Altered trends in carbon uptake in China's terrestrial ecosystems under the enhanced summer monsoon and warming hiatus
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摘要
The carbon budgets in terrestrial ecosystems in China are strongly coupled with climate changes. Over the past decade, China has experienced dramatic climate changes characterized by enhanced summer monsoon and decelerated warming. However, the changes in the trends of terrestrial net ecosystem production(NEP) in China under climate changes are not well documented. Here, we used three ecosystem models to simulate the spatiotemporal variations in China's NEP during 1982–2010 and quantify the contribution of the strengthened summer monsoon and warming hiatus to the NEP variations in four distinct climatic regions of the country. Our results revealed a decadal-scale shift in NEP from a downtrend of–5.95 TgC/yr~2(reduced sink) during 1982–2000 to an uptrend of 14.22 TgC/yr~2(enhanced sink) during 2000–10. This shift was essentially induced by the strengthened summer monsoon, which stimulated carbon uptake, and the warming hiatus, which lessened the decrease in the NEP trend. Compared to the contribution of 56.3% by the climate effect, atmospheric CO_2 concentration and nitrogen deposition had relatively small contributions(8.6 and 11.3%, respectively) to the shift. In conclusion, within the context of the global-warming hiatus, the strengthening of the summer monsoon is a critical climate factor that enhances carbon uptake in China due to the asymmetric response of photosynthesis and respiration. Our study not only revealed the shift in ecosystem carbon sequestration in China in recent decades, but also provides some insight for understanding ecosystem carbon dynamics in other monsoonal areas.
The carbon budgets in terrestrial ecosystems in China are strongly coupled with climate changes. Over the past decade, China has experienced dramatic climate changes characterized by enhanced summer monsoon and decelerated warming. However, the changes in the trends of terrestrial net ecosystem production(NEP) in China under climate changes are not well documented. Here, we used three ecosystem models to simulate the spatiotemporal variations in China's NEP during 1982–2010 and quantify the contribution of the strengthened summer monsoon and warming hiatus to the NEP variations in four distinct climatic regions of the country. Our results revealed a decadal-scale shift in NEP from a downtrend of–5.95 TgC/yr~2(reduced sink) during 1982–2000 to an uptrend of 14.22 TgC/yr~2(enhanced sink) during 2000–10. This shift was essentially induced by the strengthened summer monsoon, which stimulated carbon uptake, and the warming hiatus, which lessened the decrease in the NEP trend. Compared to the contribution of 56.3% by the climate effect, atmospheric CO_2 concentration and nitrogen deposition had relatively small contributions(8.6 and 11.3%, respectively) to the shift. In conclusion, within the context of the global-warming hiatus, the strengthening of the summer monsoon is a critical climate factor that enhances carbon uptake in China due to the asymmetric response of photosynthesis and respiration. Our study not only revealed the shift in ecosystem carbon sequestration in China in recent decades, but also provides some insight for understanding ecosystem carbon dynamics in other monsoonal areas.
The carbon budgets in terrestrial ecosystems in China are strongly coupled with climate changes. Over the past decade, China has experienced dramatic climate changes characterized by enhanced summer monsoon and decelerated warming. However, the changes in the trends of terrestrial net ecosystem production(NEP) in China under climate changes are not well documented. Here, we used three ecosystem models to simulate the spatiotemporal variations in China's NEP during 1982–2010 and quantify the contribution of the strengthened summer monsoon and warming hiatus to the NEP variations in four distinct climatic regions of the country. Our results revealed a decadal-scale shift in NEP from a downtrend of–5.95 TgC/yr~2(reduced sink) during 1982–2000 to an uptrend of 14.22 TgC/yr~2(enhanced sink) during 2000–10. This shift was essentially induced by the strengthened summer monsoon, which stimulated carbon uptake, and the warming hiatus, which lessened the decrease in the NEP trend. Compared to the contribution of 56.3% by the climate effect, atmospheric CO_2 concentration and nitrogen deposition had relatively small contributions(8.6 and 11.3%, respectively) to the shift. In conclusion, within the context of the global-warming hiatus, the strengthening of the summer monsoon is a critical climate factor that enhances carbon uptake in China due to the asymmetric response of photosynthesis and respiration. Our study not only revealed the shift in ecosystem carbon sequestration in China in recent decades, but also provides some insight for understanding ecosystem carbon dynamics in other monsoonal areas.
