1 Target Journal: Remote Sensing of Environment 2 Global Comparison of Light Use Efficiency Models for Vegetation Gross Primary 3 Production based on Eddy Covariance Towers Data 4 Wenping Yuan1, Wenwen Cai1, Jiquan Chen2, Shuguang Liu3,4, Wenjie Dong1, Dan Liu1, 5 Jiangzhou Xia1, Yang Chen1, other coauthors 6 1State 7 Change and Earth System Science, Beijing Normal University, Beijing 100875, China; 8 2Department 9 3United Key Laboratory of Earth Surface Processes and Resource Ecology, College of Global of Environmental Sciences, University of Toledo, Toledo, OH 43606, USA; States Geological Survey, Earth Resources Observation and Science Center, Sioux Falls, 10 South Dakota 57198, USA; 11 4 12 South University of Forestry and Technology, Changsha, Hunan 410004, China; 13 Abstract State Engineering Laboratory of Southern Forestry Applied Ecology and Technology, Central 14 Predicting the gross primary productivity (GPP) of terrestrial ecosystems has been a 15 major challenge in quantifying the global carbon cycle. Among all the predictive methods, the 16 light use efficiency (LUE) model may have the most potential to adequately address the spatial 17 and temporal dynamics of GPP because of its theoretical basis and practicality. Many different 18 LUE models have been developed recently, but our understanding of the relative merits of 19 different models is poor. Using carbon flux measurements data from 155 eddy covariance sites, we 20 assessed the ability of seven LUE models and compared the major model algorithms. 21 Comparisons between modeled and observed GPP showed that the model performance 1 22 substantially differed among ecosystem types. In generally, high model performance was found 23 over the deciduous broadleaf forests and mixed forests, and low performance was observed over 24 the evergreen broadleaf forests and shrublands. Except CFlux model, over the cloudy and overcast 25 days, other six models showed a significant underestimation due to ignoring the impacts of diffuse 26 radiation on light use efficiency. Among seven models, CFlux and EC-LUE showed the better 27 performance than averaged level at the 76% and 75% sites. All models were examined for 28 simulating the interannual variability of GPP observations, and the higher simulation accuracy was 29 found at CFlux and EC-LUE models. Paired comparisons showed the models differences majorly 30 resulted from the differences of environmental regulations equations compared with the fraction of 31 PAR absorbed by the vegetation canopy, and especially water stress equations substantially 32 differed among seven models. 33 Key words 34 Gross primary production; Light use efficiency; CASA; C-Fix; CFlux; EC-LUE; 35 MODIS; VPM; VPRM; 36 37 38 1. Introduction Terrestrial gross primary productivity (GPP) is the largest component flux of the global 39 carbon cycle, and is about 20 times greater than the amount of carbon from anthropogenic sources 40 (Canadell et al., 2007). Thus, even small fluctuations in GPP can cause large changes in the 41 airborne fraction of anthropogenic carbon and influence future climate warming scenarios 42 (Raupach et al., 2008). Terrestrial GPP also provides important societal services through provision 2 43 of food, fiber and energy. Regular monitoring of terrestrial GPP is therefore required to understand 44 and assess dynamics in the global carbon cycle, forecast future climate, and ensure long term 45 security in services provided by terrestrial ecosystems (Bunn and Goetz, 2006; Schimel, 2007). 46 Numerous of ecosystem models have been widely developed as a means of quantifying 47 spatial-temporal variations in GPP at large scales. However, different ecosystem models are 48 inconclusive regarding the magnitude and spatial distribution of GPP at the regional and global 49 scales. Recently, the model comparison, using standardized data from the North American Carbon 50 Program, showed none of the models consistently reproduce observed interannual variability 51 within measurement uncertainty because models can not represent the variability in spring 52 phenology, soil thaw and snowpack melting, and lagged response of ecosystems to extreme 53 climatic events (Keenan et al., 2012). Another study evaluated simulated daily average GPP from 54 26 models against estimated GPP at 39 eddy covariance flux tower sites across the United States 55 and Canada, and showed none of the models in this study match estimated GPP within the range 56 of uncertainty of observed fluxes, indicating the poor model performance (Schaefer et al., 2012). 57 These conclusions were supported by the previous comparison of 16 dynamic global vegetation 58 models that suggested the lowest estimation of global NPP (39.9 Pg C) by the Hybrid model was 59 approximately 50% smaller compared to what was estimated by the TURC model (80.5 Pg C) 60 (Cramer et al., 19999). 