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1 Chinese Journal of App lied Ecology, May 2010, 21 (5) : , ( 1, ; 2, ; 3, ) CO 2 CO 2 (N EE)., 2003 CO 2,CO 2. :CO 2,. CO 2 CO 2 CO 2. CO 2, N EE 25% 19%,, N EE 10% 25% ; CO 2,M ichaelis2menten L loyd2taylor, R ref ; CO 2,, (GPP)(R e )20%. CO 2 CO (2010) S718 A Effects of CO 2 storage flux on carbon budget of forest ecosystem. ZHANG M i 1, 2, W EN Xue2 fa 1, YU Gui2rui 1, ZHANG Lei2m ing 1, FU Yu2ling 1, SUN Xiao2m in 1, HAN Shi2jie 3 ( 1 Institute of Geographic Sciences and N atura l R esou rces Research, Chinese A cadem y of S ciences, B eijing , China; 2 Graduate U niversity of Ch inese A cadem y of Sciences, B eijing , China; 3 Institute of Applied Ecology, Ch inese A cadem y of Sciences, Shenyang , Ch ina ). 2Chin. J. A ppl. Ecol., 2010, 21 (5) : Abstract: Carbon dioxide (CO 2 ) storage flux in the air space below measurement height of eddy covariance is very important to correctly evaluate net ecosystem exchange of CO 2 (N EE ) between forest ecosystem and atmosphere. This study analyzed the dynam ic variation of CO 2 storage flux and its effects on the carbon budget of a temperate broad2leaved Korean p ine m ixed forest at Changbai Mountains, based on the eddy covariance flux data and the vertical p rofile of CO 2 concentration da2 ta. The CO 2 storage flux in this forest ecosystem had typ ical diurnal variation, with the maximum variation appeared during the transition from stable atmospheric layer to unstable atmospheric layer. The CO 2 storage flux calculated by the change in CO 2 concentration throughout a vertical p rofile was not significantly different from that calculated by the change in CO 2 concentration at the measure2 ment height of eddy covariance. The NEE of this forest ecosystem was underestimated by 25% and 19% at night and at daytime, respectively, without calculating the CO 2 storage flux at half2hour scale, and was underestim ated by 10% and 25% at daily scale and annual scale, respectively. W ithout calculating the CO 2 storage flux in this forest ecosystem, the parameters ofm ichaelis2ment2 en equation and L loyd2taylor equation were underestim ated, and the ecosystem apparent quantum yield ( ) and the ecosystem resp iration rate (R ref ) at the reference temperature were mostly affect2 ed. The gross p rimary p roductivity (GPP) and ecosystem resp iration (R e ) of this forest ecosystem were underestimated about 20% without calculating the CO 2 storage flux at half2hour, daily scale, and annual scale. Key words: forest ecosystem; CO 2 storage flux; net ecosystem CO 2 exchange. 3 ( G2010CB833500) ( ) ( KZCX22YW )( ) E2mail: yugr@ igsnrr. ac. cn ,

2 CO 2 ( net ecosystem exchange, N EE) [ 1-4 ]., CO 2 [ 5-6 ]. 400 ( FLUX2 NET) [ 1 ].,, ( z r ) CO 2 ( F c ) CO 2 [ 7-8 ].,CO 2, CO 2, CO 2 0.,, CO 2 (N EE) CO 2 ( F c ) CO 2 ( F s ) [ 6, 9-11 ]., F s ;, F s [ 12 ]., CO 2 CO 2., CO 2 CO 2 [ ] CO 2 [ ]. Hollinger [ 18 ],.,, /, CO 2, CO 2 CO 2 [ ]. (ChinaFLUX) 7, CO 2,, CO 2 CO 2., CO 2, CO 2., CO 2 N EE [ 13, 16, 22 ]. CO 2 N EE. N EE ( gross p rimary p roductivity, GPP ) ( ecosystem resp iration, R e ), M ichaelis2menten L loyd2taylor GPP R e., CO 2, GPP R e.,, CO 2 ;CO 2 ; CO ( N, E, 738 m )., 5 9, 10 4.,,,,. 316, 695 mm., < 4.,( P inus ko2 ra iensis) ( Tilia am urensis) ( Q uercus m ongolica) ( F raxinus m andshurica ) (A cer m ino)., 40%, 26 m, 560 hm - 2. [ ] m, ( CSAT3, Campbell Scientific, USA ) CO 2 / H 2 O(L i7500, L i2cor, USA ) CR5000 ( CR5000, Campbell Scientific, USA ). 10 Hz, 30 m in. CO % L i610(l i2cor, USA ) L i (HMP45C, Vaisala, Helsinki, Finland), m, (L i2190sb, L i2cor Inc1, USA ) 32 m,

