整合分析增温和增CO_2对土壤微生物的影响（英文）
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摘要
土壤微生物在陆地生态系统碳氮循环中起着重要作用。气候变暖和CO_2浓度增加是气候变化的两个重要方面。本研究整合分析了实验增温和CO_2浓度增加对土壤微生物量和群落结构的影响。生态系统类型主要包括森林生态系统和草地生态系统。增温方法包括开顶式增温小室和热红外增温。增温时间有全天增温、白天增温和晚上增温。实验增温增加了土壤放线菌和腐生真菌,而CO_2浓度增加减少了土壤革兰氏阳性细菌。实验增温对土壤革兰氏阴性细菌和总的磷脂脂肪酸量的影响随着年均温和年降水量的增加而减少。实验增温对土壤总的磷脂脂肪酸量、细菌含量、革兰氏阳性和阴性细菌的量的影响随着海拔的升高而增加。实验增温增加了草地生态系统的土壤总的磷脂脂肪酸量和放线菌含量,并增加了森林生态系统的土壤真菌和细菌的比值。开顶式增温小室增加了土壤革兰氏阴性细菌,而红外增温减少了土壤真菌和细菌的比值。白天增温增加了土壤革兰氏阴性细菌,而全天增温没有改变土壤革兰氏阴性细菌。因此,实验增温对土壤微生物的影响与生态系统类型、实验增温方法、增温时间、海拔和当地的气候条件有关。
Soil microbes play important roles in terrestrial ecosystem carbon and nitrogen cycling. Climatic warming and elevated CO_2 are two aspects of climatic change. In this study, we used a meta-analysis approach to synthesise observations related to the effects of warming and elevated CO_2 on soil microbial biomass and community structure. Ecosystem types were mainly grouped into forests and grasslands. Warming methods included open top chambers and infrared radiators. Experimental settings included all-day warming, daytime warming and nighttime warming. Warming increased soil actinomycetes and saprotrophic fungi, while elevated CO_2 decreased soil gram-positive bacteria(G~+). Mean annual temperature and mean annual precipitation were negatively correlated with warming effects on gram-negative bacteria(G~–) and total phospholipid fatty acid(PLFA), respectively. Elevation was positively correlated with the warming effect on total PLFA, bacteria, G~+ and G~–. Grassland exhibited a positive response of total PLFA and actinomycetes to warming, while forest exhibited a positive response in the ratio of soil fungi to bacteria(F/B ratio) to warming. The open top chamber method increased G~–, while the infrared radiator method decreased the F/B ratio. Daytime warming rather than all-day warming increased G~–. Our findings indicated that the effects of warming on soil microbes differed with ecosystem types, warming methods, warming times, elevation and local climate conditions.
Soil microbes play important roles in terrestrial ecosystem carbon and nitrogen cycling. Climatic warming and elevated CO_2 are two aspects of climatic change. In this study, we used a meta-analysis approach to synthesise observations related to the effects of warming and elevated CO_2 on soil microbial biomass and community structure. Ecosystem types were mainly grouped into forests and grasslands. Warming methods included open top chambers and infrared radiators. Experimental settings included all-day warming, daytime warming and nighttime warming. Warming increased soil actinomycetes and saprotrophic fungi, while elevated CO_2 decreased soil gram-positive bacteria(G~+). Mean annual temperature and mean annual precipitation were negatively correlated with warming effects on gram-negative bacteria(G~–) and total phospholipid fatty acid(PLFA), respectively. Elevation was positively correlated with the warming effect on total PLFA, bacteria, G~+ and G~–. Grassland exhibited a positive response of total PLFA and actinomycetes to warming, while forest exhibited a positive response in the ratio of soil fungi to bacteria(F/B ratio) to warming. The open top chamber method increased G~–, while the infrared radiator method decreased the F/B ratio. Daytime warming rather than all-day warming increased G~–. Our findings indicated that the effects of warming on soil microbes differed with ecosystem types, warming methods, warming times, elevation and local climate conditions.
引文
Allison S D,Treseder K K.2008.Warming and drying suppress microbial activity and carbon cycling in boreal forest soils.Global Change Biology,14(12):2898-2909.