61 The light use efficiency (LUE) model based on the satellite data may have the most 62 potential to adequately address the spatial and temporal dynamics of GPP because of its 63 theoretical basis and practicality (Running et al., 2000). Independently and as a part of integrated 64 ecosystem models, the LUE approach has been used to estimate GPP and net primary production 3 65 (NPP) at various spatial and temporal scales (Potter et al., 1993; Prince and Goward, 1995; 66 Landsberg and Waring, 1997; Running et al., 2000; Xiao et al., 2004; Coops et al., 2005). 67 Numerous of studies have validated LUE models at regional and global scales in a variety of 68 major ecosystem types (Potter et al., 1993; Turner et al., 2006). 69 LUE models are often developed based on the unique assumptions driving by different 70 environmental variables, and formulate the processes controlling vegetation production in 71 different ways. Thus, there is diversity in both the complexity of the LUE model structure and 72 formulation though all of them follow the light use efficiency principle. Each model, therefore, is 73 a complex combination of scientific hypotheses and choices, and their estimates depend on these 74 inherent assumptions (Beer et al., 2010). Recent studies showed the large model uncertainties 75 within LUE models. For example, using satellite-based models, estimated GPP for North America 76 vary considerably between 12.2 and 18.7 Pg C yr-1 (Huntzinger et al., 2012). Available individual 77 model validations, however, are not sufficient to identify the sources of model differences and 78 shortcomings due to differences at validation datasets and driving variables. Therefore, in order to 79 move towards more robust estimates of vegetation production dynamics, it is necessary to first 80 compare estimates from a variety of LUE models, as well as evaluate estimates against consistent 81 and extensive measurements that are available (Running et al., 2004; Heinsch et al., 2006). 82 In this study, we evaluated how well seven different satellite-based models capture 83 spatio-temporal variations of GPP. The overarching goals of this study are to (1) examine the 84 model performance at the numerous of eddy covariance sites and (2) compare the temperature and 85 water response curves among the seven LUE models. 4 86 2. Model and data 87 2.1 Light use efficiency model 88 The LUE model is built on two fundamental assumptions (Running et al., 2004): (1) the 89 ecosystem GPP is directly related to absorbed photosynthetically active radiation (APAR) through 90 LUE, where LUE is defined as the amount of carbon produced per unit of APAR, and (2) realised 91 LUE may be reduced below its theoretical potential value by environmental stresses, such as low 92 temperatures or water shortages (Landsberg et al., 1986). The general form of the LUE model is: 93 GPP PAR fPAR LUEmax f (Ts ,Ws , ) (1) 94 where PAR is the incident photosynthetically active radiation (MJ m-2) per time period (e.g., day or 95 month), fPAR is the fraction of PAR absorbed by the vegetation canopy, LUEmax is the potential 96 LUE (g C m-2 MJ-1 APAR) without environment stress, f is a scalar varying from 0 to 1 that 97 represents the reduction of potential LUE under limiting environmental conditions, Ts and Ws are 98 temperature and water downward regulation scalars, and the multiplication of LUEmax and f is 99 realised LUE. 100 In this study, seven LUE models were selected to conduct the global comparison of 101 model performance, including CASA (Potter et al., 1993), CFix (Veroustraete et al., 2002), CFlux 102 (Turner et al., 2006; King et al., 2011), EC-LUE (Yuan et al., 2007, 2010), MODIS-GPP (Running 103 et al., 2000), VPM (Xiao et al., 2004) and VPRM model (Mahadevan et al., 2008). The detailed 104 model introduction and model operation can be found at Supplemental Online Material (SOM). 105 2.2 Data and method 106 LaThuile FLUXNET dataset was used in this study (http://www.fluxdata.org). Totally, 5 107 155 eddy covariance (EC) towers were included in this study, from six major terrestrial biomes: 108 evergreen broadleaf forest (EBF), deciduous broadleaf forest (DBF), mixed forest (MIF), 109 evergreen needleleaf forest (ENF), shrubland (SHR) and grassland (GRA) (Table S1; Figure S1). 110 Detailed information on data processing and site information (i.e. vegetation, climate and soils) 111 are available at the LaThuile FLUXNET Internet sites. 112 We examined the model performance using calibrated parameter values. Eighty percent 113 sites were selected to calibrate model parameters for each vegetation type, and other 20% sites 114 were used to validate models. This parameterization was repeated by 100 times, and the calibrated 115 parameter values were collected within Table 1. Annual simulated and observed GPP were 116 calculated in order to investigate the model performance on interannual variability of GPP. If 117 missing daily data was 20% of entire year data, the value of this year was indicated as missing. 