3 5 : 1203 (105T, Campbell, USA ) cm. 015 Hz, ( CR23X2TD /CR10X2 TD, Campbell Scientific, USA ) 30 m in [ 23 ]. CO m 7, CO 2 CO 2 (L i2820, L i2cor Inc, USA). 60 m in 489 mol mol - 1 CO 2, 1 mol mol s,60 s. ( CR10X2TD, Campbell Scientific Inc, USA ), 30 m in CO 2 [ 22, 25 ].,CO 2,., CO 2, CO 2, z r CO 2 ( F c ) [ 7-8 ] : F c = w c ( z r ) (1) : w ; c CO 2 ;., CO 2, CO 2 N EE CO 2 ( F c )CO 2 ( F s ) : N EE = F c + F s (2) (2) FLUXNETCO 2 [ 6, 9-11 ] CO 2 CO 2,CO 2 [ 13, ] : F s z 5gc = 0 5t dz (3) : F s 40 m CO 2 (mg CO 2 m - 2 s - 1 ) ; z ; gc CO 2 ; t. CO 2, m CO m CO 2, CO m CO 2. CO 2,CO 2 [ ] : F s = c h (4) t : c CO 2 ; t, 30 m in; h ,, [ 26 ],, CO 2, 30 m in. : 1) ; 2) ; 3), ; ; ; 5 3 ; 4),, [ ] M ichaelis2menten [ ] : P ec = PAR P ec, max PAR + P ec, max - R e (5) : PAR ( mol m - 2 s - 1 ) ; (mg CO 2 mol - 1 photon) ; P ec (mg CO 2 m - 2 s - 1 ) ; P ec max (mg CO 2 m - 2 s - 1 ) ; R e (mg CO 2 m - 2 s - 1 ). ( ) L loyd2taylor [ 31 ]. R e = R ref e E 0 [ 1 / ( T ref- T 0 ) - 1 / ( T - T 0 ) ] (6) : R e CO 2 (mg CO 2 m - 2 s - 1 ) ; T5 cm; R ref T ref (28311 K) ; E 0 ; T 0 ( , K)., CO 2,CO 2 F c CO 2 N EE(5) P ec ( 6) R e

4 N EE N EE., L loyd2 Taylor ( 6) [ 31 ] R e. :N EE 5 cm (6),, R e. R e (7) GPP. GPP = R e - N EE (7) CO 2 R e GPP,(6)CO 2 R e CO 2 R es,(7) R e F c R es N EECO 2 GPP CO 2 GPP s., F c N EE CO 2, CO 2. F c N EE CO 2, CO CO 2., CO 2 F s 1., u 3 0; u 3 > 015 m s - 1, F s < 015m s - 1,, F s u 3, F s u 3, CO 2 0. Aubinet [ 13 ], u > 018 m s - 1, F s 2, CO 2, CO 2, 0. CO 2 [ 2, 13 ]., 12: 00,F s 0 ; 18: 00,, CO 2, F s, 22: 00;, F s, F s,, CO 2 [ 13 ]. CO 2, CO 2 CO 2 ; CO 2.,, F s ;, F s, 6: 00. 6: 00, F s,, F s. CO 2, F c N EE ( 2) : F s 0, F c N EE; F s 0, F c N EE. 1 CO 2 F ig. 1 Change of CO 2 storage flux with friction velocity in a temperate broad2leaved Korean p ine m ixed forest at Changbai Mountains. F s 1: CO 2 CO 2 The CO 2 storage flux calculated by the change in CO 2 concentration throughout a vertical p ro2 file; F s 2:CO 2 CO 2 The CO 2 storage flux calculated by the change in CO 2 concentration at the measurement height of eddy covariance. The same below. 2 ( 6 8 ) CO 2 F s F c N EE F ig. 2 Monthly average daily variations of F s, F c and N EE in a temperate broad2leaved Korean p ine m ixed forest at Changbai Mountains in m id2growing season ( from June to August).