Andresen L C,Dungait J A J,Bol R,et al.2014.Bacteria and fungi respond differently to multifactorial climate change in a temperate heathland,traced with 13C-glycine and FACE CO2.PLoS ONE,9(1):e85070.doi:85010.81371/journal.pone.0085070.
Baath E,Anderson T H.2003.Comparison of soil fungal/bacterial ratios in a pH gradient using physiological and PLFA-based techniques.Soil Biology&Biochemistry,35(7):955-963.
Bai E,Li S L,Xu W H,et al.2013.A meta-analysis of experimental warming effects on terrestrial nitrogen pools and dynamics.New Phytologist,199(2):441-451.
Bardgett R D,Hobbs P J,Frostegard A 1996.Changes in soil fungal:bacterial biomass ratios following reductions in the intensity of management of an upland grassland.Biology and Fertility of Soils,22(3):261-264.
Blankinship J C,Niklaus P A,Hungate B A 2011.A meta-analysis of responses of soil biota to global change.Oecologia,165(3):553-565.
Chen J,Luo Y Q,Xia J Y,et al.2015.Stronger warming effects on microbial abundances in colder regions.Scientific Reports,5:18032.
de Graaff M A,van Groenigen K J,Six J,et al.2006.Interactions between plant growth and soil nutrient cycling under elevated CO2:a metaanalysis.Global Change Biology,12(11):2077-2091.
Frostegard A,Tunlid A,Baath E.1993.Phospholipid Fatty Acid composition,biomass,and activity of microbial communities from two soil types experimentally exposed to different heavy metals.Applied and Environmental Microbiology,59(11):3605-3617.
Fu G,Shen Z X.2016.Response of alpine plants to nitrogen addition on the Tibetan Plateau:A meta-analysis.Journal of Plant Growth Regulation,35(4):974-979.
Fu G,Shen Z X.2017a.Effects of enhanced UV-B radiation on plant physiology and growth on the Tibetan Plateau:a meta-analysis.Acta Physiologiae Plantarum,39(3):doi:10.1007/s11738-11017-12387-11738.
Fu G,Shen Z X.2017b.Grazing alters soil microbial community in alpine grasslands of the Northern Tibet.Acta Aprataculturae Sinica,26(10):170-178.
Fu G,Shen Z X.2017c.Response of alpine soils to nitrogen addition on the Tibetan Plateau:A meta-analysis.Applied Soil Ecology,114(99-104.
Fu G,Shen Z X,Sun W,et al.2015.A meta-analysis of the effects of experimental warming on plant physiology and growth on the Tibetan Plateau.Journal of Plant Growth Regulation,34(1):57-65.
Gutknecht J L M,Field C B,Balser T C.2012.Microbial communities and their responses to simulated global change fluctuate greatly over multiple years.Global Change Biology,18(7):2256-2269.
Hedges L V,Gurevitch J,Curtis P S.1999.The meta-analysis of response ratios in experimental ecology.Ecology,80(4):1150-1156.
Hu S,Chapin F S,Firestone M K,et al.2001.Nitrogen limitation of microbial decomposition in a grassland under elevated CO2.Nature,409(6817):188-191.
Huang X M,Liu S R,Wang H,et al.2014.Changes of soil microbial biomass carbon and community composition through mixing nitrogen-fixing species with Eucalyptus urophylla in subtropical China.Soil Biology&Biochemistry,73:42-48.
IPCC.2013.Summary for Policymakers.In:Climate Change 2013:The Physical Science Basis.Contribution of Working Group I to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change[Stocker,T F,D Qin,G K Plattner,M.Tignor,S.K.Allen,J.Boschung,A.Nauels,Y.Xia,V.Bex and P.M.Midgley(eds.)].Cambridge University Press,Cambridge,United Kingdom and New York,NY,USA.
Kanerva T,Palojarvi A,Ramo K,et al.2008.Changes in soil microbial community structure under elevated tropospheric O3 and CO2.Soil Biology&Biochemistry,40(10):2502-2510.
