三源区分土壤呼吸组分研究
|
摘要
三源区分土壤呼吸组分是指将土壤呼吸区分为纯根呼吸、根际微生物呼吸和土壤有机质呼吸3个部分。土壤有机质呼吸、纯根呼吸和根际微生物呼吸是3种不同的生物学过程,这3种呼吸对环境变化具有不同的响应机制。区分土壤呼吸中由根系引起的自养和异养呼吸组分的研究对定量评价陆地生态系统碳平衡具有重要的意义。论述了三源区分土壤呼吸组分的意义、方法和应用,分析了不同条件下土壤呼吸组分区分的研究结果。实验室纯根和根际微生物呼吸占根源呼吸比重约为45%和55%;野外条件下约为60%和40%。最后对本研究未来的发展方向进行了展望。
Partitioning soil respiration into three components( root respiration,rhizomicrobial respiration,and basal respiration) is called "three-source partitioning of soil respiration ". These components are three different biological processes,and their responses to environment change are different. Therefore,it is important and significant to partition soil respiration into autotrophic and heterotrophic respiration to evaluate terrestrial ecosystem carbon balance quantitatively.Although there has been a general understanding of autotrophic and heterotrophic respiration affected by environment factors,the mechanisms underlying their role in the rhizosphere and their ecological significance are still not fully comprehended. To determine the mechanisms of the rhizosphere system response to environmental change,rhizosphere respiration should be partitioned into root respiration and rhizomicrobial respiration. In the present study,we discussed the significance,methods,and applications of three-source partitioning of soil respiration,and summarized the results under different conditions. Methods of three-source partitioning of soil respiration include physical excised method( such as the root excised method),isotopic tracer continuous labeling( such as13 C natural abundance),and isotopic tracer pulse labeling( such as14 C pulse labeling). The isotopic tracer pulse labeling method is more accurate than the other methods;however,it cannot be utilized in field studies and is expensive to undertake. The physical excised method is cheaper and requires no complex calculation; however,it greatly disturbs the soil and roots,which casts doubts on the relevance of theresults. The isotopic tracer continuous labeling method has advantages for in situ measurements,accurate tracing,and causes almost no disturbance. However,this method can be utilized only under special conditions( such as when the plant organics are from C3 and the basal soil organic matter are from C4) and further study is required to improve its applicability.Therefore,it is difficult to partition soil respiration into the three components under field conditions. The excised roots and trenching method and13 C natural abundance method can be utilized in field studies,especially in forest ecosystems. The results showed that the proportion of root respiration and the proportion of rhizomicrobial respiration to rhizosphere respiration was approximately 45% and 55%,respectively,under laboratory conditions,and approximately 60% and 40%,respectively,under field conditions. Root respiration,rhizomicrobial respiration,and basal respiration varied with climate,depth,soil nutrition,and other environmental factors. Decomposition of rhizoorganisms,which causes rhizomicrobial respiration,has the potential to influence greatly basal respiration. The variation of the three components reflect the turnover rate of soil carbon,and affect the acquisition of competition and symbiosis by plants and microorganisms,thus maintaining nutrient balance among the various components of an ecosystem. The conventional three-source partitioning methods might produce great uncertainty by ignoring the root-microbial system processes in the soil. These processes include the adaptation of soil microbes,rhizosphere priming effects,nutrient partitioning,etc. Rhizomicrobial respiration is an important part of soil respiration in plantations and cannot be ignored. Three-source partitioning of soil respiration is a useful approach to quantitatively evaluate forest underground CO_2 flux as the global climate changes. The outlooks of the present study on the three-source partitioning of soil respiration were also discussed and clarified.