5 5 : CO 2 t Tab. 1 Independen t2sam ple t test about sign if icance of d ifference between the CO 2 storage term ca lcula ted by two m ethods ( = 0105) Two methods Levene Levene s test for equality of variances F Sig. t df ( ) Sig. (2 Tailed) Mean difference T2test for equality of means Std. error difference 95% 95% Confidence interval of the difference Lower Upper Equal variances assumed Equal variance not assumed , CO 2,. CO 2,, CO 2 CO 2 ( F s 2) ( F s 1). CO 2, CO 2, CO 2, F s 1 ; CO 2, CO 2, CO 2,,CO 2 CO 2. ( 3),, F s 2F s 1, F s 2 F s 1 62% (, )., t 0105, 3CO 2 F ig. 3 Comparison of the CO 2 storage flux calculated by two methods in a temperate broad2leaved Korean p ine m ixed forest at ChangbaiMountains at half2hour scale.., ( 1)., CO 2, CO 2. CO 2,,CO 2, CO 2 CO CO 2 N EE 2, CO 2 F c N EE., F c N EE( 4),CO 2, F c N EE 75% 81%., CO 2 N EE 25%, N EE 19%. CO 2 CO 2.,CO 2 N EE 10% ( 5)., 2003 F c N EE g C m - 2 a - 1,CO 2 CO 2 25%., /,, CO 2, CO 2 [ 19 ]., CO 2 CO 2 [ 16, 22 ].,,CO 2, / CO 2

6 ( a)( b) F c N EE F ig. 4 Relationship between F c and N EE in a temperate broad2 leaved Korean p ine m ixed forest at Changbai Mountains at night ( a) and daytime ( b) at half2hour scale F c N EE F ig. 5 Relationship between F c and N EE in a temperate broad2 leaved Korean p ine m ixed forest at Changbai Mountains at daily scale..,,, / [ 16 ].,, /, CO 2, CO 2, CO 2.,CO 2 N EE.,CO 2 N EE,., CO 2 / N EE [ 19, 32 ]. N EE.,/ CO 2 N EE,. 213 CO 2 CO 2,. (6 8 ),( 5) F c N EE PAR, 2 : F c PAR P max R e N EE PAR,3 36% 5% 20%. CO 2, P max..,, CO 2, F c N EE ( 2), CO 2. P max,, CO 2 0, F c N EE ( 2), CO 2. (6)2003 F c N EE5 cm, 3 : F c T s R ref E 0 N EE T s, 2 18% 013%. CO 2 R ref.,co 2, F c N EE, R ref. 214 CO 2 GPP R e CO 2 N EE GPP R e. 6,,CO 2

7 5 : Tab. 2 Param eters of M ichaelis2m en ten equa tion in a tem pera te broad2leaved Korean p ine m ixed forest a t Changba i M oun ta in s from June to August in June 7 July 8 August P max R d R 2 P max R d R 2 P max R d R 2 (mg CO 2 (mg CO 2 (mg CO 2 (mg CO 2 (mg CO 2 (mg CO 2 (mg CO 2 (mg CO 2 (mg CO 2 mol - 1 m - 2 m - 2 mol - 1 m - 2 m - 2 mol - 1 m - 2 m - 2 Photon) s - 1 ) s - 1 ) Photon) s - 1 ) s - 1 ) Photon) s - 1 ) s - 1 ) PAR 2F c PAR 2N EE Tab. 3 Param eters of L loyd2taylor equa tion in a tem per2 a te broad2leaved Korean p ine m ixed forest a t Changba i M oun ta in s in 2003 R ref (mg CO 2 m - 2 s - 1 ) E 0 (J mol - 1 ) T s 2F c T s 2N EE R 2 7,CO 2 R e GPP CO 2 R es GPP s 81% 83%. CO 2 R e GPP 19% 17%., CO R e GPP R es GPP s : g C m - 2 a g C m - 2 a - 1. CO 2 19% 20%. CO 2 N EE,CO 2, 6 CO 2 R e ( a) GPP ( b) F ig. 6 Effects of CO 2 storage flux on R e ( a) and GPP ( b) in a temperate broad2leaved Korean p ine m ixed forest at Changbai Mountains at half2hour scale. R e GPP: CO 2 The values calculated without calculating CO 2 storage flux; R es GPP s : CO 2 The values calculated including CO 2 storage flux. The same below. R e GPP CO 2 R es GPP s 80% 82%.,CO 2 R e GPP 20% 18%. 7 CO R e ( a)gpp ( b) F ig. 7 Effects of CO 2 storage term on R e ( a) and GPP ( b) in a temperate broad2leaved Korean p ine m ixed forest at Changbai Mountains at daily scale.