引文
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2.Fang JY,Guo ZD and Piao SL et al.Terrestrial vegetation carbon sinks in China,1981-2000.Sci China Ser D 2007;50:1341-50.
3.Piao SL,Fang JY and Ciais P et al.The carbon balance of terrestrial ecosystems in China.Nature 2009;458:1009-13.
4.Cao MK,Prince SD and Li KR et al.Response of terrestrial carbon uptake to climate interannual variability in China.Glob Change Biol 2003;9:536-46.
5.Yu GR,Zhu XJ and Fu YL et al.Spatial patterns and climate drivers of carbon fluxes in terrestrial ecosystems of China.Glob Change Biol 2013;19:798-810.
6.Piao SL,Ciais P and Huang Y et al.The impacts of climate change on water resources and agriculture in China.Nature2010;467:43-51.
7.Ding YH,Liu YJ and Liang SJ et al.Interdecadal variability of the east asian winter monsoon and its possible links to global climate change.J Meteorol Res 2014;28:693-713.
8.IPCC:Summary for policymakers.In:Stocker TF,Qin D and Plattner GK et al.(eds).Climate Change 2013:The Physical Science Basis.Contribution of Working Group I to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change.Cambridge and New York:Cambridge University Press,2013,3-29.
9.Fyfe JC,Meehl GA and England MH et al.Making sense of the early-2000s warming slowdown.Nat Clim Change 2016;6:224-8.
10.Mu Q,Zhao M and Running SW et al.Contribution of increasing CO2and climate change to the carbon cycle in China’s ecosystems.J Geophys Res 2008;113:G01018.
11.Keenan TF,Prentice IC and Canadell JG et al.Recent pause in the growth rate of atmospheric CO2due to enhanced terrestrial carbon uptake.Nat Commun 2016;7:13428.
12.Sitch S,Friedlingstein P and Gruber N et al.Recent trends and drivers of regional sources and sinks of carbon dioxide.Biogeosciences 2015;12:653-79.
13.Ballantyne A,Smith W and Anderegg W et al.Accelerating net terrestrial carbon uptake during the warming hiatus due to reduced respiration.Nat Clim Change 2017;7:148-52.
14.Tian H,Melillo J and Lu C et al.China’s terrestrial carbon balance:contributions from multiple global change factors.Global Biogeochem Cycles 2011;25:GB1007.
15.Tao FL and Zhang Z.Dynamic responses of terrestrial ecosystems structure and function to climate change in China.J Geophys Res 2010;115:G03003.
16.Fang JY,Yu GR and Liu LL et al.Climate change,human impacts,and carbon sequestration in China.Proc Natl Acad Sci USA 2018;115:4015-20.
17.Yu GR,Ren W and Chen Z et al.Construction and progress of Chinese terrestrial ecosystem carbon,nitrogen and water fluxes coordinated observation.J Geogr Sci 2016;26:803-26.
18.Tang XL,Zhao X and Bai YF et al.Carbon pools in China’s terrestrial ecosystems:new estimates based on an intensive field survey.Proc Natl Acad Sci USA 2018;115:4021-6.
19.Editorial Committee.The 3rd National Assessment Report of Climatic Changes(in Chinese).Beijing:Science Press,2015.
20.Yu GR,Chen Z and Piao SL et al.High carbon dioxide uptake by subtropical forest ecosystems in the East Asian monsoon region.Proc Natl Acad Sci USA 2014;111:4910-5.
21.Li P,Zhu J and Hu H et al.The relative contributions of forest growth and areal expansion to forest biomass carbon.Biogeosciences 2016;13:375-88.
22.Lu F,Hu HF and Sun WJ et al.Effects of national ecological restoration projects on carbon sequestration in China from 2001to 2010.Proc Natl Acad Sci USA 2018;115:4039-44.
23.Le Quere C,Andrew RM and Canadell JG et al.Global carbon budget 2016.Earth Syst Sci Data 2016;8:605-49.
24.Schimel D,Melillo J and Tian HQ et al.Contribution of increasing CO2 and climate to carbon storage by ecosystems in the United States.Science 2000;287:2004-6.
