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Pan et al 2011

propagation are directly accessible to anyone with basic statistical knowledge.This should ul-timately open the way for a complete character-ization of the roles of direct and indirect top-down and bottom-up mechanisms involved in the reg-ulation of parasite densities(fig.S12and table S1) in the context of both single and mixed infections, and how this in turn affects transmission and disease severity.

The underlying process of bursting infected RBCs and invasion of uninfected RBCs is com-mon to blood-phase malaria across animal taxa. The methods we introduce will consequently be generally applicable.The strength of the mouse data we have used is the finely resolved measures of uninfected and infected red blood cells.We are unaware of any experimental time series in hu-man patients in which these parameters were directly measured,but our analyses suggest that future longitudinal studies of individual patients that undertake the simple assays required to di-rectly assess RBC densities in addition to parasite densities will lead to considerable insights into the factors regulating human malaria.

References and Notes

1.R.Carter,D.Walliker,Ann.Trop.Med.Parasitol.69,

187(1975).

2.L.Molineaux,M.Tr?uble,W.E.Collins,G.M.Jeffery,

K.Dietz,Trans.R.Soc.Trop.Med.Hyg.96,205(2002).

3.K.Dietz,G.Raddatz,L.Molineaux,Am.J.Trop.Med.Hyg.

75(suppl.),46(2006).

4.D.T.Haydon,L.Matthews,R.Timms,N.Colegrave,

Proc.R.Soc.B270,289(2003).

5.R.M.Ribeiro et al.,J.Virol.84,6096(2010).

6.O.N.Bj?rnstad,B.Finkenst?dt,B.T.Grenfell,

Ecol.Monogr.72,169(2002).

7.R.M.Anderson,R.M.May,Infectious Diseases of

Humans(Oxford Univ.Press,Oxford,1991).

8.M.A.Nowak,R.M.May,Virus Dynamics:Mathematical

Principles of Immunology and Virology(Oxford Univ.

Press,Oxford,2000).

9.M.M.Stevenson,E.M.Riley,Nat.Rev.Immunol.4,

169(2004).

10.M.Walther et al.,J.Immunol.177,5736(2006).

https://www.wendangku.net/doc/9312873606.html,ler,L.R?berg,A.F.Read,N.J.Savill,

PLOS Comput.Biol.6,e1000946(2010).

12.N.Mideo et al.,Am.Nat.172,E214(2008).

13.R.Antia,A.Yates,J.C.de Roode,Proc.R.Soc.B275,

1449(2008).

14.B.F.Kochin,A.J.Yates,J.C.de Roode,R.Antia,

PLoS ONE5,e10444(2010).

15.A.Handel,I.M.Longini Jr.,R.Antia,J.R.Soc.Interface

7,35(2010).

16.C.L.Ball,M.A.Gilchrist,D.Coombs,Bull.Math.Biol.

69,2361(2007).

17.A.S.Perelson,Nat.Rev.Immunol.2,28(2002).

18.R.A.Saenz et al.,J.Virol.84,3974(2010).

19.K.A.Lythgoe,L.J.Morrison,A.F.Read,J.D.Barry,

Proc.Natl.Acad.Sci.U.S.A.104,8095(2007).

20.L.Molineaux,K.Dietz,Parassitologia41,221

(1999).

21.R.Killick-Kendrick,W.Peters,Eds.,Rodent Malaria

(Academic Press,London,1978).

22.V.C.Barclay et al.,Proc.R.Soc.B275,1171

(2008).

23.S.Huijben,thesis,University of Edinburgh(2010).

24.G.H.Long,B.H.K.Chan,J.E.Allen,A.F.Read,

A.L.Graham,BMC Evol.Biol.8,128(2008).

25.See supporting material on Science Online.

26.P.G.McQueen,F.E.McKenzie,Proc.Natl.Acad.

Sci.U.S.A.101,9161(2004).

27.B.Hellriegel,Proc.R.Soc.B250,249(1992).

28.C.Hetzel,R.M.Anderson,Parasitology113,25

(1996).

29.W.Jarra,K.N.Brown,Parasitology99,157(1989).

https://www.wendangku.net/doc/9312873606.html,mikanra et al.,Blood110,18(2007).

31.S.S.Pilyugin,R.Antia,Bull.Math.Biol.62,869

(2000).

32.R.Antia,J.C.Koella,J.Theor.Biol.168,141(1994).

33.J.R.Glynn,D.J.Bradley,Parasitology110,7(1995).

34.M.S.Russell et al.,J.Immunol.179,211(2007).

35.D.L.Chao,M.P.Davenport,S.Forrest,A.S.Perelson,

Immunol.Cell Biol.82,55(2004).

36.R.Stephens,https://www.wendangku.net/doc/9312873606.html,nghorne,PLoS Pathog.6,e1001208

(2010).

37.C.Othoro et al.,J.Infect.Dis.179,279(1999).

38.F.P.Mockenhaupt et al.,Blood104,2003(2004).

39.S.Wambua,J.Mwacharo,S.Uyoga,A.Macharia,

T.N.Williams,Br.J.Haematol.133,206(2006).

40.K.Baer,C.Klotz,S.H.Kappe,T.Schnieder,U.Frevert,

PLoS Pathog.3,e171(2007).

41.N.J.Savill,W.Chadwick,S.E.Reece,PLOS Comput.Biol.

5,e1000416(2009).

42.Z.Su,A.Fortin,P.Gros,M.M.Stevenson,J.Infect.Dis.

186,1321(2002).