Partitioning soil respiration into three components( root respiration,rhizomicrobial respiration,and basal respiration) is called "three-source partitioning of soil respiration ". These components are three different biological processes,and their responses to environment change are different. Therefore,it is important and significant to partition soil respiration into autotrophic and heterotrophic respiration to evaluate terrestrial ecosystem carbon balance quantitatively.Although there has been a general understanding of autotrophic and heterotrophic respiration affected by environment factors,the mechanisms underlying their role in the rhizosphere and their ecological significance are still not fully comprehended. To determine the mechanisms of the rhizosphere system response to environmental change,rhizosphere respiration should be partitioned into root respiration and rhizomicrobial respiration. In the present study,we discussed the significance,methods,and applications of three-source partitioning of soil respiration,and summarized the results under different conditions. Methods of three-source partitioning of soil respiration include physical excised method( such as the root excised method),isotopic tracer continuous labeling( such as13 C natural abundance),and isotopic tracer pulse labeling( such as14 C pulse labeling). The isotopic tracer pulse labeling method is more accurate than the other methods;however,it cannot be utilized in field studies and is expensive to undertake. The physical excised method is cheaper and requires no complex calculation; however,it greatly disturbs the soil and roots,which casts doubts on the relevance of theresults. The isotopic tracer continuous labeling method has advantages for in situ measurements,accurate tracing,and causes almost no disturbance. However,this method can be utilized only under special conditions( such as when the plant organics are from C3 and the basal soil organic matter are from C4) and further study is required to improve its applicability.Therefore,it is difficult to partition soil respiration into the three components under field conditions. The excised roots and trenching method and13 C natural abundance method can be utilized in field studies,especially in forest ecosystems. The results showed that the proportion of root respiration and the proportion of rhizomicrobial respiration to rhizosphere respiration was approximately 45% and 55%,respectively,under laboratory conditions,and approximately 60% and 40%,respectively,under field conditions. Root respiration,rhizomicrobial respiration,and basal respiration varied with climate,depth,soil nutrition,and other environmental factors. Decomposition of rhizoorganisms,which causes rhizomicrobial respiration,has the potential to influence greatly basal respiration. The variation of the three components reflect the turnover rate of soil carbon,and affect the acquisition of competition and symbiosis by plants and microorganisms,thus maintaining nutrient balance among the various components of an ecosystem. The conventional three-source partitioning methods might produce great uncertainty by ignoring the root-microbial system processes in the soil. These processes include the adaptation of soil microbes,rhizosphere priming effects,nutrient partitioning,etc. Rhizomicrobial respiration is an important part of soil respiration in plantations and cannot be ignored. Three-source partitioning of soil respiration is a useful approach to quantitatively evaluate forest underground CO_2 flux as the global climate changes. The outlooks of the present study on the three-source partitioning of soil respiration were also discussed and clarified.
引文
[1]宋文琛.自然13C丰度法区分剌槐人工林土壤呼吸三组分的研究[D].北京:北京林业大学,2015.
[2]刘佳,同小娟,张劲松,孟平,李俊,郑宁.太阳辐射对黄河小浪底人工混交林净生态系统碳交换的影响.生态学报,2014,34(8):2118-2127.
[3]Le QuéréC,Peters G P,Andrew RM,Boden T A,Ciais P,Friedlingstein P,Houghton R A,Marland G,Moriarty R,Sitch S,Tans P,Arneth A,Arvanitis A,Bakker D C E,Bopp L,Canadell J G,Chini L P,Doney S C,Harper A,Harris I,House J I,Jain A K,Jones S D,Kato E,Keeling R F,Goldewijk K,K9rtzinger A,Koven C,Lefèvre N,Maignan F,Omar A,Ono T,Park G H,Pfeil B,Poulter B,Raupach M R,Regnier P,R9denbeck C,Saito S,Schwinger J,Segschneider J,Stocker B,Tilbrook B,Van Heuven S M A C,Viovy N,Wanninkhof R,Wiltshire A,Zaehle S,Yue C.Global carbon budget 2013.Earth System Science Data,2014,6(7):235-263.
[4]Atarashi-Andoh M,Koarashi J,Ishizuka S,Hirai K.Seasonal patterns and control factors of CO2effluxes from surface litter,soil organic carbon,and root-derived carbon estimated using radiocarbon signatures.Agricultural and Forest Meteorology,2012,152:149-158.
[5]Cheng W X,Parton W J,Gonzalez-Meler MA,Phillips R,Asao S,McN ickle G G,Brzostek E,Jastrow J D.Synthesis and modeling perspectives of rhizosphere priming.New Phytologist,2014,201(1):31-44.
[6]Hopkins F,Gonzalez-Meler M A,Flower C E,Lynch D J,Czimczik C,Tang J W,Subke J A.Ecosystem-level controls on root-rhizosphere respiration.New Phytologist,2013,199(2):339-351.
[7]Chapin F S III,Matson P A,Vitousek P M.Principles of Terrestrial Ecosystem Ecology.2nd ed.New York:Springer Press,2012.
[8]Kuzyakov Y,Friedel J K,Stahr K.Review of mechanisms and quantification of priming effects.Soil Biology and Biochemistry,2000,32(11-12):1485-1498.