25.Hayes DJ,Turner DP and Stinson G et al.Reconciling estimates of the contemporary North American carbon balance among terrestrial biosphere models,atmospheric inversions,and a new approach for estimating net ecosystem exchange from inventory-based data.Glob Change Biol 2012;18:1282-99.
26.Schulze ED,Luyssaert S and Ciais P et al.Importance of methane and nitrous oxide for Europe’s terrestrial greenhouse-gas balance.Nat Geosci 2009;2:842-50.
27.Janssens IA,Freibauer A and Ciais P et al.Europe’s terrestrial biosphere absorbs 7 to 12%of European anthropogenic CO2emissions.Science 2003;300:1538-42.
28.Piao SL,Yin GD and Tan JG et al.Detection and attribution of vegetation greening trend in China over the last 30 years.Glob Change Biol 2015;21:1601-9.
29.Jung M,Reichstein M and Margolis HA et al.Global patterns of landatmosphere fluxes of carbon dioxide,latent heat,and sensible heat derived from eddy covariance,satellite,and meteorological observations.J Geophys Res 2011;116:G00J7.
30.Chevallier F,Ciais P and Conway T et al.CO2surface fluxes at grid point scale estimated from a global 21 year reanalysis of atmospheric measurements.JGeophys Res 2010;115:D21307.
31.Zhang H,Chen B and vd I Laan-Luijkx et al.Net terrestrial CO2exchange over China during 2001-2010 estimated with an ensemble data assimilation system for atmospheric CO2.J Geophys Res:Atmospheres 2014;119:3500-15.
32.Liu HW,Zhou TJ and Zhu YX et al.The strengthening East Asia summer monsoon since the early 1990s.Chin Sci Bull 2012;57:1553-8.
33.Zhang RH.Changes in East Asian summer monsoon and summer rainfall over eastern China during recent decades.Sci Bull 2015;60:1222-4.
34.Zhou TJ,Gong DY and Li J et al.Detecting and understanding the multi-decadal variability of the East Asian Summer Monsoon:recent progress and state of affairs.metz 2009;18:455-67.
35.Ahlstrom A,Raupach MR and Schurgers G et al.The dominant role of semi-arid ecosystems in the trend and variability of the land CO2sink.Science 2015;348:895-9.
36.Poulter B,Frank D and Ciais P et al.Contribution of semi-arid ecosystems to interannual variability of the global carbon cycle.Nature 2014;509:600-3.
37.Tan Z-H,Zhang Y-P and Liang N et al.An observational study of the carbon-sink strength of East Asian subtropical evergreen forests.Environ Res Lett 2012;7:044017.
38.Wang WL,Ciais P and Nemani RR et al.Variations in atmospheric CO2growth rates coupled with tropical temperature.Proc Nat Acad Sci USA 2013;110:13061-6.
39.Liu XJ,Zhang Y and Han WX et al.Enhanced nitrogen deposition over China.Nature 2013;494:459-62.
40.Gu FX,Zhang YD and Huang M et al.Nitrogen deposition and its effect on carbon storage in Chinese forests during 1981-2010.Atmos Environ 2015;123:171-9.
41.Schimel D,Stephens BB and Fisher JB.Effect of increasing CO2on the terrestrial carbon cycle.Proc Natl Acad Sci USA 2015;112:436-41.
42.Fang JY,Chen AP and Peng CH et al.Changes in forest biomass carbon storage in China between 1949 and 1998.Science 2001;292:2320-2.
43.Houghton RA and Hackler JL.Sources and sinks of carbon from land-use change in China.Global Biogeochem Cycles 2003;17.
44.Liu ML and Tian HQ.China’s land cover and land use change from 1700 to 2005:estimations from high-resolution satellite data and historical archives.Global Biogeochem Cycles 2010;24:GB3003.
45.Wang SQ,Zhou L and Chen JM et al.Relationships between net primary productivity and stand age for several forest types and their influence on China’s carbon balance.J Environ Manage 2011;92:1651-62.
46.Bao Q.Projected changes in Asian Summer Monsoon in RCP scenarios of CMIP5.Atmos Oceanic Sci Lett 2012;05:43-8.
47.Chen HP and Sun JQ.Projected change in East Asian summer monsoon precipitation under RCP scenario.Meteorol Atmos Phys 2013;121:55-77.