118 For a site to be included for evaluating interannual variability, it had to have minimum of 3 years 119 of GPP observations and simulations. Based on this criterion, 100 sites consisting of 462 years 120 were included into the analysis (Table S1). We calculated the standard deviations of annual 121 averaged GPP observations and simulations for each site, and examined the correlations through 122 all 100 sites. Moreover, cloudiness index (CL), which was calculated by the ratio of PAR and 123 potential PAR, was used to indicate the friction of cloud cover. The days when CL is less than 0.3 124 were indicated as clear days, the CL ranges 0.3 to 0.6 for cloudy days and more than 0.6 were 125 indicated as overcast days. Similarly, water stress was separated into three levels (i.e. drought, 126 normal and wet conditions) based on the water stress scalars of seven models as the following 127 equations 6 Drought, Ws < Wsmin + 128 Normal , (Wsmin + { Wet Wsmax −Wsmin 3 Wsmax −Wsmin , Ws > Wsmin + ) < Ws < 3 2×(Wsmax −Wsmin ) (Wsmin + 2×(Wsmax −Wsmin ) 3 (2) 3 129 where Wsmin and Wsmax are the minimum and maximum values of Ws at each site. 130 <<Table 1>> 131 ) Two pairwise comparisons were conducted on the model components in order to 132 investigate the differences of model structure. First, we identified the impacts of fraction of PAR 133 absorbed by the vegetation canopy on GPP simulations by comparing the two correlations: 134 (a) the correlation of simulated GPP among seven models; 135 (b) the correlation of potential light energy use (PLUE) (i.e. PAR×fPAR×LUEmax); 136 Then, we diagnosed the primary environment variables by the second pairwise 137 138 139 140 comparison: (a) the correlation of realized light energy use only considering temperature stress (RLUEtem) (i.e. PAR×fPAR×LUEmax×Ts); (b) the correlation of realized light energy use only considering water stress (RLUEwater) 141 (i.e. PAR×fPAR×LUEmax×Ws). 142 2.3 Statistical analysis 143 The nonlinear regression procedure (Proc NLIN) in the Statistical Analysis System 144 (SAS, SAS Institute Inc., Cary, NC, USA) was applied to optimize the model parameters of seven 145 LUE models across the calibration sites. Four metrics were used to evaluate the performance of 146 the LUE models in this study, including coefficient of determination (R2), root mean square error 147 (RMSE), mean predictive error (PE, difference between mean observations and simulations), and 7 148 relative predictive error (RPE, the ratio between PE and mean observations). 149 3. Results and discussion 150 3.1 Comparison of model performance 151 All of seven LUE models showed the substantial difference of model performance 152 among various ecosystem types according to the R2, RMSE, PE and RPE (Figure S1). Over the 153 shrublands and evergreen broadleaf forests, almost all models showed obvious low performance 154 with low R2 and high RMSE. The highest model performance was observed over deciduous 155 broadleaf forests within seven LUE models. Parameter calibration significantly improved the 156 model performance over almost all ecosystem types (Figure 1; Table 1). Similarly, all of seven 157 calibrated models showed the highest model performance over the deciduous broadleaf forests, 158 intermediate over the evergreen needleleaf forests, mixed forests and grasslands, lowest at 159 shrublands and evergreen broadleaf forests (Figure 1). For a given vegetation type, models 160 performance differed among seven models. For example, CFlux and EC-LUE models were found 161 higher R2 and lower RMSE compared with other five models (Figure 1). 162 <<Figure 1>> 163 From the spatial scales, EC-LUE and CFlux models explained higher GPP variations 164 with determination coefficient of 0.55 and 0.44 respectively (Figure 2). All models appeared the 165 overestimation at the low GPP regions, and underestimation at the high GPP regions (Figure 2). 166 Moreover, we calculated the mean R2 and RMSE values of seven models at each site, and 167 compared the R2 and RMSE of individual model with means of seven models. On average, at 80% 168 and 75% sites, EC-LUE and CFlux models showed higher R2 compared with mean level of seven 8 169 models, while they showed the lower RMSE at 76% and 75% sites respectively (Figure 3). 