[9]Kuzyakov Y,Blagodatskaya E V.Microbial hotspots and hot moments in soil:Concept&review.Soil Biology and Biochemistry,2015,83:184-199.
[10]Kuzyakov Y.Priming effects:interactions between living and dead organic matter.Soil Biology and Biochemistry,2010,42(9):1363-1371.
[11]Blagodatskaya E V,Kuzyakov Y.Active microorganisms in soil:critical review of estimation criteria and approaches.Soil Biology and Biochemistry,2013,67:192-211.
[12]Kuzyakov Y.Sources of CO2efflux from soil and review of partitioning methods.Soil Biology and Biochemistry,2006,38(3):425-448.
[13]Werth M,Kuzyakov Y.Three-source partitioning of CO2efflux from maize field soil by13C natural abundance.Journal of Plant Nutrition and Soil Science,2009,172(4):487-499.
[14]Kuzyakov Y V,Larionova A A.Contribution of rhizomicrobial and root respiration to the CO2emission from soil(A Review).Eurasian Soil Science,2006,39(7):753-764.
[15]金钊,董云社,齐玉春.区分纯根呼吸和根际微生物呼吸的争议.土壤,2008,40(4):517-522.
[16]Hanson P J,Edwards N T,Garten C T,Andrews J A.Separating root and soil microbial contributions to soil respiration:A review of methods and observations.Biogeochemistry,2000,48(1):115-146.
[17]Anderson J M.The effects of climate change on decomposition processes in grassland and coniferous forests.Ecological Applications,1991,1(3):326-347.
[18]Li P,Yang Y H,Fang J Y.Variations of root and heterotrophic respiration along environmental gradients in Chinas forests.Journal of Plant Ecology,2013,6(5):358-367.
[19]H9gberg P,Buchmann N,Read D J.Comments on Yakov Kuzyakov's review‘Sources of CO2efflux from soil and review of partitioning methods':[Soil Biology&Biochemistry 38,425–448].Soil Biology and Biochemistry,2006,38(9):2997-2998.
[20]Kuzyakov Y,Larionova A A.Root and rhizomicrobial respiration:A review of approaches to estimate respiration by autotrophic and heterotrophic organisms in soil.Journal of Plant Nutrition and Soil Science,2005,168(4):503-520.
[21]孙悦,徐兴良,Kuzyakov Y.根际激发效应的发生机制及其生态重要性.植物生态学报,2014,38(1):62-75.
[22]Kuzyakov Y,Xu X L.Competition between roots and microorganisms for nitrogen:mechanisms and ecological relevance.New Phytologist,2013,198(3):656-669.
[23]Xu M,Shang H.Contribution of soil respiration to the global carbon equation.Journal of Plant Physiology,2016,203:16-28.
[24]Gerber S,Joos F,Prentice I C.Sensitivity of a dynamic global vegetation model to climate and atmospheric CO2.Global Change Biology,2004,10(8):1223-1239.
[25]Macdonald C A,Anderson I C,Bardgett R D,Singh B K.Role of nitrogen in carbon mitigation in forest ecosystems.Current Opinion in Environmental Sustainability,2011,3(5):303-310.
[26]Phillips R P,Finzi A C,Bernhardt E S.Enhanced root exudation induces microbial feedbacks to N cycling in a pine forest under long-term CO2fumigation.Ecology Letters,2011,14(2):187-194.
[27]Sayer E J,Heard M S,Grant H K,Marthews T R,Tanner E V J.Soil carbon release enhanced by increased tropical forest litter fall.Nature Climate Change,2011,1(6):304-307.
[28]Bengtson P,Barker J,Grayston S J.Evidence of a strong coupling between root exudation,C and N availability,and stimulated SOM decomposition caused by rhizosphere priming effects.Ecology and Evolution,2012,2(8):1843-1852.
[29]Yuan H Z,Ge T D,Chen C Y,O'Donnell A G,Wu J S.Significant role for microbial autotrophy in the sequestration of soil carbon.Applied and Environmental Microbiology,2012,78(7):2328-2336.
[30]Von Lützow M,K9gel-Knabner I.Temperature sensitivity of soil organic matter decomposition-What do we know?Biology and Fertility of Soils,2009,46(1):1-15.