48.Christensen JH,Kanikicharla KK and Marshall G et al.Climate phenomena and their relevance for future regional climate change.In:Stocker TF,Qin D and Plattner GK et al.(eds).Climate Change 2013:The Physical Science Basis.Contribution of Working Group I to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change.Cambridge and New York:Cambridge University Press,2013,1217-308.
49.Roberts CD,Palmer MD and Mc Neall D et al.Quantifying the likelihood of a continued hiatus in global warming.Nat Clim Change 2015;5:337-42.
50.Knutson TR,Zhang R and Horowitz LW.Prospects for a prolonged slowdown in global warming in the early 21st century.Nat Commun 2016;7:13676.
51.Liu Y,Zhou Y and Ju W et al.Impacts of droughts on carbon sequestration by China’s terrestrial ecosystems from 2000 to 2011.Biogeosciences 2014;11:2583-99.
52.Yan H,Wang SQ and Billesbach D et al.Global estimation of evapotranspiration using a leaf area index-based surface energy and water balance model.Remote Sens Environ 2012;124:581-95.
53.Wang Q,Tenhunen J and Falge E et al.Simulation and scaling of temporal variation in gross primary production for coniferous and deciduous temperate forests.Glob Change Biol 2004;10:37-51.
54.Wang J,Wang J and Ye H et al.An interpolated temperature and precipitation dataset at 1-km grid resolution in China(2000-2012).China Scientific Data2017;2:88-95.
55.Wu BF(ed).Land Cover Atlas of the People’s Republic of China(1:1 000 000)(in Chinese).Beijing:China Map Publishing House,2017
56.Zhu Z,Bi J and Pan Y et al.Global data sets of vegetation LAI3g and FPAR3g derived from GIMMS NDVI3g for the period 1981 to 2011.Remote Sens 2013;5:927-48.
57.Liu Y,Liu R and Chen JM.Retrospective retrieval of long-term consistent global leaf area index(1981-2011)from combined AVHRR and MODIS data.J Geophys Res 2012;117:G04003.
58.Piao SL,Sitch S and Ciais P et al.Evaluation of terrestrial carbon cycle models for their response to climate variability and to CO2 trends.Glob Change Biol2013;19:2117-32.
2.Fang JY,Guo ZD and Piao SL et al.Terrestrial vegetation carbon sinks in China,1981-2000.Sci China Ser D 2007;50:1341-50.
3.Piao SL,Fang JY and Ciais P et al.The carbon balance of terrestrial ecosystems in China.Nature 2009;458:1009-13.
4.Cao MK,Prince SD and Li KR et al.Response of terrestrial carbon uptake to climate interannual variability in China.Glob Change Biol 2003;9:536-46.
5.Yu GR,Zhu XJ and Fu YL et al.Spatial patterns and climate drivers of carbon fluxes in terrestrial ecosystems of China.Glob Change Biol 2013;19:798-810.
6.Piao SL,Ciais P and Huang Y et al.The impacts of climate change on water resources and agriculture in China.Nature2010;467:43-51.
7.Ding YH,Liu YJ and Liang SJ et al.Interdecadal variability of the east asian winter monsoon and its possible links to global climate change.J Meteorol Res 2014;28:693-713.
8.IPCC:Summary for policymakers.In:Stocker TF,Qin D and Plattner GK et al.(eds).Climate Change 2013:The Physical Science Basis.Contribution of Working Group I to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change.Cambridge and New York:Cambridge University Press,2013,3-29.
9.Fyfe JC,Meehl GA and England MH et al.Making sense of the early-2000s warming slowdown.Nat Clim Change 2016;6:224-8.
10.Mu Q,Zhao M and Running SW et al.Contribution of increasing CO2and climate change to the carbon cycle in China’s ecosystems.J Geophys Res 2008;113:G01018.
11.Keenan TF,Prentice IC and Canadell JG et al.Recent pause in the growth rate of atmospheric CO2due to enhanced terrestrial carbon uptake.Nat Commun 2016;7:13428.
12.Sitch S,Friedlingstein P and Gruber N et al.Recent trends and drivers of regional sources and sinks of carbon dioxide.Biogeosciences 2015;12:653-79.
13.Ballantyne A,Smith W and Anderegg W et al.Accelerating net terrestrial carbon uptake during the warming hiatus due to reduced respiration.Nat Clim Change 2017;7:148-52.