43.G.H.Long,B.H.K.Chan,J.E.Allen,A.F.Read,

A.L.Graham,Parasitology133,673(2006).

44.K.-H.Chang,M.Tam,M.M.Stevenson,J.Infect.Dis.

189,735(2004).

Acknowledgments:Our empirical work was funded by the

Wellcome Trust(A.F.R.,V.C.B.,G.H.L.),the Darwin

Trust of the University of Edinburgh(S.H.),and the UK

Biotechnology and Biological Sciences Research Council

(A.L.G.,G.H.L.),and the theoretical work by the Bill

and Melinda Gates Foundation(C.J.E.M.,B.T.G.,O.N.B.),

the RAPIDD program of the Science and Technology

Directorate(B.T.G.,A.L.G.,A.F.R.),and National Institute

of General Medical Sciences grant R01GM089932

(B.G.,O.N.B.,A.F.R.).We thank N.Mideo and P.Klepac

for extensive discussion.All authors discussed the

results and implications and commented on the

manuscript at all stages.C.J.E.M.and O.N.B.developed

the statistical approach;A.F.R.,V.B.,and S.H.designed

and performed the dose-dependent and CD4+T cell–

depleted mice experiments;A.L.G.and G.H.L.designed

and performed the innate immunity experiments.

The authors declare no competing interests.

Supporting Online Material

https://www.wendangku.net/doc/9312873606.html,/cgi/content/full/333/6045/984/DC1

Materials and Methods

SOM Text

Figs.S1to S12

Table S1

References

21February2011;accepted22June2011

10.1126/science.1204588

A Large and Persistent Carbon Sink in the World’s Forests

Yude Pan,1*Richard A.Birdsey,1Jingyun Fang,2,3Richard Houghton,4Pekka E.Kauppi,5 Werner A.Kurz,6Oliver L.Phillips,7Anatoly Shvidenko,8Simon L.Lewis,7Josep G.Canadell,9 Philippe Ciais,10Robert B.Jackson,11Stephen W.Pacala,12A.David McGuire,13Shilong Piao,2 Aapo Rautiainen,5Stephen Sitch,7Daniel Hayes14

The terrestrial carbon sink has been large in recent decades,but its size and location remain https://www.wendangku.net/doc/9312873606.html,ing forest inventory data and long-term ecosystem carbon studies,we estimate a total forest sink of2.4T0.4petagrams of carbon per year(Pg C year–1)globally for1990to2007. We also estimate a source of1.3T0.7Pg C year–1from tropical land-use change,consisting of a gross tropical deforestation emission of2.9T0.5Pg C year–1partially compensated by a carbon sink in tropical forest regrowth of1.6T0.5Pg C year–1.Together,the fluxes comprise a net global forest sink of1.1T0.8Pg C year–1,with tropical estimates having the largest uncertainties.Our total forest sink estimate is equivalent in magnitude to the terrestrial sink deduced from fossil fuel emissions and land-use change sources minus ocean and atmospheric sinks.

F orests have an important role in the global

carbon cycle and are valued globally for the

services they provide to society.International negotiations to limit greenhouse gases require an understanding of the current and potential future role of forest C emissions and sequestra-tion in both managed and unmanaged forests.

Estimates by the Intergovernmental Panel on Cli-

mate Change(IPCC)show that the net uptake by

terrestrial ecosystems ranges from less than1.0

to as much as2.6Pg C year–1for the1990s(1).

More recent global C analyses have estimated a

terrestrial C sink in the range of2.0to3.4Pg C

year–1on the basis of atmospheric CO2obser-

vations and inverse modeling,as well as land

observations(2–4).Because of this uncertainty

and the possible change in magnitude over time,

constraining these estimates is critically impor-

tant to support future climate mitigation actions.

1U.S.Department of Agriculture Forest Service,Newtown

Square,PA19073,USA.2Key Laboratory for Earth Surface Pro-

cesses,Ministry of Education,Peking University,Beijing,100871

China.3State Key Laboratory of Vegetation and Environmental

Change,Institute of Botany,Chinese Academy of Sciences,

Beijing,100093China.4Woods Hole Research Center,Falmouth,

MA02543,USA.5University of Helsinki,Helsinki,Finland.6Natural

Resources Canada,Canadian Forest Service,Victoria,BC,V8Z

1M5,Canada.7School of Geography,University of Leeds,LS2

9JT,UK.8International Institute for Applied Systems Analysis,

Laxenburg,Austria.9Global Carbon Project,Commonwealth Sci-

entific and Industrial Research Organization Marine and Atmo-

spheric Research,Canberra,Australia.10Laboratoire des Sciences

du Climat et de l’Environnement CEA-UVSQ-CNRS,Gif sur Yvette,

France.11Duke University,Durham,NC27708,USA.12Prince-

ton University,Princeton,NJ08544,USA.13U.S.Geological

Survey,Alaska Cooperative Fish and Wildlife Research Unit,

University of Alaska,Fairbanks,AK99775,USA.14Oak Ridge

National Laboratory,Oak Ridge,TN37831,USA.