Lange M,Habekost M,Eisenhauer N,et al.2014.Biotic and abiotic properties mediating plant diversity effects on soil microbial communities in an experimental grassland.PLoS ONE,9(5):e96182.doi:96110.91371/journal.pone.0096182.
Li Q,Bai H H,Liang W J,et al.2013.Nitrogen addition and warming independently influence the belowground micro-food web in a temperate steppe.PLoS ONE,8(3):doi:10.1371/journal.pone.0060441.
Liu Y,Li M,Zheng J W,et al.2014.Short-term responses of microbial community and functioning to experimental CO2 enrichment and warming in a Chinese paddy field.Soil Biology&Biochemistry,77:58-68.
Lu M,Zhou X H,Yang Q,et al.2013.Responses of ecosystem carbon cycle to experimental warming:a meta-analysis.Ecology,94(3):726-738.
Luo Y Q,Hui D F,Zhang D Q.2006.Elevated CO2 stimulates net accumulations of carbon and nitrogen in land ecosystems:A meta-analysis.Ecology,87(1):53-63.
Ma L N,Lu X T,Liu Y,et al.2011.The effects of warming and nitrogen addition on soil nitrogen cycling in a temperate grassland,Northeastern China.PLoS ONE,6(11):e27645.doi:27610.21371/journal.pone.0027645.
Rinnan R,Michelsen A,Baath E,et al.2007.Fifteen years of climate change manipulations alter soil microbial communities in a subarctic heath ecosystem.Global Change Biology,13(1):28-39.
Ross D J,Newton P C D,Tate K R,et al.2013.Impact of a low level of CO2 enrichment on soil carbon and nitrogen pools and mineralization rates over ten years in a seasonally dry,grazed pasture.Soil Biology&Biochemistry:58(265-274.
Rousk J,Smith A R,Jones D L.2013.Investigating the long-term legacy of drought and warming on the soil microbial community across five European shrubland ecosystems.Global Change Biology,19(12):3872-3884.
Schindlbacher A,Rodler A,Kuffner M,et al.2011.Experimental warming effects on the microbial community of a temperate mountain forest soil.Soil Biology&Biochemistry,43(7):1417-1425.
Shen R C,Xu M,Chi Y G,et al.2014.Soil microbial responses to experimental warming and nitrogen addition in a temperate steppe of Northern China.Pedosphere,24(4):427-436.
Wang X J,Zhou Y M,Jiang X J,et al.2014.Effects of warming on soil microbial community structure in Changbai Mountain tundra.Acta Ecologica Sinica,34(20):5706-5713.
Yang Y,Luo Y,Lu M,et al.2011.Terrestrial C:N stoichiometry in response to elevated CO2 and N addition:a synthesis of two meta-analyses.Plant and Soil,343(1-2):393-400.
Zhang B,Chen S,He X,et al.2014.Responses of soil microbial communities to experimental warming in alpine grasslands on the Qinghai-Tibet Plateau.PLoS ONE,9(8):e103859.doi:103810.101371/journal.pone.0103859.
Zhang N,Liu W,Yang H,et al.2013.Soil microbial responses to warming and increased precipitation and their implications for ecosystem C cycling.Oecologia,173(3):1125-1142.
Zhang W,Parker K M,Luo Y,et al.2005.Soil microbial responses to experimental warming and clipping in a tallgrass prairie.Global Change Biology,11(2):266-277.
Zhang W J,Xu Q,Wang X K,et al.2004.Impacts of experimental atmospheric warming on soil microbial community structure in a tallgrass prairie.Acta Ecologica Sinica,24(8):1742-1747.
Zhang X Z,Shen Z X,Fu G.2015.A meta-analysis of the effects of experimental warming on soil carbon and nitrogen dynamics on the Tibetan Plateau.Applied Soil Ecology,87:32-38.
Zhao C Z,Zhu L Y,Liang J,et al.2014.Effects of experimental warming and nitrogen fertilization on soil microbial communities and processes of two subalpine coniferous species in Eastern Tibetan Plateau,China.Plant and Soil,382(1-2):189-201.
Zhou J Z,Xue K,Xie J P,et al.2012.Microbial mediation of carboncycle feedbacks to climate warming.Nature Climate Change,2(2):106-110.