14.Tian H,Melillo J and Lu C et al.China’s terrestrial carbon balance:contributions from multiple global change factors.Global Biogeochem Cycles 2011;25:GB1007.
15.Tao FL and Zhang Z.Dynamic responses of terrestrial ecosystems structure and function to climate change in China.J Geophys Res 2010;115:G03003.
16.Fang JY,Yu GR and Liu LL et al.Climate change,human impacts,and carbon sequestration in China.Proc Natl Acad Sci USA 2018;115:4015-20.
17.Yu GR,Ren W and Chen Z et al.Construction and progress of Chinese terrestrial ecosystem carbon,nitrogen and water fluxes coordinated observation.J Geogr Sci 2016;26:803-26.
18.Tang XL,Zhao X and Bai YF et al.Carbon pools in China’s terrestrial ecosystems:new estimates based on an intensive field survey.Proc Natl Acad Sci USA 2018;115:4021-6.
19.Editorial Committee.The 3rd National Assessment Report of Climatic Changes(in Chinese).Beijing:Science Press,2015.
20.Yu GR,Chen Z and Piao SL et al.High carbon dioxide uptake by subtropical forest ecosystems in the East Asian monsoon region.Proc Natl Acad Sci USA 2014;111:4910-5.
21.Li P,Zhu J and Hu H et al.The relative contributions of forest growth and areal expansion to forest biomass carbon.Biogeosciences 2016;13:375-88.
22.Lu F,Hu HF and Sun WJ et al.Effects of national ecological restoration projects on carbon sequestration in China from 2001to 2010.Proc Natl Acad Sci USA 2018;115:4039-44.
23.Le Quere C,Andrew RM and Canadell JG et al.Global carbon budget 2016.Earth Syst Sci Data 2016;8:605-49.
24.Schimel D,Melillo J and Tian HQ et al.Contribution of increasing CO2 and climate to carbon storage by ecosystems in the United States.Science 2000;287:2004-6.
25.Hayes DJ,Turner DP and Stinson G et al.Reconciling estimates of the contemporary North American carbon balance among terrestrial biosphere models,atmospheric inversions,and a new approach for estimating net ecosystem exchange from inventory-based data.Glob Change Biol 2012;18:1282-99.
26.Schulze ED,Luyssaert S and Ciais P et al.Importance of methane and nitrous oxide for Europe’s terrestrial greenhouse-gas balance.Nat Geosci 2009;2:842-50.
27.Janssens IA,Freibauer A and Ciais P et al.Europe’s terrestrial biosphere absorbs 7 to 12%of European anthropogenic CO2emissions.Science 2003;300:1538-42.
28.Piao SL,Yin GD and Tan JG et al.Detection and attribution of vegetation greening trend in China over the last 30 years.Glob Change Biol 2015;21:1601-9.
29.Jung M,Reichstein M and Margolis HA et al.Global patterns of landatmosphere fluxes of carbon dioxide,latent heat,and sensible heat derived from eddy covariance,satellite,and meteorological observations.J Geophys Res 2011;116:G00J7.
30.Chevallier F,Ciais P and Conway T et al.CO2surface fluxes at grid point scale estimated from a global 21 year reanalysis of atmospheric measurements.JGeophys Res 2010;115:D21307.
31.Zhang H,Chen B and vd I Laan-Luijkx et al.Net terrestrial CO2exchange over China during 2001-2010 estimated with an ensemble data assimilation system for atmospheric CO2.J Geophys Res:Atmospheres 2014;119:3500-15.
32.Liu HW,Zhou TJ and Zhu YX et al.The strengthening East Asia summer monsoon since the early 1990s.Chin Sci Bull 2012;57:1553-8.
33.Zhang RH.Changes in East Asian summer monsoon and summer rainfall over eastern China during recent decades.Sci Bull 2015;60:1222-4.
34.Zhou TJ,Gong DY and Li J et al.Detecting and understanding the multi-decadal variability of the East Asian Summer Monsoon:recent progress and state of affairs.metz 2009;18:455-67.
35.Ahlstrom A,Raupach MR and Schurgers G et al.The dominant role of semi-arid ecosystems in the trend and variability of the land CO2sink.Science 2015;348:895-9.
