增温对南亚热带季风常绿阔叶林土壤微生物群落的影响
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摘要
土壤微生物是森林生态系统中重要的分解者,参与生物圈的物质循环和能量流动,对温度变化响应较为敏感。以鼎湖山南亚热带季风常绿阔叶林为研究对象,基于野外增温实验平台,采集0-10 cm和10-20 cm土层的土壤样品,采用磷脂脂肪酸(PLFA)方法并结合土壤理化性质的监测,探究气温上升对土壤微生物群落的影响。结果表明:(1)增温处理使0-10 cm和10-20 cm土层月均温分别显著上升1.24℃和1.17℃,土层湿度变化不显著;(2)增温显著增加了土壤硝氮含量,但对其他理化性质作用不明显;(3)增温组土壤微生物生物量碳(MBC)、微生物生物量氮(MBN)、微生物生物量碳氮比(C/N)以及微生物总磷脂脂肪酸含量与对照组差异不显著;(4)增温显著改变了土壤微生物群落结构,使细菌相对丰度、细菌真菌之比(B/F)以及革兰氏阳性菌革兰氏阴性菌之比(G~+/G~-)显著增加,降低了真菌和丛枝菌根真菌的相对丰度;(5)进一步分析表明,土壤硝态氮和有机碳是影响土壤微生物群落结构变异的主要因子,两者共同解释了微生物群落结构60.5%的变异度。以上研究结果表明,尽管增温对南亚热带季风常绿阔叶林土壤微生物生物量作用不明显,但可通过对土壤硝氮和土壤有机碳含量的影响引起土壤微生物群落结构及其相对丰度的改变,微生物群落结构和相对丰度的变化又将通过影响微生物对土壤碳氮的同化作用,最终影响土壤的碳氮过程。
Soil microorganism, as an important decomposer, plays critical roles in the nutrient cycling and energy flow in forest ecosystem, which is often dependence of temperature. We established a field warming experiment to explore the effects of warming on soil microbial communities of the monsoon evergreen broad-leaved forest in Dinghushan National Nature Reserve in south subtropical region of China. Soil samples were collected in layers of 0-10 cm and 10-20 cm to monitor structure of soil microbial community by using the phospholipid fatty acids(PLFA) as biomarkers. The results showed that:(1) The warming treatment did not significantly alter soil moisture in both soil layers, while averagely increased soil temperature by 1.24 ℃ in 0-10 cm and 1.17 ℃ in 10-20 cm soil layers, respectively.(2) The warming treatment significantly increased soil nitrate content, but no significant effect was found on other soil physical and chemical properties.(3) The warming treatment has no significant effect on soil microbial biomass carbon(MBC), microbial biomass nitrogen(MBN), the ratio of MBC to MBN or microbial total PLFAs content.(4) The warming treatment significantly changed soil microbial community structure, with increases in the relative abundance of bacteria, the ratio of bacteria to fungi and the ratio of Gram-positive bacteria to Gram-negative bacteria but decreases in the relative abundance of fungi and arbuscular mycorrhizal fungi. And(5) further analysis showed that soil nitrate nitrogen and soil organic carbon was the most important influencing factors, explaining 60.5% variability of microbial community structure under the warming treatment.These findings indicate that climate warming may alter soil microbial community structure instead of soil microbial biomass by influencing soil nitrate nitrogen and soil organic carbon in the southern subtropical monsoon evergreen broad-leaved forest, in turn resulting in an effect on soil carbon and nitrogen processes by microbial assimilation.