[31]Conant R T,Ryan M G,gren G I,Birge H E,Davidson E A,Eliasson P E,Evans S E,Frey S D,Giardina C P,Hopkins F M,Hyv9nen R,Kirschbaum M U F,Lavallee J M,Leifeld J,Parton W J,Steinweg J M,Wallenstein M D,Wetterstedt JM,Bradford M A.Temperature and soil organic matter decomposition rates-synthesis of current knowledge and a way forward.Global Change Biology,2011,17(11):3392-3404.
[32]王兵,姜艳,郭浩,赵广东,白秀兰.土壤呼吸及其三个生物学过程研究.土壤通报,2011,42(2):483-490.
[33]陈敏鹏,夏旭,李银坤,梅旭荣.土壤呼吸组分分离技术研究进展.生态学报,2013,33(22):7067-7077.
[34]Remus R,Hüve K,P9rschmann J,Augustin J.Determining the timepoint when14C tracer accurately reflect photosynthate use in the plant-soil system.Plant and Soil,2016,408(1-2):457-474.
[35]Remus R,Augustin J.Dynamic linking of14C partitioning with shoot growth allows a precise determination of plant-derived C input to soil.Plant and Soil,2016,408(1-2):493-513.
[36]Carbone M S,Richardson A D,Chen M,Davidson E A,Hughes H,Savage K E,Hollinger D Y.Constrained partitioning of autotrophic and heterotrophic respiration reduces model uncertainties of forest ecosystem carbon fluxes but not stocks.Journal of Geophysical Research:Biogeosciences,2016,121(9):2476-2492.
[37]Kuzyakov Y.Separating microbial respiration of exudates from root respiration in non-sterile soils:A comparison of four methods.Soil Biology and Biochemistry,2002,34(11):1621-1631.
[38]Kelting D L,Burger J A,Edwards G S.Estimating root respiration,microbial respiration in the rhizosphere,and root-free soil respiration in forest soils.Soil Biology and Biochemistry,1998,30(7):961-968.
[39]Chen C R,Condron L M,Xu Z H,Davis M R,Sherlock R R.Root,rhizosphere and root-free respiration in soils under grassland and forest plants.European Journal of Soil Science,2006,57(1):58-66.
[40]Craine J M,Wedin D A,Chapin F S.Predominance of ecophysiological controls on soil CO2flux in a Minnesota grassland.Plant and Soil,1999,207(1):77-86.
[41]Larionova A A,Sapronov D V,de Gerenyu L,Kuznetsova L G,Kudeyarov V N.Contribution of plant root respiration to the CO2emission from soil.Eurasian Soil Science,2006,39(10):1127-1135.
[42]Barba J,Yuste J C,Poyatos R,Janssens I A,Lloret F.Strong resilience of soil respiration components to drought-induced die-off resulting in forest secondary succession.Oecologia,2016,182(1):27-41.
[43]Cheng W X,Zhang Q L,Coleman D C,Carroll C R,Hoffman C A.Is available carbon limiting microbial respiration in the rhizosphere?Soil Biology and Biochemistry,1996,28(10-11):1283-1288.
[44]Kuzyakov Y,Cheng W.Photosynthesis controls of rhizosphere respiration and organic matter decomposition.Soil Biology and Biochemistry,2001,33(14):1915-1925.
[45]Kuzyakov Y,Domanski G.Model for rhizodeposition and CO2efflux from planted soil and its validation by14C pulse labelling of ryegrass.Plant and Soil,2002,239(1):87-102.
[46]Sapronov D V,Kuzyakov Y V.Separation of root and microbial respiration:comparison of three methods.Eurasian Soil Science,2007,40(7):775-784.
[47]O'Leary M H.Carbon isotopes in photosynthesis.Bioscience,1988,38(5):328-336.
[48]Kuzyakov Y.Theoretical background for partitioning of root and rhizomicrobial respiration byδ13C of microbial biomass.European Journal of Soil Biology,2005,41(1-2):1-9.
[49]Werth M,Subbotina I,Kuzyakov Y.Three-source partitioning of CO2efflux from soil planted with maize by13C natural abundance fails due to inactive microbial biomass.Soil Biology and Biochemistry,2006,38(9):2772-2781.
[50]Blagodatskaya E,Khomyakov N,Myachina O,Bogomolova I,Blagodatsky S,Kuzyakov Y.Microbial interactions affect sources of priming induced by cellulose.Soil Biology and Biochemistry,2014,74:39-49.
[51]KerréB,Hernandez-Soriano M C,Smolders E.Partitioning of carbon sources among functional pools to investigate short-term priming effects of biochar in soil:A13C study.Science of Total Environment,2016,547:30-38.