170 <<Figure 2>> 171 <<Figure 3>> 172 Seven LUE models, expect CFlux model, significantly underestimated GPP at the 173 overcastting and cloudy days (Figure 4). For example, the averaged predictive errors of CASA 174 model was about -1.12 g C m-2 day-1 at overcastting days, however, the predictive errors was 0.15 175 g C m-2 day-1 at clear days. Previous of studies have found that increased fraction of diffuse 176 radiation at the cloudy days enhanced the plant photosynthesis (Gu et al., 2002, 2003; Urban et al., 177 2007; Alton et al., 2007). For example, Gu et al (2003) reported increase in diffuse radiation 178 because of volcanic aerosols alone enhanced noontime photosynthesis of a deciduous forest by 23% 179 in 1992 and 8% in 1993. This finding contributed to the temporary increase of terrestrial 180 ecosystems carbon sink after the eruption of Mount Pinatubo (15.1ºN, 121.4ºE) on 15 June 1991 181 (Ciais et al., 1995; Bousquet et al., 2000; Battle et al., 2000). Besides volcanic, cloud reduces the 182 global solar radiation but increases the relative proportion of diffuse radiation at the Earth surface 183 too. 184 <<Figure 4>> 185 It is a case that increased fraction of diffuse radiation can be the cause of changes in 186 many atmospheric factors such as temperature, moisture, and latent heating etc. These factors all 187 have direct or indirect influences on terrestrial ecosystem carbon dynamics (Gu et al., 1999). 188 Therefore some researchers emphasized decreases in the respiration of sunlit leaves due to 189 reduced leaf temperature and vapor pressure deficit (Baldocchi, 1997). However, direct impacts of 190 diffuse radiation had been found: (1) Diffuse radiation results in higher light use efficiencies by 9 191 plant canopies (Gu et al., 2002; Alton et al., 2007). (2) Diffuse radiation penetrates to lower depths 192 of the canopy more efficiently than does the direct radiation (Matsuda et al., 2004). This increased 193 the potential leaf area available for photosynthesis. (3) An increase of blue/red light ratio may lead 194 to higher photosynthesis rates per unit leaf area with increasing faction of diffuse radiation 195 (Matsuda et al., 2004). 196 Among the investigated seven LUE models, CFlux is only one model to integrate the 197 impacts of diffuse radiation on plant photosynthesis (Turner et al., 2006). CFlux model assumed a 198 maximum potential LUE at the overcast condition and minimum LUE value at the clear days, and 199 then a linear decreased trend with cloud cover (Turner et al., 2006). The results showed the simply 200 linear equation successfully reflected the impacts of diffuse radiation. The latest study developed a 201 two-leaf light use efficiency (TL-LUE) model based on the MOD17 algorithm, which separates 202 the canopy into sunlit and shaded leaf groups and calculates GPP separately for them with 203 different maximum light use efficiencies (He et al., 2013). The newly developed TL-LUE model 204 shows lower sensitivity to sky conditions than the MOD17 algorithm. 205 3.2 Comparison of interannual variability 206 The interannual variability of vegetation production exists at different scales from global, 207 regional to plot/stand levels and has been linked with the interannual variability of global carbon 208 balance as well as atmospheric CO2 concentration (Goulden et al., 1996; Bousquet et al., 2000; 209 Zhao et al., 2010). Previous study indicated interannual variability of the global vegetation 210 production is a major driver of the interannual CO2 growth rate (Zhao et al., 2010). Therefore, 211 understanding the cause and degree of interannual variability is important for both ecological 10 212 213 theory and global carbon cycle. In this study, the ability of simulating interannual variability was investigated at 100 214 sites with more than three-year observations. The results showed the poor ability of seven models 215 to identify the interannual variability. The correlation coefficient (R2) of standard deviation 216 between simulations and observations through all sites ranged from 0.14 to 0.54, and EC-LUE, 217 CFlux and CASA models showed the highest R2 (Figure 5). 218 <<Figure 5>> 219 Our result confirmed a previous study that modeling interannual variation in GPP has 220 proven challenging (Richardson et al., 2007). Keenan et al (2012) assessed the performance of 16 221 terrestrial biosphere models and 3 remote sensing products against long-term measurements of 222 biosphere-atmosphere CO2 exchange made with eddy-covariance flux towers at 11 forested sites 223 in North America. The results showed none of the models consistently reproduce observed 224 interannual variability within measurement uncertainty. Compared with the process-based models, 225 the remote sensing GPP products performed comparably to the average process-based model when 226 assessed against interannual variability (Keenan et al., 2012). Although the response of terrestrial 227 ecosystems to mean climatic drivers is relatively well captured, sensitivity to the impact of 228 variability in climatic drivers may not be, leading to the accumulation of high frequency model 229 error (Dietze et al., 2011) over longer time scales (Schwalm et al., 2010). Although estimates of 230 GPP based on remote sensing have been used to evaluate process-based models, results herein 231 suggest that estimates of interannual variability from both approaches are subject to similar 232 magnitudes of error (Poulter et al., 2011). 