8 %. 3, CO 2,,. CO 2 CO 2 CO 2, CO 2 CO 2. CO 2 M ichaelis2menten L loyd2taylor, ;CO 2 N EE, N EE ;CO 2 GPP R e 20%. CO 2. CO 2 /,,, CO 2 CO 2, CO 2 ( / ), CO 2 CO 2. [ 1 ] Baldocchi D. B reathing of the terrestrial biosphere: Lessons learned from a global network of carbon dioxide flux measurement system s. ny, 2008, 56: 1-26 A ustralian Journal of B ota2 [ 2 ] Goulden ML, Munger JM, Fan SM, et al. Measurement of carbon sequestration by long2term eddy covariance: Methods and a critical evaluation of accuracy. Change B iology. 1996, 2: Global [ 3 ] Yu G2R (), Sun X2M (). Theory and Methodology for FluxM easurement in Terrestrial Ecosys2 tem. Beijing: H igher Education Press, 2006 ( in Chi2 nese) [ 4 ] Yu GR, W en XF, Sun XM, et al. Overview of China2 FLUX and evaluation of its eddy covariance measure2 ment. A gricultural and Forest M eteorology, 2006, 137: [ 5 ] Berbigier P, Bonnefond JM, Mellmann P. CO 2 and wa2 ter vapor fluxes for 2 years above Euroflux forest site. A gricultural and Forest M eteorology, 2001, 108: [ 6 ] Wofsy SC, Goulden ML, Munger JW, et al. Net ex2 change of CO 2 in a m id2latitude forest. Science, 1993, 260: [ 7 ] Moncrieff JB, Malhi Y, Leuning R. The p ropagation of errors in long2term measurement of land2atmosphere flu2 xes of carbon and water. Global Change B iology, 1996, 2: [ 8 ] Moncrieff JB, M assheder JM, Elbers J. A system to measure surface fluxes of momentum, sensible heat, wa2 ter vapor and carbon dioxide. 1997, 188: Journal of Hydrology, [ 9 ] AubinetM, Grelle A, Ibrom A, et al. Estimates of the annual net carbon and water exchange of European for2 ests: The EUROFLUX methodology. logical Research, 2000, 30: A dvanced in Eco2 [ 10 ] B lack TA, Hartog G, Neumann HH, et al. Annual cy2 cles of water vapour and carbon dioxide fluxes in and above a boreal aspen forest. 1996, 2: Global Change B iology, [ 11 ] Greco S, Baldocchi D. Seasonal variations of CO 2 and water vapour exchange rates over a temperate deciduous forest. Global Change B iology, 1996, 2: [ 12 ] Goulden ML, Munger JW, Fan SM, et al. Exchange of carbon dioxide by a deciduous forest: Response to inter2 annual climate variability. Science, 1996b, 271: [ 13 ] AubinetM, Berbigier P, Bernhofer CH, et al. Compa2 ring CO 2 storage and advection conditions at night at dif2 ferent CarboEuroFLUX sites. B oundary LayerM eteorolo2 gy, 2005, 116: [ 14 ] Finnigan J. Response to comment by D r. A. S. Kowals2 ki on The storage term in eddy flux calculations. A gricultural and Forest M eteorology, 2009, 149: [ 15 ] Finnigan J. The storage term in eddy flux calculations. A gricultural and Forest M eteorology, 2006, 136: [ 16 ] Lee X. On m icrometeorological observation of surface2air exchange over tall vegetation. M eteorology, 1998, 91: A gricultural and Forest [ 17 ] Griffis TJ, B lack TA, Morgenstern K, et al. Ecophysio2 logical controls on the carbon balances of three southern boreal forests. A gricultural and Forest M eteorology, 2003, 117: [ 18 ] Hollinger DY, Kelliher FM, Byers JN, et al. Carbon di2