[52]宋文琛,同小娟,张劲松,孟平,李俊.用自然13C丰度法区分人工林根源呼吸的原理与应用.中国水土保持科学,2015,13(4):37-43.
[53]Song W C,Tong X J,Zhang J S,Meng P.Three-source partitioning of soil respiration by13C natural abundance and its variation with soil depth in a plantation.Journal of Forestry Research,2016,27(3):533-540.
[54]TomèE,Ventura M,Folegot S,Zanotelli D,Montagnani L,Mimmo T,Tonon G,Tagliavini M,Scandellari F.Mycorrhizal contribution to soil respiration in an apple orchard.Applied Soil Ecology,2016,101:165-173.
[55]Werth M,Kuzyakov Y.13C fractionation at the root-microorganisms-soil interface:A review and outlook for partitioning studies.Soil Biology and Biochemistry,2010,42(9):1372-1384.
[56]Tian J,Pausch J,Yu G R,Blagodatskaya E,Kuzyakov Y.Aggregate size and glucose level affect priming sources:A three-source-partitioning study.Soil Biology and Biochemistry,2016,97:199-210.
[57]Dijkstra F A,Carrillo Y,Pendall E,Morgan J A.Rhizosphere priming:a nutrient perspective.Frontiers in Microbiology,2013,4(1):65-72.
[58]Sullivan B W,Hart S C.Evaluation of mechanisms controlling the priming of soil carbon along a substrate age gradient.Soil Biology and Biochemistry,2013,58:293-301.
[59]Burns R G,DeF orest J L,Marxsen J,Sinsabaugh R L,Stromberger M E,Wallenstein M D,Weintraub M N,Zoppini A.Soil enzymes in a changing environment:current knowledge and future directions.Soil Biology and Biochemistry,2013,58:216-234.
[60]Spohn M,P9tsch E M,Eichorst S A,Woebken D,Wanek W,Richter A.Soil microbial carbon use efficiency and biomass turnover in a long-term fertilization experiment in a temperate grassland.Soil Biology and Biochemistry,2016,97:168-175.
[61]Zang H D,Wang J Y,Kuzyakov Y.N fertilization decreases soil organic matter decomposition in the rhizosphere.Applied Soil Ecology,2016,108:47-53.
[62]耿元波,史晶晶.碳同位素在草地土壤呼吸区分中的应用.中国农业科学,2012,45(17):3541-3550.
[63]魏书精,罗碧珍,魏书威,孙龙,文正敏,胡海清.森林生态系统土壤呼吸测定方法研究进展.生态环境学报,2014,23(3):504-514.
[64]Qiao N,Schaefer D,Blagodatskaya E,Zou X M,Xu X L,Kuzyakov Y.Labile carbon retention compensates for CO2released by priming in forest soils.Global Change Biology,2014,20(6):1943-1954.
[65]Pan Y D,Birdsey R A,Fang J Y,Houghton R,Kauppi P E,Kurz W A,Phillips O L,Shvidenko A,Lewis S L,Canadell J G,Ciais P,Jackson P B,Pacala S W,McG uire A D,Piao S L,Rautiainen A,Sitch S,Hayes D.A large and persistent carbon sink in the world's forests.Science,2011,333(6045):988-993.
[2]刘佳,同小娟,张劲松,孟平,李俊,郑宁.太阳辐射对黄河小浪底人工混交林净生态系统碳交换的影响.生态学报,2014,34(8):2118-2127.
[3]Le QuéréC,Peters G P,Andrew RM,Boden T A,Ciais P,Friedlingstein P,Houghton R A,Marland G,Moriarty R,Sitch S,Tans P,Arneth A,Arvanitis A,Bakker D C E,Bopp L,Canadell J G,Chini L P,Doney S C,Harper A,Harris I,House J I,Jain A K,Jones S D,Kato E,Keeling R F,Goldewijk K,K9rtzinger A,Koven C,Lefèvre N,Maignan F,Omar A,Ono T,Park G H,Pfeil B,Poulter B,Raupach M R,Regnier P,R9denbeck C,Saito S,Schwinger J,Segschneider J,Stocker B,Tilbrook B,Van Heuven S M A C,Viovy N,Wanninkhof R,Wiltshire A,Zaehle S,Yue C.Global carbon budget 2013.Earth System Science Data,2014,6(7):235-263.