*To whom correspondence should be addressed.E-mail:

ypan@https://www.wendangku.net/doc/9312873606.html,

RESEARCH ARTICLES

Here,we present bottom-up estimates of C stocks and fluxes for the world’s forests based on recent inventory data and long-term field obser-vations coupled to statistical or process models (table S1).We advanced our analyses by including comprehensive C pools of the forest sector(dead wood,harvested wood products,living biomass, litter,and soil)and report past trends and changes in C stocks across countries,regions,and conti-nents representing boreal,temperate,and tropical forests(5,6).To gain full knowledge of the trop-ical C balance,we subdivided tropical forests in-to intact and regrowth forests(Table1).The latter is an overlooked category,and its C uptake is usually not reported but is implicit in the tropical land-use change emission estimates.Although deforestation,reforestation,afforestation and the carbon outcomes of various management prac-tices are included in the assessments of boreal and temperate forest C sink estimates,we sepa-rately estimated three major fluxes in the tropics: C uptake by intact forests,losses from deforesta-tion,and C uptake of forest regrowth after an-thropogenic disturbances.The area of global forests used as a basis for estimating C stocks and fluxes is3.9billion ha,representing95%of the world’s forests(7)(table S2).

Global forest C stocks and changes.The current C stock in the world’s forests is estimated to be861T66Pg C,with383T30Pg C(44%)in soil(to1-m depth),363T28Pg C(42%)in live biomass(above and below ground),73T6Pg C (8%)in deadwood,and43T3Pg C(5%)in litter (table S3).Geographically,471T93Pg C(55%)

is stored in tropical forests,272T23Pg C(32%)

in boreal,and119T6Pg C(14%)in temperate

forests.The C stock density in tropical and boreal

forests is comparable(242versus239Mg C ha–1),

whereas the density in temperate forests is~60%

of the other two biomes(155Mg C ha–1).

Although tropical and boreal forests store the

most carbon,there is a fundamental difference in

their carbon structures:Tropical forests have56%

of carbon stored in biomass and32%in soil,

whereas boreal forests have only20%in biomass

and60%in soil.

The average annual change in the C stock of

established forests(Table1)indicates a large

uptake of2.5T0.4Pg C year–1for1990to1999

and a similar uptake of2.3T0.5Pg C year–1for

2000to2007.Adding the C uptake in tropical

regrowth forests to those values indicates a

persistent global gross forest C sink of4.0T0.7

Pg C year–1over the two periods(Tables1and2).

Despite the consistency of the global C sink since

1990,our analysis revealed important regional

and temporal differences in sink sizes.The C sink

in temperate forests increased by17%in2000to

2007compared with1990to1999,in contrast to

C uptake in intact tropical forests,which de-

creased by23%(but nonsignificantly).Boreal

forests,on average,showed little difference be-

tween the two time periods(Fig.1).Subtract-

ing C emission losses from tropical deforestation

and degradation,the global net forest C sink

was1.0T0.8and1.2T0.9Pg C year–1for

1990to1999and2000to2007,respectively

(Table1).

Forest carbon sinks by regions,biomes,and

pools.Boreal forests(1135Mha)had a consistent

average sink of0.5T0.1Pg C year–1for two dec-

ades(Table2,20and22%of the global C sinks

in established forests).However,the overall sta-

bility of the boreal forest C sink is the net result

of contrasting carbon dynamics in different boreal

countries and regions associated with natural dis-

turbances and forest https://www.wendangku.net/doc/9312873606.html,n Russia

had the largest boreal sink,but that sink showed

no overall change,even with increased emissions

from wildfire disturbances(8).In contrast,there

was a notable sink increase of35%in European

Russia(Fig.1)attributed to several factors:in-

creased areas of forests after agricultural aban-

donment,reduced harvesting,and changes of

forest age structure to more productive stages,

particularly for the deciduous forests(8).In con-

trast to the large increase of biomass sinks in

European Russia and northern Europe(8,9),the

biomass C sink in Canadian managed forests was

reduced by half between the two periods,mostly

due to the biomass loss from intensified wildfires

and insect outbreaks(10,11).A net loss of soil C

in northern Europe was attributed to shifts of

forest to nonforest in some areas.Overall,the

relatively stable boreal C sink is the sum of a net

reduction in Canadian biomass sink offset by

increased biomass sink in all other boreal regions,

and a balance between decreased litter and soil C

sinks in northern Eurasia and a region-wide in-

crease in the accumulation of dead wood(Table2).

Temperate forests(767Mha)contributed0.7T

0.1and0.8T0.1Pg C year–1(27and34%)to

the global C sinks in established forests for two

decades(Table2).The primary reasons for the

increased C sink in temperate forests are the

increasing density of biomass and a substantial

increase in forest area(12,13).The U.S.forest

C sink increased by33%from the1990s to

2000s,caused by increasing forest area;growth

of existing immature forests that are still recover-

ing from historical agriculture,grazing,harvesting

(12,14);and environmental factors such as CO2

fertilization and N deposition(15).However,for-

ests in the western United States have shown

considerably increased mortality over the past

few decades,related to drought stress,and in-

creased mortality from insects and fires(16,17).

The European temperate forest sink was stable

between1990to1999and2000to2007.There

was a large C sink in soil due to expansion of

forests in the1990s,but this trend slowed in the

2000s(7,18).However,the increased C sink in

biomass during the second period(+17%)

helped to maintain the stability of the total C sink.

China’s forest C sink increased by34%between

1990to1999and2000to2007,with the biomass

sink almost doubling(Table2).This was caused

primarily by increasing areas of newly planted

forests,the consequence of an intensive national

afforestation/reforestation program in the past

few decades(table S2)(19).

Table1.Global forest carbon budget(Pg C year–1)over two time periods.Sinks are positive values;

sources are negative values.