9 5 : 1209 oxide exchange between an undisturbed old2growth tem2 perate forest and the atmosphere. Ecology, 1994, 75: [ 19 ] Aubinet M, Heinesch B, Longdoz B. Estimation of the carbon sequestration by a heterogeneous forest: N ight flux corrections, heterogeneity of the site and inter2an2 nual variability. Global Change B iology, 2002, 8: [ 20 ] Hutyra LR, Munger JW, Hammond2Pyle E. et al. Re2 solving systematic errors in estimates of net ecosystem exchange of CO 2 and ecosystem resp iration in a trop ical forest biome. A gricultural and Forest M eteorology, 2008, 148: [ 21 ] Staebler RM, Fitzjarrald DR. Observing subcanopy CO 2 advection. A gricultural and Forest M eteorology, 2004, 122: [ 22 ] W u J2B (), Guan D2X (), Zhao X2S (), et al. CO 2 concentration character in broad2 leaved Korean p ine forest of Changbai Mountain. Chi2 nese Journal of A pplied Ecology ( ), 2005, 16 (1) : ( in Chinese) [ 23 ] Guan D2X (), W u J2B (), Yu G2R (), et al. Meteorological control on CO 2 flux above broadleaved Korean p ine m ixed forest in Changbai Mountains. Science in China, Series D ( D ), 2004, 34 ( Supp l. ) : ( in Chinese) [ 24 ] Zhang M ( ), Guan D2X (), Han S2J (), et al. Climatic dynam ics of broadleaved Ko2 rean p ine forest in ChangbaiMountain during the last 22 Year. Chinese Journal of Ecology ( ), 2005, 24 ( 9) : ( in Chinese) [ 25 ] Han SJ, Sha LQ, Yu GR, et al. Dynam ics of p rofiles and storage of carbon dioxide in broadleaved / Korean for2 est in ChangbaiMountain. Journal of Forestry Research, 2003, 14: [ 26 ] Falge E, Baldocchi D, O lson R, et al. Gap filling strat2 egies for defensible annual sum s of net ecosystem ex2 change. A gricultural and Forest M eteorology, 2001, 107: [ 27 ] Zhang L2M (), Yu G2R (), Sun X2M (), et al. Seasonal variation of carbon exchange of typ ical forest ecosystem s along the eastern forest tran2 sect in China. Science in China, Series D ( D ), 2006, 36 ( Supp l. I) : ( in Chinese) [ 28 ] AubinetM, Chermanne B, Vandenhaute M, et al. Long term carbon dioxide exchange above a m ixed forest in the Belgian A rdennes. A gricultural and Forest M eteorol2 ogy, 2001, 108: [ 29 ] Goulden ML, Daube BC, Fan SM, et al. Physiological responses of a black sp ruce forest to weather. Journal of Geophysical R esearch, 1997, 102: [ 30 ] Law BE, Falge E, Gu LH, et al. Environmental con2 trols over carbon dioxide and water vapor exchange of terrestrial vegetation. A gricultural and Forest M eteorolo2 gy, 2002, 113: [ 31 ] L loyd J, Taylor JA. On the temperature dependence of soil resp iration. Functional Ecology, 1994, 8: [ 32 ] W en X2F (), Yu G2R (), Sun X2M ( ). Uncertainties in long2term studies of net ecosystem CO 2 eddy covariance technique. 2004, 19: exchange with the atmosphere based on A dvances in Earth Science,,, 1980,., 5. E2mail: zhangm. igsnrr. ac. cn

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