Soil microorganism, as an important decomposer, plays critical roles in the nutrient cycling and energy flow in forest ecosystem, which is often dependence of temperature. We established a field warming experiment to explore the effects of warming on soil microbial communities of the monsoon evergreen broad-leaved forest in Dinghushan National Nature Reserve in south subtropical region of China. Soil samples were collected in layers of 0-10 cm and 10-20 cm to monitor structure of soil microbial community by using the phospholipid fatty acids(PLFA) as biomarkers. The results showed that:(1) The warming treatment did not significantly alter soil moisture in both soil layers, while averagely increased soil temperature by 1.24 ℃ in 0-10 cm and 1.17 ℃ in 10-20 cm soil layers, respectively.(2) The warming treatment significantly increased soil nitrate content, but no significant effect was found on other soil physical and chemical properties.(3) The warming treatment has no significant effect on soil microbial biomass carbon(MBC), microbial biomass nitrogen(MBN), the ratio of MBC to MBN or microbial total PLFAs content.(4) The warming treatment significantly changed soil microbial community structure, with increases in the relative abundance of bacteria, the ratio of bacteria to fungi and the ratio of Gram-positive bacteria to Gram-negative bacteria but decreases in the relative abundance of fungi and arbuscular mycorrhizal fungi. And(5) further analysis showed that soil nitrate nitrogen and soil organic carbon was the most important influencing factors, explaining 60.5% variability of microbial community structure under the warming treatment.These findings indicate that climate warming may alter soil microbial community structure instead of soil microbial biomass by influencing soil nitrate nitrogen and soil organic carbon in the southern subtropical monsoon evergreen broad-leaved forest, in turn resulting in an effect on soil carbon and nitrogen processes by microbial assimilation.
引文
BILLINGS S A,2012.How interactions between microbial resource demands,soil organic matter stoichiometry,and substrate reactivity determine the direction and magnitude of soil respiratory responses to warming[J].Global Change Biology,19(1):90-102.
BLIGH E,1959.A rapid method of total lipid extraction and purification[J].Canadian journal of biochemistry and physiology,37(8):911-917.
BOSSIO D A,SCOW K M,GUNAPALA N,et al.,1998.Determinants of soil microbial communities:effects of agricultural management,season,and soil type on phospholipid fatty acid profiles[J].Microbial Ecology,36(1):1-12.
BREUER L R,KIESE R,BUTTERBACH-BAHL K,2002.Temperature and moisture effects on nitrification rates in tropical rainforest soils[J].Soil Science Society of America Journal,66(3):399-402.
BROOKES P C,LANDMAN A,PRUDEN G,et al.,1985.Chloroform fumigation and released of soil nitrogen a rapid direct extraction method to measure microbial biomass nitrogen in soil[J].Soil Biology and Biochemistry,17(6):837-842.
CONANT R T,RYAN M G,AGREN G I,et al.,2011.Temperature and soil organic matter decomposition rates-synthesis of current knowledge and a way forward[J].Global Change Biology,17(11):3392-3404.
FANG X,ZHOU G Y,LI Y L,et al.,2016.Warming effects on biomass and composition of microbial communities and enzyme activities within soil aggregates in subtropical forest[J].Biology and Fertility of Soils,52(3):353-365.
FREY S D,DRIJBER R,SMITH H,et al.,2008.Microbial biomass,functional capacity,and community structure after 12 years of soil warming[J].Soil Biology&Biochemistry,40(11):2904-2907.
FROSTEG?RD A,BAATH E,1996.The use of phospholipid fatty acid analysis to estimate bacterial and fungal biomass in soil[J].Biology&Fertility of Soils,22(1-2):59-65.
FU G,ZHANG H R,LI S W,et al.,2019.A Meta-analysis of the effects of warming and elevated CO2 on soil microbes[J].Journal of Resources and Ecology,10(1):69-76.
INGWERSEN J K,BUTTERHACH-BAHL K,GASCHE R,et al.,1999.Barinetric process separation:New method for quantifying nitrification,denitrification,and nitrous oxide sources in soils[J].Soil Science Society of Americia Journal,63(1):117-128
IPCC,2018:Summary for Policymakers.In:Global Warming of 1.5℃.An IPCC Special Report on the impacts of global warming of 1.5℃above pre-industrial levels and related global greenhouse gas emission pathways,in the context of strengthening the global response to the threat of climate change,sustainable development,and efforts to eradicate poverty.MASSON-DELMOTTE,ZHAI P,PORTNER H-O,ROBERTS D,et al.,Eds.World Meteorological Organization,Geneva,Switzerland,32 pp.
OLSSON P A,1999.Signature fatty acids provide tools for determination of the distribution and interactions of mycorrhizal fungi in soil[J].FEMSMicrobiology Ecology,29(4):303-310.
RILLIG M C,MUMMEY D L,2006.Mycorrhizas and soil structure[J].New Phytologist,171(1):41-53.
RINNAN R,BAATH E,2009.Differential utilization of carbon substrates by bacteria and fungi in tundra soil[J].Applied and Environmental Microbiology,75(11):3611-3620.