Carbon sink and source in biomes1990–19992000–20071990–2007

Boreal forest0.50T0.080.50T0.080.50T0.08

Temperate forest0.67T0.080.78T0.090.72T0.08

Tropical intact forest* 1.33T0.35 1.02T0.47 1.19T0.41

Total sink in global established forests? 2.50T0.36 2.30T0.49 2.41T0.42

Tropical regrowth forest? 1.57T0.50 1.72T0.54 1.64T0.52

Tropical gross deforestation emission§–3.03T0.49–2.82T0.45–2.94T0.47

Tropical land-use change emission||–1.46T0.70–1.10T0.70–1.30T0.70

Global gross forest sink? 4.07T0.62 4.02T0.73 4.05T0.67

Global net forest sink# 1.04T0.79 1.20T0.85 1.11T0.82

Equations of global forest C fluxes

F established forests=F boreal forests+F temperate forests+F tropical intact forests(Eq.1)

F tropical land-use change=F tropical gross deforestation+F tropical regrowth forests(Eq.2)

F gross forest sink=F established forests+F tropical regrowth forests(Eq.3)

F net forest sink=F established forests+F tropical land-use change(Eq.4)

*Tropical intact forests:tropical forests that have not been substantially affected by direct human activities;flux accounts for the

dynamics of natural disturbance-recovery processes.?Global established forests:the forest remaining forest over the study periods

plus afforested land in boreal and temperate biomes,in addition to intact forest in the tropics(Eq.1).?Tropical regrowth forests:

tropical forests that are recovering from past deforestation and logging.§Tropical gross deforestation:the total C emissions from

tropical deforestation and logging,not counting the uptake of C in tropical regrowth forests.||Tropical land-use change:emissions

from tropical land-use change,which is a net balance of tropical gross deforestation emissions and C uptake in regrowth forests(Eq.2).

It may be referenced as a tropical net deforestation emission in the literature.?Global gross forest sink:the sum of total sinks in

global established forests and tropical regrowth forests(Eq.3).#Global net forest sink:the net budget of global forest fluxes

(Eq.4).It can be calculated in two ways:(i)total sink in global established forests minus tropical land-use change emission or(ii)total

global gross forest sink minus tropical gross deforestation emission.

RESEARCH ARTICLES

Tropical intact forests(1392Mha)represent ~70%of the total tropical forest area(1949Mha) that accounts for the largest area of global forest biomes(~50%).We used two networks of per-manent monitoring sites spanning intact tropical forest across Africa(20)and South America(21)

and assumed that forest C stocks of Southeast

Asia(9%of total intact tropical forest area)are

changing at the mean rate of Africa and South

America,as we lack sufficient data in Southeast

Asia to make robust estimates.These networks

are large enough to capture the disturbance-

recovery dynamics of intact forests(6,20,22).

We estimate a sink of1.3T0.3and1.0T0.5Pg C

year–1for1990to1999and2000to2007,

Table2.Estimated annual change in C stock(Tg C year–1)by biomes by country or region for the time periods of1990to1999and2000to2007.Estimates include C stock changes on“forest land remaining forest land”and“new forest land”(afforested land).The uncertainty calculation refers to the supporting online material.ND,data not available;[1],litter is included in soils.

Biome and country/

region

1990–19992000–2007

Biomass

Dead

wood Litter Soil

Harvested

wood

product

Total

stock

change

Uncertainty

(T)

Stock

change

per area Biomass

Dead

wood Litter Soil

Harvested

wood

product

Total

stock

change

Uncertainty

(T)

Stock

change

per area (Tg C year–1)

(Mg C ha–1

year–1)(Tg C year–1)

(Mg C ha–1

year–1)

Boreal*

Asian

Russia6166634519255640.396997434213264660.39 European

Russia3710223641146370.93841935352619950 1.21 Canada6–24146232670.11–5316197211030.04 European

boreal?130338116516 1.122104–10132770.45 Subtotal1175310312594493760.451201321017473499830.44

Temperate*

United

States?118613933179340.721479183728239450.94 Europe11728812423258 1.7113729652723960 1.68 China602215317135340.9611524828718245 1.22 Japan249ND1925414 2.28235ND82379 1.59 South

Korea62ND50144 2.14122ND40185 2.86 Australia17ND1015850130.3317ND10141051130.34 New

Zealand10015720.911001692 1.05 Other

countries1ND ND ND0110.0720000320.18 Subtotal345424616080673780.9145442451568077789 1.03

Tropical intact

Asia125132ND5144380.88100102ND6117300.90 Africa469487ND95323020.94425436ND84822740.94 Americas573489ND226521660.77345455ND234183860.53 Subtotal116710917ND3513283470.848709813ND3610174740.71 Global

subtotal§163020416628620924943630.73144427315823018922944890.69

Tropical regrowth

Asia498ND[1]27ND526263 3.52564ND[1]30ND593297 3.53 Africa169ND[1]73ND242121 1.48188ND[1]83ND271135 1.47 Americas694ND[1]113ND807403 4.67745ND[1]113ND858429 4.56 Subtotal1361ND[1]213ND1574496 3.241497ND[1]226ND1723539 3.19

All tropics||

Asia623132275670266 2.14664102306711298 2.38 Africa638487739774325 1.06613436838753305 1.08 Americas1267489113221458436 1.421090455113231276577 1.30 Subtotal252910917213352903605 1.4023679813226362740718 1.38 Global

total?29912041664982094068615 1.0429412731584561894017728 1.04 *Carbon outcomes of forest land-use changes(deforestation,reforestation,afforestation,and management practices)are included in the estimates in boreal and temperate forests.?Estimates for the area that includes Norway,Sweden,and Finland.?Estimates for the continental U.S.and a small area in southeast Alaska.§Estimates for global established forests.||Estimates for all tropical forests including tropical intact and regrowth forests.?Areas excluded from this table include interior Alaska(51Mha in2007),northern Canada(118Mha in2007),and“other wooded land”reported to the Food and Agriculture Organization.