[4]Atarashi-Andoh M,Koarashi J,Ishizuka S,Hirai K.Seasonal patterns and control factors of CO2effluxes from surface litter,soil organic carbon,and root-derived carbon estimated using radiocarbon signatures.Agricultural and Forest Meteorology,2012,152:149-158.
[5]Cheng W X,Parton W J,Gonzalez-Meler MA,Phillips R,Asao S,McN ickle G G,Brzostek E,Jastrow J D.Synthesis and modeling perspectives of rhizosphere priming.New Phytologist,2014,201(1):31-44.
[6]Hopkins F,Gonzalez-Meler M A,Flower C E,Lynch D J,Czimczik C,Tang J W,Subke J A.Ecosystem-level controls on root-rhizosphere respiration.New Phytologist,2013,199(2):339-351.
[7]Chapin F S III,Matson P A,Vitousek P M.Principles of Terrestrial Ecosystem Ecology.2nd ed.New York:Springer Press,2012.
[8]Kuzyakov Y,Friedel J K,Stahr K.Review of mechanisms and quantification of priming effects.Soil Biology and Biochemistry,2000,32(11-12):1485-1498.
[9]Kuzyakov Y,Blagodatskaya E V.Microbial hotspots and hot moments in soil:Concept&review.Soil Biology and Biochemistry,2015,83:184-199.
[10]Kuzyakov Y.Priming effects:interactions between living and dead organic matter.Soil Biology and Biochemistry,2010,42(9):1363-1371.
[11]Blagodatskaya E V,Kuzyakov Y.Active microorganisms in soil:critical review of estimation criteria and approaches.Soil Biology and Biochemistry,2013,67:192-211.
[12]Kuzyakov Y.Sources of CO2efflux from soil and review of partitioning methods.Soil Biology and Biochemistry,2006,38(3):425-448.
[13]Werth M,Kuzyakov Y.Three-source partitioning of CO2efflux from maize field soil by13C natural abundance.Journal of Plant Nutrition and Soil Science,2009,172(4):487-499.
[14]Kuzyakov Y V,Larionova A A.Contribution of rhizomicrobial and root respiration to the CO2emission from soil(A Review).Eurasian Soil Science,2006,39(7):753-764.
[15]金钊,董云社,齐玉春.区分纯根呼吸和根际微生物呼吸的争议.土壤,2008,40(4):517-522.
[16]Hanson P J,Edwards N T,Garten C T,Andrews J A.Separating root and soil microbial contributions to soil respiration:A review of methods and observations.Biogeochemistry,2000,48(1):115-146.
[17]Anderson J M.The effects of climate change on decomposition processes in grassland and coniferous forests.Ecological Applications,1991,1(3):326-347.
[18]Li P,Yang Y H,Fang J Y.Variations of root and heterotrophic respiration along environmental gradients in Chinas forests.Journal of Plant Ecology,2013,6(5):358-367.
[19]H9gberg P,Buchmann N,Read D J.Comments on Yakov Kuzyakov's review‘Sources of CO2efflux from soil and review of partitioning methods':[Soil Biology&Biochemistry 38,425–448].Soil Biology and Biochemistry,2006,38(9):2997-2998.
[20]Kuzyakov Y,Larionova A A.Root and rhizomicrobial respiration:A review of approaches to estimate respiration by autotrophic and heterotrophic organisms in soil.Journal of Plant Nutrition and Soil Science,2005,168(4):503-520.
[21]孙悦,徐兴良,Kuzyakov Y.根际激发效应的发生机制及其生态重要性.植物生态学报,2014,38(1):62-75.
[22]Kuzyakov Y,Xu X L.Competition between roots and microorganisms for nitrogen:mechanisms and ecological relevance.New Phytologist,2013,198(3):656-669.
[23]Xu M,Shang H.Contribution of soil respiration to the global carbon equation.Journal of Plant Physiology,2016,203:16-28.
[24]Gerber S,Joos F,Prentice I C.Sensitivity of a dynamic global vegetation model to climate and atmospheric CO2.Global Change Biology,2004,10(8):1223-1239.
[25]Macdonald C A,Anderson I C,Bardgett R D,Singh B K.Role of nitrogen in carbon mitigation in forest ecosystems.Current Opinion in Environmental Sustainability,2011,3(5):303-310.
[26]Phillips R P,Finzi A C,Bernhardt E S.Enhanced root exudation induces microbial feedbacks to N cycling in a pine forest under long-term CO2fumigation.Ecology Letters,2011,14(2):187-194.