Andresen L C,Dungait J A J,Bol R,et al.2014.Bacteria and fungi respond differently to multifactorial climate change in a temperate heathland,traced with 13C-glycine and FACE CO2.PLoS ONE,9(1):e85070.doi:85010.81371/journal.pone.0085070.
Baath E,Anderson T H.2003.Comparison of soil fungal/bacterial ratios in a pH gradient using physiological and PLFA-based techniques.Soil Biology&Biochemistry,35(7):955-963.
Bai E,Li S L,Xu W H,et al.2013.A meta-analysis of experimental warming effects on terrestrial nitrogen pools and dynamics.New Phytologist,199(2):441-451.
Bardgett R D,Hobbs P J,Frostegard A 1996.Changes in soil fungal:bacterial biomass ratios following reductions in the intensity of management of an upland grassland.Biology and Fertility of Soils,22(3):261-264.
Blankinship J C,Niklaus P A,Hungate B A 2011.A meta-analysis of responses of soil biota to global change.Oecologia,165(3):553-565.
Chen J,Luo Y Q,Xia J Y,et al.2015.Stronger warming effects on microbial abundances in colder regions.Scientific Reports,5:18032.
de Graaff M A,van Groenigen K J,Six J,et al.2006.Interactions between plant growth and soil nutrient cycling under elevated CO2:a metaanalysis.Global Change Biology,12(11):2077-2091.
Frostegard A,Tunlid A,Baath E.1993.Phospholipid Fatty Acid composition,biomass,and activity of microbial communities from two soil types experimentally exposed to different heavy metals.Applied and Environmental Microbiology,59(11):3605-3617.
Fu G,Shen Z X.2016.Response of alpine plants to nitrogen addition on the Tibetan Plateau:A meta-analysis.Journal of Plant Growth Regulation,35(4):974-979.
Fu G,Shen Z X.2017a.Effects of enhanced UV-B radiation on plant physiology and growth on the Tibetan Plateau:a meta-analysis.Acta Physiologiae Plantarum,39(3):doi:10.1007/s11738-11017-12387-11738.
Fu G,Shen Z X.2017b.Grazing alters soil microbial community in alpine grasslands of the Northern Tibet.Acta Aprataculturae Sinica,26(10):170-178.
Fu G,Shen Z X.2017c.Response of alpine soils to nitrogen addition on the Tibetan Plateau:A meta-analysis.Applied Soil Ecology,114(99-104.
Fu G,Shen Z X,Sun W,et al.2015.A meta-analysis of the effects of experimental warming on plant physiology and growth on the Tibetan Plateau.Journal of Plant Growth Regulation,34(1):57-65.
Gutknecht J L M,Field C B,Balser T C.2012.Microbial communities and their responses to simulated global change fluctuate greatly over multiple years.Global Change Biology,18(7):2256-2269.
Hedges L V,Gurevitch J,Curtis P S.1999.The meta-analysis of response ratios in experimental ecology.Ecology,80(4):1150-1156.
Hu S,Chapin F S,Firestone M K,et al.2001.Nitrogen limitation of microbial decomposition in a grassland under elevated CO2.Nature,409(6817):188-191.
Huang X M,Liu S R,Wang H,et al.2014.Changes of soil microbial biomass carbon and community composition through mixing nitrogen-fixing species with Eucalyptus urophylla in subtropical China.Soil Biology&Biochemistry,73:42-48.
IPCC.2013.Summary for Policymakers.In:Climate Change 2013:The Physical Science Basis.Contribution of Working Group I to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change[Stocker,T F,D Qin,G K Plattner,M.Tignor,S.K.Allen,J.Boschung,A.Nauels,Y.Xia,V.Bex and P.M.Midgley(eds.)].Cambridge University Press,Cambridge,United Kingdom and New York,NY,USA.
Kanerva T,Palojarvi A,Ramo K,et al.2008.Changes in soil microbial community structure under elevated tropospheric O3 and CO2.Soil Biology&Biochemistry,40(10):2502-2510.