233 The possible causes of the errors for modeling interannual variability of GPP could be: 11 234 (1) Incomplete integration of environmental regulations. Most of LUE models only integrate the 235 impacts of temperature, water and radiation. Few LUE models consider the impacts of stand age, 236 phenology and CO2 fertilization on vegetation production, however, previous of studies have 237 showed the significant regulations of those environmental variables to vegetation production 238 (White et al., 1999; DeLucia and Thomas, 2000). (2) Limited understands on the key 239 physiological processes. Braswell et al. (1997) showed that climate induced physiological changes 240 are greater than the direct effect of climatic variability on the carbon cycle. Hui et al. (2003) used 241 a sum-of-squares approach to separate interannual variability in carbon cycle into four different 242 sources: functional change, interannual climatic variability, seasonal climatic variability and 243 random error, and seasonal climatic variability can explain mostly interannual variability in 244 carbon cycles. However, current LUE models do not integrated any responses of physiological 245 processes to environment changes. 246 3.3 Comparison of model structure 247 Pairwise comparison showed higher correlations of PLUE (i.e. PAR×fPAR×LUEmax) 248 among seven models compared with those of GPP simulations (Fig.6). For example, the 249 correlations of CASA model and other six models on PLUE ranged from 0.75 to 0.96, however, 250 correlations of GPP simulations were found from 0.37 to 0.43 (Fig.6). Generally, the pairwise 251 comparison of PLUE and GPP simulations can essentially indicate the contributions of fPAR and 252 environment regulation scalars to the differences of GPP simulations. The result implied larger 253 contribution of environment stress equations on model differences compared with the fraction of 254 PAR absorbed by the vegetation canopy. 12 255 256 <<Figure 6>> Within LUE models, the fraction of solar radiation intercepted by terrestrial vegetation 257 (fPAR) is calculated from remote sensing data, which is a critical variable for estimating vegetation 258 production. Various methods were developed for calculating fPAR among the LUE models. At the 259 CASA model, fPAR was calculated as a linear function of the simple ratio (SR), which was derived 260 from NDVI (Potter et al., 1993). CFlux and MODIS models directly utilized MODIS-fPAR 261 products. Other four models calculated fPAR based on vegetation index. CFix and EC-LUE 262 models used linear equation with NDVI to estimate fPAR (Veroustraete et al., 2002; Yuan et al., 263 2007), and VPM and VPRM directly used EVI to indicate fPAR (Xiao, et. al., 2004). CASA, CFix 264 and EC-LUE applied the equation with NDVI to calculate fPAR and showed the high consistence 265 through most sites (Fig.6). Low correlations were observed among MODIS-fPAR, EVI and 266 NDVI-based fPAR (Fig.6) 267 The second pairwise comparison indicated the larger impacts of water stress equation on 268 GPP simulations than that of temperature stress equation (Fig.7). On average, the correlations of 269 RLUEtem between two models were 0.80±0.08, while the mean value of correlations of RLUEwater 270 was 0.65±0.12. In generally, temperature and water response equations were the important two 271 down-regulation factors for LUE models. 272 <<Figure 7>> 273 We compared the consistence of water stress derived from seven models within sites. 274 Water stress scalars of seven models were separated into three levels: drought, normal and wet. 275 The results showed the more than 50% water stress levels were not consistent among models 276 (Table2). The largest inconsistence of water levels was found between MODIS and EC-LUE/CFix 13 277 with 65% differences (Table 2). Moreover, results also showed large friction of inconsistent 278 identifying drought and wet conditions. As shown at Table 2, on average among all models, more 279 than 15% days were wrongly identified between drought and wet days (Table 2). Comparison of 280 water stress equations among LUE models showed the substantial differences through the almost 281 all sites (Fig.8). Moisture availability of CASA showed the largest correlation with that of 282 EC-LUE model through the sites, and the weak relationship with that of MODIS-GPP. Among 283 other four models, moisture availability hardly showed significant relationship. For example, 284 EC-LUE models indicated the site with similar moisture availability, on the contrary, MODIS-GPP 285 product showed the large difference (Fig.8). The moisture response curve of VPM and VPRM is 286 unique and showed the different spatial distribution compare with those of other models. 