[27]Sayer E J,Heard M S,Grant H K,Marthews T R,Tanner E V J.Soil carbon release enhanced by increased tropical forest litter fall.Nature Climate Change,2011,1(6):304-307.
[28]Bengtson P,Barker J,Grayston S J.Evidence of a strong coupling between root exudation,C and N availability,and stimulated SOM decomposition caused by rhizosphere priming effects.Ecology and Evolution,2012,2(8):1843-1852.
[29]Yuan H Z,Ge T D,Chen C Y,O'Donnell A G,Wu J S.Significant role for microbial autotrophy in the sequestration of soil carbon.Applied and Environmental Microbiology,2012,78(7):2328-2336.
[30]Von Lützow M,K9gel-Knabner I.Temperature sensitivity of soil organic matter decomposition-What do we know?Biology and Fertility of Soils,2009,46(1):1-15.
[31]Conant R T,Ryan M G,gren G I,Birge H E,Davidson E A,Eliasson P E,Evans S E,Frey S D,Giardina C P,Hopkins F M,Hyv9nen R,Kirschbaum M U F,Lavallee J M,Leifeld J,Parton W J,Steinweg J M,Wallenstein M D,Wetterstedt JM,Bradford M A.Temperature and soil organic matter decomposition rates-synthesis of current knowledge and a way forward.Global Change Biology,2011,17(11):3392-3404.
[32]王兵,姜艳,郭浩,赵广东,白秀兰.土壤呼吸及其三个生物学过程研究.土壤通报,2011,42(2):483-490.
[33]陈敏鹏,夏旭,李银坤,梅旭荣.土壤呼吸组分分离技术研究进展.生态学报,2013,33(22):7067-7077.
[34]Remus R,Hüve K,P9rschmann J,Augustin J.Determining the timepoint when14C tracer accurately reflect photosynthate use in the plant-soil system.Plant and Soil,2016,408(1-2):457-474.
[35]Remus R,Augustin J.Dynamic linking of14C partitioning with shoot growth allows a precise determination of plant-derived C input to soil.Plant and Soil,2016,408(1-2):493-513.
[36]Carbone M S,Richardson A D,Chen M,Davidson E A,Hughes H,Savage K E,Hollinger D Y.Constrained partitioning of autotrophic and heterotrophic respiration reduces model uncertainties of forest ecosystem carbon fluxes but not stocks.Journal of Geophysical Research:Biogeosciences,2016,121(9):2476-2492.
[37]Kuzyakov Y.Separating microbial respiration of exudates from root respiration in non-sterile soils:A comparison of four methods.Soil Biology and Biochemistry,2002,34(11):1621-1631.
[38]Kelting D L,Burger J A,Edwards G S.Estimating root respiration,microbial respiration in the rhizosphere,and root-free soil respiration in forest soils.Soil Biology and Biochemistry,1998,30(7):961-968.
[39]Chen C R,Condron L M,Xu Z H,Davis M R,Sherlock R R.Root,rhizosphere and root-free respiration in soils under grassland and forest plants.European Journal of Soil Science,2006,57(1):58-66.
[40]Craine J M,Wedin D A,Chapin F S.Predominance of ecophysiological controls on soil CO2flux in a Minnesota grassland.Plant and Soil,1999,207(1):77-86.
[41]Larionova A A,Sapronov D V,de Gerenyu L,Kuznetsova L G,Kudeyarov V N.Contribution of plant root respiration to the CO2emission from soil.Eurasian Soil Science,2006,39(10):1127-1135.
[42]Barba J,Yuste J C,Poyatos R,Janssens I A,Lloret F.Strong resilience of soil respiration components to drought-induced die-off resulting in forest secondary succession.Oecologia,2016,182(1):27-41.
[43]Cheng W X,Zhang Q L,Coleman D C,Carroll C R,Hoffman C A.Is available carbon limiting microbial respiration in the rhizosphere?Soil Biology and Biochemistry,1996,28(10-11):1283-1288.
[44]Kuzyakov Y,Cheng W.Photosynthesis controls of rhizosphere respiration and organic matter decomposition.Soil Biology and Biochemistry,2001,33(14):1915-1925.
[45]Kuzyakov Y,Domanski G.Model for rhizodeposition and CO2efflux from planted soil and its validation by14C pulse labelling of ryegrass.Plant and Soil,2002,239(1):87-102.