RESEARCH ARTICLES

respectively(Table2).An average C sink of1.2T 0.4Pg C year–1for1990to2007is approx-imately half of the total global C sink in estab-lished forests(2.4T0.4Pg C year–1)(Table1). When only the biomass sink is considered,about two-thirds of the global biomass C sink in estab-lished forests is from tropical intact forests(1.0 versus1.5Pg C year–1).The sink reduction in the period2000to2007(–23%)was caused by deforestation reducing intact forest area(–8%) and a severe Amazon drought in2005(21),which appeared strong enough to affect the tropics-wide decadal C sink estimate(–15%).Except for the Amazon drought,the recent excess of biomass C gain(growth)over loss(death)in tropical intact for-ests appears to result from progressively enhanced productivity(20,21,23).Increased dead biomass production should lead to enhanced soil C seques-tration,but we lack data about changes in soil C stocks for tropical intact forests,so the C sink for tropical intact forests may be underestimated.

Tropical land-use changes have caused net C releases in tropical regions by clearing forests for agriculture,pasture,and timber(24),second in magnitude to fossil fuel emissions(Table3).

Tropical land-use change emissions are a net balance of C fluxes consisting of gross tropical de-forestation emissions partially compensated by C sinks in tropical forest regrowth.They declined from1.5T0.7Pg C year–1in the1990s to1.1T0.7 Pg C year–1for2000to2007(Table1)due to reduced rates of deforestation and increased for-est regrowth(25).The tropical land-use change emissions were approximately equal to the total global land-use emissions(Tables1and3),be-

cause effects of land-use changes on C were

roughly balanced in extratropics(7,24,25).

Tropical deforestation produced significant

gross C emissions of3.0T0.5and2.8T0.5Pg

C year–1,respectively,for1990to1999and2000

to2007,~40%of the global fossil fuel emissions.

However,these large emission numbers are usu-

ally neglected because more than one half was

offset by large C uptake in tropical regrowth for-

ests recovering from the deforestation,logging,

or abandoned agriculture.

Tropical regrowth forests(557Mha)repre-

sent~30%of the total tropical forest area.The

C uptake by tropical regrowth forests is usually

implicitly included in estimated net emissions

of tropical land-use changes rather than estimated

independently as a sink(24).We estimate that

Boreal

Canada

N. Europe

Asian Russia European Russia

Tropical Regrowth

Carbon Flux 2000-2007

Forest Carbon Flux

2000-2007

Tropical Regrowth

Carbon Flux 1990-1999

Forest Carbon Flux

1990-1999

Tropical Gross Deforestation

C Emissions 1990-1999

Tropical Gross Deforestation

C Emissions 2000-2007

Fig.1.Carbon sinks and sources(Pg C year–1)in the world’s forests.Colored bars in the down-facing direction represent C sinks,whereas bars in the upward-facing direction represent C sources.Light and dark purple,global established forests(boreal,temperate,and intact tropical forests);light and dark green,tropical regrowth forests after anthropogenic disturbances;and light and dark brown,tropical gross deforestation emissions.

Table3.The global carbon budget for two time periods(Pg C year?1).There are different arrangements to account for elements of the global C budget(see also table S6).Here,the accounting was based on global C sources and sinks.The terrestrial sink was the residual derived from constraints of two major anthropogenic sources and the sinks in the atmosphere and oceans.We used the C sink in global established forests as a proxy for the terrestrial sink.

Sources and sinks1990–19992000–2007

Sources(C emissions)

Fossil fuel and cement* 6.5T0.47.6T0.4

Land-use change? 1.5T0.7 1.1T0.7

Total sources8.0T0.88.7T0.8

Sinks(C uptake)

Atmosphere? 3.2T0.1 4.1T0.1 Ocean? 2.2T0.4 2.3T0.4 Terrestrial(established forests)§ 2.5T0.4 2.3T0.5

Total sinks7.9T0.68.7T0.7

Global residuals||0.1T1.00.0T1.0

*See(2).?See(4,7,25).The global land-use change emission is approximately equal to the tropical land-use change emission, because the net carbon balance of land-use changes in temperate and boreal regions is neutral(24,38).?See(4).§Estimates

of C sinks in the global established forests(that are outside the areas of tropical land-use changes)from this study.Note that the carbon sink in tropical regrowth forests is excluded because it is included in the term of land-use change emission(see above and Table1).||Global C residuals are close to zero when averaged over a decade.Uncertainties in the global residuals indicate either a

land sink or source in the212Mha of forest not included here,on nonforest land,or systematic error in other source(overestimate)or

sink(underestimate)terms,or both.

RESEARCH ARTICLES

the C sink by tropical regrowth forests was1.6T 0.5and1.7T0.5Pg C year–1,respectively,for 1990to1999and2000to2007.Our results in-dicate that tropical regrowth forests were stronger C sinks than the intact forests due to rapid bio-mass accumulation under succession,but these estimates are poorly constrained because of sparse data(table S4)(6).Although distinguishing a C sink in tropical regrowth forests does not affect the estimated net emissions from tropical land-use changes,an explicit estimate of this compo-nent facilitates evaluating the complete C sink capacity of all tropical and global forests.