SCHINDLBACHER A,RODLER A,KUFFNER M,2011.Experimental warming effects on the microbial community of a temperate mountain forest soil[J].Soil Biology&Biochemistry,43(7):1417-1425.
SCHLESINGER W H,1977.Carbon balance in terrestrial detritus[J].Annual Review of Ecology&Systematics,8(1):51-81.
SEDJO R A,1993.The carbon cycle and global forest ecosystem[J].Water Air and Soil Pollution,70(1-4):295-307.
TUNLID A H,HOITINK A C,LOW,et al.,1989.Characterization of bacteria that suppress rhizoctonia damping-off in bark compost media by analysis of Fatty Acid biomarkers[J].Applied&Environmental Microbiology,55(6):1368-1374.
VANCE E D,BROOKES P C,JENKINSON D S,1987.An extraction method for measuring soil microbial biomass C[J].Soil Biology&Biochemistry,19(6):703-707.
WALTHER G R,POST E P,CONVEY,et al.,2002.Ecological responses to recent climate change[J].Nature,416(6879):389-395.
WILKINSON S C,ANDERSON J M,SCARDELIS S P,et al.,2002.PLFAprofiles of microbial communities in decomposing conifer litters subject to moisture stress[J].Soil Biology&Biochemistry,34(2):189-200.
YU H Y,MA Q H,LIU X D,et al.,2018.Short-and long-term warming alters soil microbial community and relates to soil traits[J].Applied Soil Ecology,131:22-28.
ZHANG Q C,SHAMSI I H,XU D T,et al.,2012.Chemical fertilizer and organic manure inputs in soil exhibit a vice versa pattern of microbial community structure[J].Applied Soil Ecology,57:1-8.5
ZHANG W,PARKER K M,LUO Y,et al.,2005.Soil microbial responses to experimental warming and clipping in a tallgrass prairie[J].Global Change Biology,11(2):266-277.
ZHOU G Y,GUAN L L,WEI X H,et al.,2007.Litterfall production along successional and altitudinal gradients of subtropical monsoon evergreen broadleaved forests in Guangdong,China[J].Plant Ecology,188(1):77-89.
ZIEGLER S E,BILLINGS S A,LANE C S,et al.,2013.Warming alters routing of labile and slower-turnover carbon through distinct microbial groups in boreal forest organic soils[J].Soil Biology&Biochemistry,60:23-32.
鲍士旦,2010.土壤农化分析[M].北京:中国农业出版社.BAO S D,2000.Soil agro-chemistrical analysis[M].Beijing:China Agriculture Press.
李倩,马琨,冶秀香,等,2018.不同培肥方式对土壤有机碳与微生物群落结构的影响[J].中国生态农业学报,26(12):1866-1875.LI Q,MA K,YE X X,et al.,2018.Effect of fertilization managements on soil organic carbon and microbial community structure[J].Chinese Journal of Eco-Agriculture,26(12):1866-1875.
李义勇,2016.南亚热带针阔叶混交林植物生理生态过程对气候变暖的响应[D].广州:中国科学院华南植物园.LI Y Y,2016.Responses of plant physiological and ecological processes in coniferous-broadleaved mixed forest to climate warming[D].Guangzhou:South China Botanical Garden,CAS.
刘光崧,1996.土壤理化分析与剖面描述[M].北京:中国标准出版社.LIU G S,1996.Soil physical and chemical analysis and description of soil profiles[M].Beijing:Standards Press of China.
刘菊秀,李跃林,刘世忠,等,2013.气温上升对模拟森林生态系统影响实验的介绍[J].植物生态学报,37(6):558-565.LIU J X,LI Y L,LIU S Z,et al.,2013.An introduction to an experimental design for studying effects of air temperature rise on model forest ecosystems[J].Chinese Journal of Plant Ecology,37(6):558-565.
沈菊培,贺纪正,2011.微生物介导的碳氮循环过程对全球气候变化的响应[J].生态学报,31(11):2957-2967.SHEN J P,HE J Z,2011.Responses of microbes-mediated carbon and nitrogen cycles to global climate change[J].Acta Ecologica Sinica,31(11):2957-2967.
卫云燕,尹华军,刘庆,等,2009.气候变暖背景下森林土壤碳循环研究进展[J].应用与环境生物学报,15(6):888-894.WEI Y Y,YIN H J,LIU Q,et al.,2009.Advance in research of forest carbon cycling under climate warming[J].Chinese Journal of Applied and Environmental Biology,15(6):888-894.