[46]Sapronov D V,Kuzyakov Y V.Separation of root and microbial respiration:comparison of three methods.Eurasian Soil Science,2007,40(7):775-784.
[47]O'Leary M H.Carbon isotopes in photosynthesis.Bioscience,1988,38(5):328-336.
[48]Kuzyakov Y.Theoretical background for partitioning of root and rhizomicrobial respiration byδ13C of microbial biomass.European Journal of Soil Biology,2005,41(1-2):1-9.
[49]Werth M,Subbotina I,Kuzyakov Y.Three-source partitioning of CO2efflux from soil planted with maize by13C natural abundance fails due to inactive microbial biomass.Soil Biology and Biochemistry,2006,38(9):2772-2781.
[50]Blagodatskaya E,Khomyakov N,Myachina O,Bogomolova I,Blagodatsky S,Kuzyakov Y.Microbial interactions affect sources of priming induced by cellulose.Soil Biology and Biochemistry,2014,74:39-49.
[51]KerréB,Hernandez-Soriano M C,Smolders E.Partitioning of carbon sources among functional pools to investigate short-term priming effects of biochar in soil:A13C study.Science of Total Environment,2016,547:30-38.
[52]宋文琛,同小娟,张劲松,孟平,李俊.用自然13C丰度法区分人工林根源呼吸的原理与应用.中国水土保持科学,2015,13(4):37-43.
[53]Song W C,Tong X J,Zhang J S,Meng P.Three-source partitioning of soil respiration by13C natural abundance and its variation with soil depth in a plantation.Journal of Forestry Research,2016,27(3):533-540.
[54]TomèE,Ventura M,Folegot S,Zanotelli D,Montagnani L,Mimmo T,Tonon G,Tagliavini M,Scandellari F.Mycorrhizal contribution to soil respiration in an apple orchard.Applied Soil Ecology,2016,101:165-173.
[55]Werth M,Kuzyakov Y.13C fractionation at the root-microorganisms-soil interface:A review and outlook for partitioning studies.Soil Biology and Biochemistry,2010,42(9):1372-1384.
[56]Tian J,Pausch J,Yu G R,Blagodatskaya E,Kuzyakov Y.Aggregate size and glucose level affect priming sources:A three-source-partitioning study.Soil Biology and Biochemistry,2016,97:199-210.
[57]Dijkstra F A,Carrillo Y,Pendall E,Morgan J A.Rhizosphere priming:a nutrient perspective.Frontiers in Microbiology,2013,4(1):65-72.
[58]Sullivan B W,Hart S C.Evaluation of mechanisms controlling the priming of soil carbon along a substrate age gradient.Soil Biology and Biochemistry,2013,58:293-301.
[59]Burns R G,DeF orest J L,Marxsen J,Sinsabaugh R L,Stromberger M E,Wallenstein M D,Weintraub M N,Zoppini A.Soil enzymes in a changing environment:current knowledge and future directions.Soil Biology and Biochemistry,2013,58:216-234.
[60]Spohn M,P9tsch E M,Eichorst S A,Woebken D,Wanek W,Richter A.Soil microbial carbon use efficiency and biomass turnover in a long-term fertilization experiment in a temperate grassland.Soil Biology and Biochemistry,2016,97:168-175.
[61]Zang H D,Wang J Y,Kuzyakov Y.N fertilization decreases soil organic matter decomposition in the rhizosphere.Applied Soil Ecology,2016,108:47-53.
[62]耿元波,史晶晶.碳同位素在草地土壤呼吸区分中的应用.中国农业科学,2012,45(17):3541-3550.
[63]魏书精,罗碧珍,魏书威,孙龙,文正敏,胡海清.森林生态系统土壤呼吸测定方法研究进展.生态环境学报,2014,23(3):504-514.
[64]Qiao N,Schaefer D,Blagodatskaya E,Zou X M,Xu X L,Kuzyakov Y.Labile carbon retention compensates for CO2released by priming in forest soils.Global Change Biology,2014,20(6):1943-1954.
[65]Pan Y D,Birdsey R A,Fang J Y,Houghton R,Kauppi P E,Kurz W A,Phillips O L,Shvidenko A,Lewis S L,Canadell J G,Ciais P,Jackson P B,Pacala S W,McG uire A D,Piao S L,Rautiainen A,Sitch S,Hayes D.A large and persistent carbon sink in the world's forests.Science,2011,333(6045):988-993.