When all tropical forests,both intact and regrowth,are combined,the tropical sinks sum to2.9T0.6and2.7T0.7Pg C year–1over the two periods(Table1),and on average account for ~70%of the gross C sink in the world forests (~4.0Pg C year–1).However,with equally significant gross emissions from tropical de-forestation(Table1),tropical forests were nearly carbon-neutral.In sum,the tropics have the world’s largest forest area,the most intense contemporary land-use change,and the highest C uptake,but also the greatest uncertainty,showing that invest-ment in better understanding carbon cycling in the tropics should be a high priority in the future.

Deadwood,litter,soil,and harvested wood products together accounted for35%of the global sink and60%of the global forest C stock,showing the importance of including these components (Table2and table S3).Compared with biomass, estimates of these terrestrial carbon pools are gen-erally less certain because of insufficient data. For deadwood,there was a large sink increase in boreal forests over the past decade,caused by the recent increase in natural disturbances in Siberia and Canada.Increased deadwood carbon thus makes a major(27%)but possibly transient con-tribution to the total C sink in the boreal zone. Changes in litter C accounted for a relatively small and stable portion of the global forest C sink.However,litter C accumulation contributed 20%of the total C sink in boreal forests and,like deadwood,is vulnerable to wildfire disturbances. Changes in soil C stocks accounted for more than10%of the total sink in the world’s forests, largely driven by land-use changes.We may un-derestimate global soil C stocks and fluxes be-cause the standard1-m soil depth excludes some deep organic soils in boreal and tropical peat for-ests(26–28).We estimate the net C change in har-vested wood products(HWP),including wood in use and disposed in landfills,as described in the IPCC guidelines(29),attributing changes in stock to the region where the wood was harvested.Car-bon sequestration in HWP accounted for~8%of the total sink in established forests.This sink re-mained stable for temperate and tropical regions but declined dramatically in boreal regions because of reduced harvest in Russia in the past decade.

Data gaps,uncertainty,and suggested im-provements in global forest monitoring.We es-timated uncertainties based on a combination of quantitative methods and expert opinions(6).There are critical data gaps that affected both the

results presented here and our ability to report

and verify changes in forest C stocks in the fu-

ture.Data are substantially lacking for areas of

the boreal forest in North America,including

Alaska(51Mha)and Canadian unmanaged for-

ests(118Mha)(table S5).The forests in these

regions could be a small C source or sink,based

on the estimate of Canadian managed forests(10)

and modeling studies in Alaska(30).There is also

a lack of measurement data of soil C flux in trop-

ical intact forests,which may cause uncertainty

of10to20%of the estimated total C sink in these

forest areas.In addition,there is a large uncertainty

associated with the estimate of C stocks and fluxes

in tropical Asia,due to the absence of long-term

field measurements,and a notable lack of data

about regrowth rates of tropical forests worldwide.

Prioritized recommendations for improve-

ments in regional forest inventories to assess C

density,uptake,and emissions for global-scale

aggregation include the following:(i)Land mon-

itoring should be greatly expanded in the tropics

and in unsampled regions of northern boreal

forests.(ii)Globally consistent remote sensing of

land-cover change and forest-area is required to

combine the strengths of two observation sys-

tems:solid ground truth of forest C densities

from inventories and reliable forest areas from

remote sensing.(iii)Improved methods and greater

sampling intensity are needed to estimate non-

living C pools,including soil,litter,and dead wood.

(iv)Better data are required in most regions for

estimating lateral C transfers in harvested wood

products and rivers.

Forest carbon in the global context.The new

C sink estimates from world’s forests can con-

tribute to the much needed detection and attri-

bution that is required in the context of the global

carbon budget(2,4,25).Our results suggest that,

within the limits of reported uncertainty,the en-

tire terrestrial C sink is accounted for by C uptake

of global established forests(Table3),as the

balanced global budget yields near-zero residuals

with T1.0Pg C year–1uncertainty for both1990

to1999and2000to2007(Table3).Consequent-

ly,our results imply that nonforest ecosystems

are collectively neither a major(>1Pg)C sink

nor a major source over the two time periods that

we monitored.Because the tropical gross de-

forestation emission is mostly compensated by

the C uptakes in both tropical intact and regrowth

forests(Fig.1and Table1),the net global forest

C sink(1.1T0.8Pg C year–1)resides mainly in

the temperate and boreal forests,consistent with

previous estimates(31,32).Notably,the total

gross C uptake by the world’s established and

tropical regrowth forests is4.0Pg C year–1,

which is equivalent to half of the fossil fuel C

emissions in2009(4).Over the period that we

studied(1990to2007),the cumulative C sink

into the world’s established forests was~43Pg C

and73Pg C for the established plus regrowing

forests;the latter equivalent to60%of cumula-

tive fossil emissions in the period(i.e.,126Pg C).

Clearly,forests play a critical role in the Earth’s

terrestrial C sinks and exert strong control on the

evolution of atmospheric CO2.

Drivers and outlook of forest carbon sink.

The mechanisms affecting the current C sink in

global forests are diverse,and their dynamics will

determine its future longevity.The C balance of

boreal forests is driven by changes in harvest

patterns,regrowth over abandoned farmlands,

and increasing disturbance regimes.The C balance

of temperate forests is primarily driven by forest

management,through low harvest rates(Europe)

(33),recovery from past harvesting and agricul-

tural abandonment(U.S.)(34),and large-scale

afforestation(China)(19).For tropical forests,

deforestation and forest degradation are dom-

inant causes of C emissions,with regrowth and

an increase in biomass in intact forests being the

main sinks balancing the emissions(23,24).