张明乾韩证仿陈金,等,2012.夜间增温对冬小麦土壤微生物量碳氮及其活性的影响[J].中国生态农业学报,20(11):1464-1470.ZHANG M Q,HAN Z F,CHEN J,et al.,2012.Impact of night time warming on soil microbial biomass carbon/nitrogen and activity in main winter wheat cropping areas in China[J].Chinese Journal of Eco-Agriculture,20(11):1464-1470.
BLIGH E,1959.A rapid method of total lipid extraction and purification[J].Canadian journal of biochemistry and physiology,37(8):911-917.
BOSSIO D A,SCOW K M,GUNAPALA N,et al.,1998.Determinants of soil microbial communities:effects of agricultural management,season,and soil type on phospholipid fatty acid profiles[J].Microbial Ecology,36(1):1-12.
BREUER L R,KIESE R,BUTTERBACH-BAHL K,2002.Temperature and moisture effects on nitrification rates in tropical rainforest soils[J].Soil Science Society of America Journal,66(3):399-402.
BROOKES P C,LANDMAN A,PRUDEN G,et al.,1985.Chloroform fumigation and released of soil nitrogen a rapid direct extraction method to measure microbial biomass nitrogen in soil[J].Soil Biology and Biochemistry,17(6):837-842.
CONANT R T,RYAN M G,AGREN G I,et al.,2011.Temperature and soil organic matter decomposition rates-synthesis of current knowledge and a way forward[J].Global Change Biology,17(11):3392-3404.
FANG X,ZHOU G Y,LI Y L,et al.,2016.Warming effects on biomass and composition of microbial communities and enzyme activities within soil aggregates in subtropical forest[J].Biology and Fertility of Soils,52(3):353-365.
FREY S D,DRIJBER R,SMITH H,et al.,2008.Microbial biomass,functional capacity,and community structure after 12 years of soil warming[J].Soil Biology&Biochemistry,40(11):2904-2907.
FROSTEG?RD A,BAATH E,1996.The use of phospholipid fatty acid analysis to estimate bacterial and fungal biomass in soil[J].Biology&Fertility of Soils,22(1-2):59-65.
FU G,ZHANG H R,LI S W,et al.,2019.A Meta-analysis of the effects of warming and elevated CO2 on soil microbes[J].Journal of Resources and Ecology,10(1):69-76.
INGWERSEN J K,BUTTERHACH-BAHL K,GASCHE R,et al.,1999.Barinetric process separation:New method for quantifying nitrification,denitrification,and nitrous oxide sources in soils[J].Soil Science Society of Americia Journal,63(1):117-128
IPCC,2018:Summary for Policymakers.In:Global Warming of 1.5℃.An IPCC Special Report on the impacts of global warming of 1.5℃above pre-industrial levels and related global greenhouse gas emission pathways,in the context of strengthening the global response to the threat of climate change,sustainable development,and efforts to eradicate poverty.MASSON-DELMOTTE,ZHAI P,PORTNER H-O,ROBERTS D,et al.,Eds.World Meteorological Organization,Geneva,Switzerland,32 pp.
OLSSON P A,1999.Signature fatty acids provide tools for determination of the distribution and interactions of mycorrhizal fungi in soil[J].FEMSMicrobiology Ecology,29(4):303-310.
RILLIG M C,MUMMEY D L,2006.Mycorrhizas and soil structure[J].New Phytologist,171(1):41-53.
RINNAN R,BAATH E,2009.Differential utilization of carbon substrates by bacteria and fungi in tundra soil[J].Applied and Environmental Microbiology,75(11):3611-3620.
SCHINDLBACHER A,RODLER A,KUFFNER M,2011.Experimental warming effects on the microbial community of a temperate mountain forest soil[J].Soil Biology&Biochemistry,43(7):1417-1425.
SCHLESINGER W H,1977.Carbon balance in terrestrial detritus[J].Annual Review of Ecology&Systematics,8(1):51-81.
SEDJO R A,1993.The carbon cycle and global forest ecosystem[J].Water Air and Soil Pollution,70(1-4):295-307.