Lange M,Habekost M,Eisenhauer N,et al.2014.Biotic and abiotic properties mediating plant diversity effects on soil microbial communities in an experimental grassland.PLoS ONE,9(5):e96182.doi:96110.91371/journal.pone.0096182.
Li Q,Bai H H,Liang W J,et al.2013.Nitrogen addition and warming independently influence the belowground micro-food web in a temperate steppe.PLoS ONE,8(3):doi:10.1371/journal.pone.0060441.
Liu Y,Li M,Zheng J W,et al.2014.Short-term responses of microbial community and functioning to experimental CO2 enrichment and warming in a Chinese paddy field.Soil Biology&Biochemistry,77:58-68.
Lu M,Zhou X H,Yang Q,et al.2013.Responses of ecosystem carbon cycle to experimental warming:a meta-analysis.Ecology,94(3):726-738.
Luo Y Q,Hui D F,Zhang D Q.2006.Elevated CO2 stimulates net accumulations of carbon and nitrogen in land ecosystems:A meta-analysis.Ecology,87(1):53-63.
Ma L N,Lu X T,Liu Y,et al.2011.The effects of warming and nitrogen addition on soil nitrogen cycling in a temperate grassland,Northeastern China.PLoS ONE,6(11):e27645.doi:27610.21371/journal.pone.0027645.
Rinnan R,Michelsen A,Baath E,et al.2007.Fifteen years of climate change manipulations alter soil microbial communities in a subarctic heath ecosystem.Global Change Biology,13(1):28-39.
Ross D J,Newton P C D,Tate K R,et al.2013.Impact of a low level of CO2 enrichment on soil carbon and nitrogen pools and mineralization rates over ten years in a seasonally dry,grazed pasture.Soil Biology&Biochemistry:58(265-274.
Rousk J,Smith A R,Jones D L.2013.Investigating the long-term legacy of drought and warming on the soil microbial community across five European shrubland ecosystems.Global Change Biology,19(12):3872-3884.
Schindlbacher A,Rodler A,Kuffner M,et al.2011.Experimental warming effects on the microbial community of a temperate mountain forest soil.Soil Biology&Biochemistry,43(7):1417-1425.
Shen R C,Xu M,Chi Y G,et al.2014.Soil microbial responses to experimental warming and nitrogen addition in a temperate steppe of Northern China.Pedosphere,24(4):427-436.
Wang X J,Zhou Y M,Jiang X J,et al.2014.Effects of warming on soil microbial community structure in Changbai Mountain tundra.Acta Ecologica Sinica,34(20):5706-5713.
Yang Y,Luo Y,Lu M,et al.2011.Terrestrial C:N stoichiometry in response to elevated CO2 and N addition:a synthesis of two meta-analyses.Plant and Soil,343(1-2):393-400.
Zhang B,Chen S,He X,et al.2014.Responses of soil microbial communities to experimental warming in alpine grasslands on the Qinghai-Tibet Plateau.PLoS ONE,9(8):e103859.doi:103810.101371/journal.pone.0103859.
Zhang N,Liu W,Yang H,et al.2013.Soil microbial responses to warming and increased precipitation and their implications for ecosystem C cycling.Oecologia,173(3):1125-1142.
Zhang W,Parker K M,Luo Y,et al.2005.Soil microbial responses to experimental warming and clipping in a tallgrass prairie.Global Change Biology,11(2):266-277.
Zhang W J,Xu Q,Wang X K,et al.2004.Impacts of experimental atmospheric warming on soil microbial community structure in a tallgrass prairie.Acta Ecologica Sinica,24(8):1742-1747.
Zhang X Z,Shen Z X,Fu G.2015.A meta-analysis of the effects of experimental warming on soil carbon and nitrogen dynamics on the Tibetan Plateau.Applied Soil Ecology,87:32-38.
Zhao C Z,Zhu L Y,Liang J,et al.2014.Effects of experimental warming and nitrogen fertilization on soil microbial communities and processes of two subalpine coniferous species in Eastern Tibetan Plateau,China.Plant and Soil,382(1-2):189-201.
Zhou J Z,Xue K,Xie J P,et al.2012.Microbial mediation of carboncycle feedbacks to climate warming.Nature Climate Change,2(2):106-110.
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