Changes in climate and atmospheric drivers

(CO2,N-deposition,ozone,diffuse light)affect the

C balance of forests,but it is difficult to separate

their impacts from other factors using ground

observations.For Europe,the U.S.,China,and

the tropics,evidence from biogeochemical pro-

cess models suggests that climate change,in-

creasing atmospheric CO2,and N deposition are,

at different levels,important factors driving the

long-term C sink(15,18,20,23,34).Drought

in all regions and warmer winters in boreal re-

gions reduce the forest sink through suppressed

gross primary production,increased tree mortal-

ity,increased fires,and increased insect damage

(8,10,18,21,30,35,36).

Our estimates suggest that currently the glob-

al established forests,which are outside the areas

of tropical land-use changes,alone can account

for the terrestrial C sink(~2.4Pg C year–1).The

tropics are the dominant terms in the exchange

of CO2between the land and the atmosphere.A

large amount of atmospheric CO2has been se-

questrated by the natural system of forested lands

(~4.0Pg C year–1),but the benefit is substantially

offset by the C losses from tropical deforestation

(~2.9Pg C year–1).This result highlights the po-

tential for the United Nations Reducing Emissions

from Deforestation and Degradation program to

lessen the risk of climate change.However,an

important caveat is that adding geological carbon

from fossil fuels into the contemporary carbon

cycle and then relying on biospheric sequestra-

tion is not without risk,because such sequestra-

tion is reversible from either climate changes,direct

human actions,or a combination of both.

Nonetheless,C sinks in almost all forests

across the world(Fig.1)may suggest overall fa-

vorable conditions for increasing stocks in forests

and wood products.Our analysis also suggests

that there are extensive areas of relatively young

forests with potential to continue sequestering

C in the future in the absence of accelerated

natural disturbance,climate variability,and land-

use change.As a result of the large C stocks in

both boreal forest soils and tropical forest bio-

mass,warming in the boreal zone,deforestation,

RESEARCH ARTICLES

and occasional extreme drought,coincident with fires in the tropics,represent the greatest risks to the continued large C sink in the world’s for-ests(21,24,30,37).A better understanding of the role of forests in biosphere C fluxes and mech-anisms responsible for forest C changes is critical for projecting future atmospheric CO2growth and guiding the design and implementation of mitigation policies.

Reference and Notes

1.G.J.Nabuurs et al.,in Climate Change2007:Mitigation,

B.Metz,O.R.Davidson,P.R.Bosch,R.Dave,L.A.Meyer,Eds.

(Cambridge Univ.Press,Cambridge,2007),pp.542–584.

2.J.G.Canadell et al.,Proc.Natl.Acad.Sci.U.S.A.104,

18866(2007).

3.S.Khatiwala,F.Primeau,T.Hall,Nature462,346(2009).

4.C.Le Quéréet al.,Nat.Geosci.2,831(2009).

5.R.K.Dixon et al.,Science263,185(1994).

6.Details of data sources,accounting,and estimation

methods used for each country,region,and C component are provided in the supporting online material.

7.Food and Agriculture Organization,Global Forest

Resources Assessment2010(Food and Agriculture

Organization,Rome,2010),forestry paper163.

8.A.Z.Shvidenko,D.G.Schepaschenko,S.Nilsson,in

Basic Problems of Transition to Sustainable Forest

Management in Russia,V.A.Sokolov,A.Z.Shvidenko,

O.P.Vtorina,Eds.(Russian Academy of Sciences,

Krasnoyarsk,Russia,2007),pp.5–35.

9.P.E.Kauppi et al.,For.Ecol.Manage.259,1239(2010).

10.W.A.Kurz,G.Stinson,G.J.Rampley,C.C.Dymond,

E.T.Neilson,Proc.Natl.Acad.Sci.U.S.A.105,1551(2008).

11.G.Stinson et al.,Glob.Change Biol.17,2227(2011).12.R.Birdsey,K.Pregitzer,A.Lucier,J.Environ.Qual.35,

1461(2006).

13.P.E.Kauppi et al.,Proc.Natl.Acad.Sci.U.S.A.103,

17574(2006).

14.Y.Pan et al.,Biogeosciences8,715(2011).

15.Y.Pan,R.Birdsey,J.Hom,K.McCullough,For.Ecol.Manage.

259,151(2009).

16.P.J.van Mantgem et al.,Science323,521(2009).

17.D.D.Breshears et al.,Proc.Natl.Acad.Sci.U.S.A.102,

15144(2005).

18.P.Ciais et al.,Nat.Geosci.1,425(2008).

19.J.Fang,A.Chen,C.Peng,S.Zhao,L.Ci,Science292,

2320(2001).

20.S.L.Lewis et al.,Nature457,1003(2009).

21.O.L.Phillips et al.,Science323,1344(2009).

22.M.Gloor et al.,Glob.Change Biol.15,2418(2009).

23.S.L.Lewis,J.Lloyd,S.Sitch,E.T.A.Mitchard,

https://www.wendangku.net/doc/9312873606.html,urance,Annu.Rev.Ecol.Syst.40,529(2009).

24.R.A.Houghton,Annu.Rev.Earth Planet.Sci.35,313(2007).

25.P.Friedlingstein et al.,Nat.Geosci.3,811(2010).

26.C.Tarnocai et al.,Global Biogeochem.Cycles23,

GB2023(2009).

27.A.Hooijer et al.,Biogeosciences7,1505(2010).

28.S.E.Page,J.O.Rieley,C.J.Banks,Glob.Change Biol.

17,798(2011).

