciences[J].Journal of Desert Research,2016,36(2):247-264(in Chinese)李新荣,周海燕,王新平,等.中国干旱沙区的生态重建与恢复:沙坡头站60年重要研究进展综述[J].中国沙漠,2016,36(2):247-264
[5]Zhang P,Li XR,Zhang ZS,et al.Nitrogen fixation potential of biological soil crusts in southeast edge of Tengger Desert,Northwest China[J].Chinese Journal of Applied Ecology,2012,23(8):2157-2164(in Chinese)张鹏,李新荣,张志山,等.腾格里沙漠东南缘生物土壤结皮的固氮潜力[J].应用生态学报,2012,23(8):2157-2164
[6]Pepe-Ranney C,Koechli C,Potrafka R,et al.Non-cyanobacterial diazotrophs mediate dinitrogen fixation in biological soil crusts during early crust formation[J].The ISME Journal,2016,10(2):287-298
[7]JanatkováK,?ehákováK,Dole?al J,et al.Community structure of soil phototrophs along environmental gradients in arid Himalaya[J].Environmental Microbiology,2013,15(9):2505-2516
[8]Zhang JE.A Common Experimental Study of Methods andTechniques on Ecology[M].Beijing:Chemical Industry Publishing House,2007:225(in Chinese)章家恩.生态学常用实验研究方法与技术[M].北京:化学工业出版社,2007:225
[9]Bao SD.Soil Agrochemical Analysis[M].3rd ed.Beijing:Agricultural Publishing House,2000:34(in Chinese)鲍士旦.土壤农化分析[M].3版.北京:农业出版社,2000:34
[10]Zhou X,Smith H,Silva AG,et al.Differential responses of dinitrogen fixation,diazotrophic cyanobacteria and ammonia oxidetion reveal a potential warming-induced imbalance of the N-Cycle in biological soil crusts[J].PLo S One,2016,11(10):e0164932
[11]Johnson SL,Neuer S,Garcia-Pichel F.Export of nitrogenous compounds due to incomplete cycling within biological soil crusts of arid lands[J].Environmental Microbiology,2007,9(3):680-689
[12]Yeager CM,Kornosky JL,Housman DC,et al.Diazotrophic community structure and function in two successional stages of biological soil crusts from the Colorado Plateau and Chihuahuan Desert[J].Applied and Environmental Microbiology,2004,70(2):973-983
[13]Yeager CM,Kuske CR,Carney TD,et al.Response of biological soil crust diazotrophs to season,altered summer precipitation,and year-round increased temperature in an arid grassland of the Colorado Plateau,USA[J].Frontiers in Microbiology,2012,3:358
[14]Wang J,Bao JT,Li XR,et al.Molecular ecology of nif H genes and transcripts along a chronosequence in revegetated areas of the Tengger desert[J].Microbial Ecology,2016,71(1):150-163
[15]Beraldi-Campesi H,Hartnett HE,Anbar A,et al.Effect of biological soil crusts on soil elemental concentrations:implications for biogeochemistry and as traceable biosignatures of ancient life on land[J].Geobiology,2009,7(3):348-359
[16]Rajeev L,da Rocha UN,Klitgord N,et al.Dynamic cyanobacterial response to hydration and dehydration in a desert biological soil crust[J].The ISME Journal,2013,7(11):2178-2191
[17]Tahon G,Tytgat B,Stragier P,et al.Analysis of cbb L,nif H,and puf LM in soils from the S?r Rondane Mountains,Antarctica,reveals a large diversity of autotrophic and phototrophic bacteria[J].Microbial Ecology,2016,71(1):131-149
[18]K?berl M,Erlacher A,Ramadan EM,et al.Comparisons of diazotrophic communities in native and agricultural desert ecosystems reveal plants as important drivers in diversity[J].FEMS Microbiology Ecology,2016,92(2):fiv166
[19]Zhang BC,Kong WD,Wu N,et al.Bacterial diversity and community along the succession of biological soil crusts in the Gurbantunggut Desert,Northern China[J].Journal of Basic Microbiology,2016,56(6):670-679
[20]Yachi S,Loreau M.Biodiversity and ecosystem productivity in a fluctuating environment:the insurance hypothesis[J].Proceedings of the National Academy of Sciences of the United States of America,1999,96(4):1463-1468