TUNLID A H,HOITINK A C,LOW,et al.,1989.Characterization of bacteria that suppress rhizoctonia damping-off in bark compost media by analysis of Fatty Acid biomarkers[J].Applied&Environmental Microbiology,55(6):1368-1374.
VANCE E D,BROOKES P C,JENKINSON D S,1987.An extraction method for measuring soil microbial biomass C[J].Soil Biology&Biochemistry,19(6):703-707.
WALTHER G R,POST E P,CONVEY,et al.,2002.Ecological responses to recent climate change[J].Nature,416(6879):389-395.
WILKINSON S C,ANDERSON J M,SCARDELIS S P,et al.,2002.PLFAprofiles of microbial communities in decomposing conifer litters subject to moisture stress[J].Soil Biology&Biochemistry,34(2):189-200.
YU H Y,MA Q H,LIU X D,et al.,2018.Short-and long-term warming alters soil microbial community and relates to soil traits[J].Applied Soil Ecology,131:22-28.
ZHANG Q C,SHAMSI I H,XU D T,et al.,2012.Chemical fertilizer and organic manure inputs in soil exhibit a vice versa pattern of microbial community structure[J].Applied Soil Ecology,57:1-8.5
ZHANG W,PARKER K M,LUO Y,et al.,2005.Soil microbial responses to experimental warming and clipping in a tallgrass prairie[J].Global Change Biology,11(2):266-277.
ZHOU G Y,GUAN L L,WEI X H,et al.,2007.Litterfall production along successional and altitudinal gradients of subtropical monsoon evergreen broadleaved forests in Guangdong,China[J].Plant Ecology,188(1):77-89.
ZIEGLER S E,BILLINGS S A,LANE C S,et al.,2013.Warming alters routing of labile and slower-turnover carbon through distinct microbial groups in boreal forest organic soils[J].Soil Biology&Biochemistry,60:23-32.
鲍士旦,2010.土壤农化分析[M].北京:中国农业出版社.BAO S D,2000.Soil agro-chemistrical analysis[M].Beijing:China Agriculture Press.
李倩,马琨,冶秀香,等,2018.不同培肥方式对土壤有机碳与微生物群落结构的影响[J].中国生态农业学报,26(12):1866-1875.LI Q,MA K,YE X X,et al.,2018.Effect of fertilization managements on soil organic carbon and microbial community structure[J].Chinese Journal of Eco-Agriculture,26(12):1866-1875.
李义勇,2016.南亚热带针阔叶混交林植物生理生态过程对气候变暖的响应[D].广州:中国科学院华南植物园.LI Y Y,2016.Responses of plant physiological and ecological processes in coniferous-broadleaved mixed forest to climate warming[D].Guangzhou:South China Botanical Garden,CAS.
刘光崧,1996.土壤理化分析与剖面描述[M].北京:中国标准出版社.LIU G S,1996.Soil physical and chemical analysis and description of soil profiles[M].Beijing:Standards Press of China.
刘菊秀,李跃林,刘世忠,等,2013.气温上升对模拟森林生态系统影响实验的介绍[J].植物生态学报,37(6):558-565.LIU J X,LI Y L,LIU S Z,et al.,2013.An introduction to an experimental design for studying effects of air temperature rise on model forest ecosystems[J].Chinese Journal of Plant Ecology,37(6):558-565.
沈菊培,贺纪正,2011.微生物介导的碳氮循环过程对全球气候变化的响应[J].生态学报,31(11):2957-2967.SHEN J P,HE J Z,2011.Responses of microbes-mediated carbon and nitrogen cycles to global climate change[J].Acta Ecologica Sinica,31(11):2957-2967.
卫云燕,尹华军,刘庆,等,2009.气候变暖背景下森林土壤碳循环研究进展[J].应用与环境生物学报,15(6):888-894.WEI Y Y,YIN H J,LIU Q,et al.,2009.Advance in research of forest carbon cycling under climate warming[J].Chinese Journal of Applied and Environmental Biology,15(6):888-894.
张明乾韩证仿陈金,等,2012.夜间增温对冬小麦土壤微生物量碳氮及其活性的影响[J].中国生态农业学报,20(11):1464-1470.ZHANG M Q,HAN Z F,CHEN J,et al.,2012.Impact of night time warming on soil microbial biomass carbon/nitrogen and activity in main winter wheat cropping areas in China[J].Chinese Journal of Eco-Agriculture,20(11):1464-1470.
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