29.Intergovernmental Panel on Climate Change,IPCC

Guidelines for National Greenhouse Gas Inventories

(Institute for Global Environmental Strategies,Japan,2006);

www.ipcc-nggip.iges.or.jp/public/2006gl/index.html.

30.A.D.McGuire et al.,Ecol.Monogr.79,523(2009).

31.C.L.Goodale et al.,Ecol.Appl.12,891(2002).

32.J.L.Sarmiento et al.,Biogeosciences7,2351(2010).

33.E.D.Schulze et al.,Nat.Geosci.2,842(2009).

34.S.W.Pacala et al.,Science292,2316(2001).

35.O.L.Phillips et al.,Philos.Trans.R.Soc.London Ser.B

359,381(2004).

36.J.M.Metsaranta,W.A.Kurz,E.T.Neilson,G.Stinson,

Tellus62B,719(2010).

37.M.Zhao,S.W.Running,Science329,940(2010).

38.R.A.Houghton,Tellus55B,378(2003).

Acknowledgments:This study is the major output of two

workshops at Peking Univ.and Princeton Univ.Y.P.,

R.A.B.,and J.F.were lead authors and workshop

organizers;Y.P.,R.A.B.,J.F.,R.H.,P.E.K.,W.A.K.,O.L.P.,

A.S.,and S.L.L.contributed primary data sets and

analyses;J.G.C.,P.C.,R.B.J.,and S.W.P.contributed

noteworthy ideas to improve the study;A.D.M.,S.P.,

A.R.,S.S.,and D.H.provided results of modeling or

data analysis relevant to the study;and all authors

contributed in writing,discussions,or comments.We

thank K.McCullough for helping to make the map in

Fig.1and C.Wayson for helping to develop a

Monte-Carlo analysis.This work was supported in part

by the U.S.Forest Service,NASA(grant31021001),the

National Basic Research Program of China on Global

Change(2010CB50600),the Gordon and Betty Moore

Foundation,Peking Univ.,and Princeton Univ.This work

is a contribution toward the Global Carbon Project’s aim

of fostering an international framework to study the

global carbon cycle.

Supporting Online Material

https://www.wendangku.net/doc/9312873606.html,/cgi/content/full/science.1201609/DC1

Materials and Methods

SOM Text

Tables S1to S6

References

13December2010;accepted29June2011

Published online14July2011;

10.1126/science.1201609

REPORTS

Detection of Emerging Sunspot Regions in the Solar Interior

Stathis Ilonidis,*Junwei Zhao,Alexander Kosovichev

Sunspots are regions where strong magnetic fields emerge from the solar interior and where major eruptive events occur.These energetic events can cause power outages,interrupt telecommunication and navigation services,and pose hazards to astronauts.We detected subsurface signatures of emerging sunspot regions before they appeared on the solar disc.Strong acoustic travel-time anomalies of an order of12to16seconds were detected as deep as65,000kilometers.These anomalies were associated with magnetic structures that emerged with an average speed of0.3to0.6kilometer per second and caused high peaks in the photospheric magnetic flux rate1to2days after the detection of the anomalies.Thus,synoptic imaging of subsurface magnetic activity may allow anticipation of large sunspot regions before they become visible,improving space weather forecast.

U nderstanding solar magnetism is among the most important problems of solar phys-

ics and astrophysics(1–5).Modern theo-ries assume that sunspot regions are generated by a dynamo action at the bottom of the convection zone,about200Mm below the photosphere.How-ever,there is no convincing observational evidence to support this idea,and dynamo mechanisms op-erating in the bulk of the convection zone or even

in the near-surface shear layer have been pro-

posed as well(6,7).Investigation of emerging

magnetic flux could possibly determine the depth

of this process and set the foundations for a better

understanding of sunspots and active regions.

Active regions on the Sun produce flares and

mass eruptions that may cause power outages

on Earth,satellite failures,and interruptions of

telecommunication and navigation services.Moni-

toring solar subsurface processes and predict-

ing magnetic activity would also improve space

weather forecasts.

Time-distance helioseismology(8)is one of

the local helioseismology techniques that image

acoustic perturbations in the interior of the Sun

(9).Acoustic waves are excited by turbulent con-

vection near the surface,propagate deep inside

the Sun,and are refracted back to the surface

(Fig.1).Time-distance helioseismology measures

travel times of acoustic waves propagating to dif-

ferent distances by computing cross-covariances

between the oscillation signals observed at pairs

of locations on the solar photosphere.Varia-

tions in acoustic travel times are caused mainly

by thermal perturbations,magnetic fields,and

flows.Previous studies of emerging sunspot re-

gions(10–14)have found difficulties in detect-

ing signals deeper than30Mm and before the

initial magnetic field becomes visible on the sur-

face because of the fast emergence speed and low

signal-to-noise ratio(15).Here,we present a deep-

focus time-distance measurement scheme,which

allows us to detect signals of emerging magnetic

regions in the deep solar interior(16,17).

We have used Doppler observations(18)from

Michelson Doppler Imager(MDI)(19)onboard

the Solar and Heliospheric Observatory(SOHO)

and computed travel-time maps of four emerging

flux regions and nine quiet regions.In Fig.2,we

present the results of our analysis for Active Re-

gion(AR)10488,which started emerging on the

solar disc at09:30UT,26October2003,about

W.W.Hansen Experimental Physics Laboratory,Stanford University,Stanford,CA94305–4085,USA

*To whom correspondence should be addressed.E-mail: ilonidis@https://www.wendangku.net/doc/9312873606.html,

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