36.Poulter B,Frank D and Ciais P et al.Contribution of semi-arid ecosystems to interannual variability of the global carbon cycle.Nature 2014;509:600-3.
37.Tan Z-H,Zhang Y-P and Liang N et al.An observational study of the carbon-sink strength of East Asian subtropical evergreen forests.Environ Res Lett 2012;7:044017.
38.Wang WL,Ciais P and Nemani RR et al.Variations in atmospheric CO2growth rates coupled with tropical temperature.Proc Nat Acad Sci USA 2013;110:13061-6.
39.Liu XJ,Zhang Y and Han WX et al.Enhanced nitrogen deposition over China.Nature 2013;494:459-62.
40.Gu FX,Zhang YD and Huang M et al.Nitrogen deposition and its effect on carbon storage in Chinese forests during 1981-2010.Atmos Environ 2015;123:171-9.
41.Schimel D,Stephens BB and Fisher JB.Effect of increasing CO2on the terrestrial carbon cycle.Proc Natl Acad Sci USA 2015;112:436-41.
42.Fang JY,Chen AP and Peng CH et al.Changes in forest biomass carbon storage in China between 1949 and 1998.Science 2001;292:2320-2.
43.Houghton RA and Hackler JL.Sources and sinks of carbon from land-use change in China.Global Biogeochem Cycles 2003;17.
44.Liu ML and Tian HQ.China’s land cover and land use change from 1700 to 2005:estimations from high-resolution satellite data and historical archives.Global Biogeochem Cycles 2010;24:GB3003.
45.Wang SQ,Zhou L and Chen JM et al.Relationships between net primary productivity and stand age for several forest types and their influence on China’s carbon balance.J Environ Manage 2011;92:1651-62.
46.Bao Q.Projected changes in Asian Summer Monsoon in RCP scenarios of CMIP5.Atmos Oceanic Sci Lett 2012;05:43-8.
47.Chen HP and Sun JQ.Projected change in East Asian summer monsoon precipitation under RCP scenario.Meteorol Atmos Phys 2013;121:55-77.
48.Christensen JH,Kanikicharla KK and Marshall G et al.Climate phenomena and their relevance for future regional climate change.In:Stocker TF,Qin D and Plattner GK et al.(eds).Climate Change 2013:The Physical Science Basis.Contribution of Working Group I to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change.Cambridge and New York:Cambridge University Press,2013,1217-308.
49.Roberts CD,Palmer MD and Mc Neall D et al.Quantifying the likelihood of a continued hiatus in global warming.Nat Clim Change 2015;5:337-42.
50.Knutson TR,Zhang R and Horowitz LW.Prospects for a prolonged slowdown in global warming in the early 21st century.Nat Commun 2016;7:13676.
51.Liu Y,Zhou Y and Ju W et al.Impacts of droughts on carbon sequestration by China’s terrestrial ecosystems from 2000 to 2011.Biogeosciences 2014;11:2583-99.
52.Yan H,Wang SQ and Billesbach D et al.Global estimation of evapotranspiration using a leaf area index-based surface energy and water balance model.Remote Sens Environ 2012;124:581-95.
53.Wang Q,Tenhunen J and Falge E et al.Simulation and scaling of temporal variation in gross primary production for coniferous and deciduous temperate forests.Glob Change Biol 2004;10:37-51.
54.Wang J,Wang J and Ye H et al.An interpolated temperature and precipitation dataset at 1-km grid resolution in China(2000-2012).China Scientific Data2017;2:88-95.
55.Wu BF(ed).Land Cover Atlas of the People’s Republic of China(1:1 000 000)(in Chinese).Beijing:China Map Publishing House,2017
56.Zhu Z,Bi J and Pan Y et al.Global data sets of vegetation LAI3g and FPAR3g derived from GIMMS NDVI3g for the period 1981 to 2011.Remote Sens 2013;5:927-48.
57.Liu Y,Liu R and Chen JM.Retrospective retrieval of long-term consistent global leaf area index(1981-2011)from combined AVHRR and MODIS data.J Geophys Res 2012;117:G04003.
58.Piao SL,Sitch S and Ciais P et al.Evaluation of terrestrial carbon cycle models for their response to climate variability and to CO2 trends.Glob Change Biol2013;19:2117-32.
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