287 <<Figure 8>> 288 Significant differences of model performance were found among seven LUE models at 289 the three water stress condition (i.e. drought, normal and wet) (Fig.9). Except CFlux, VPRM and 290 VPM, other four models showed low R2 at the drought days. For example, CFix model explained 291 about 50±25% of the variation of GPP estimated at the wet conditions averagely, however, only 292 explained 22±15% variation at the drought days (Fig. 9). Moreover, no consistent predictive errors 293 were found under the different water stresses among the various models. CASA model tended to 294 underestimate GPP at the drought days, on the contrary, CFix model showed obvious 295 overestimation of GPP (Fig. 9). 296 <<Figure 9>> 297 298 Defining a function for quantifying the control of moisture availability on plant photosynthesis has long been a challenge. The effects of water on plant photosynthesis have been 14 299 estimated as a function of soil moisture, evapotranspiration friction and water vapor pressure 300 deficit (VPD) in a number of LUE models (Field et al., 1995; Prince and Goward, 1995; Running 301 et al., 2000). For instance, in the EC-LUE model, water stress was estimated using the ratio of 302 actual evapotranspiration to net shortwave radiation energy. This ratio was considered to be a very 303 good indicator of soil or vegetation moisture conditions because decreasing amounts of energy 304 partitioned into latent heat flux suggests a stronger moisture limitation (Kurc and Small, 2004; 305 Zhang et al., 2004; Suleiman and Crago, 2004). Other of models, such as VPM and VPRM, used a 306 satellite-derived water index (Land Surface Water Index) to estimate the seasonal dynamics of 307 water stress (Xiao et al., 2004). 308 These variables, which have been used into LUE models, had their weaknesses. For 309 example, it is difficult to characterize soil moisture conditions over large areas from either 310 modeling or remote sensing. This limits the predictive power of any spatial GPP model that relies 311 on soil moisture. VPD is not a good indicator of the spatial heterogeneity of soil moisture 312 conditions across the landscape (e.g., slope versus valley) and it is not likely to be linearly related 313 to soil water availability for which it is often used as a proxy. Moreover, evapotranspiration 314 friction needs an ET model for simulating ecosystem evapotranspiration, and any uncertainties 315 within ET models will reduce the model performance of LUE models. 316 4. Summary 317 We evaluated seven satellite-driven light use efficiency models against 155 eddy 318 covariance sites globally including six major biomes. All seven models showed similar model 319 performance over the vegetation types. The best model performance was observed at deciduous 15 320 broadleaf forests and mixed forests, intermediate at grasslands and evergreen needleleaf forests, 321 lowest at evergreen broadleaf forests. From the spatial respective, CFlux and EC-LUE models 322 showed higher correlations between site-averaged GPP observations and simulations, and were 323 represented with higher performance to simulate interannual variability of GPP. fPAR, temperature 324 curves and water stress equations largely differed among the seven LUE models. Comparably, 325 water stress equations differed largestly which was the major cause for GPP simulations 326 difference. 327 328 329 Acknowledgments This study was supported by the National Natural Science Foundation of China 330 (41201078), the National High Technology Research and Development Program of China (863 331 Program) (2013AA122003), Program for New Century Excellent Talents in University 332 (NCET-12-0060) and the Fundamental Research Funds for the Central Universities. 333 334 References 335 Alton, P. B., North, P. R. & Los, S. O. (2007). The impact of diffuse sunlight on canopy light-use 336 efficiency, gross photosynthetic product and net ecosystem exchange in three forest biomes. 337 Global Change Biology, 14(3), 776–787. 338 Baldocchi, D. (1997). Measuring and modeling carbon dioxide and water vapor exchange over a 339 temperate broad-leaved forest during the 1995 summer drought. Plant, Cell & Environment. 340 20(9), 1108–1122. 16 341 Battle, M., Bender, M. L., Tans, P. P., White, J. W. C., Ellis, J. T., Conway, T., et al (2000). Global 342 carbon sinks and their variability inferred from atmospheric O2 and δ13C. Science, 287, 343 2467–2470. 344 Baldocchi, D., Falge, E., Gu, L., Olson, R., Hollinger, D., Running, S., et al. (2001). FLUXNET: a 345 new tool to study the temporal and spatial variability of ecosystem-scale carbon dioxide, water 346 vapor and energy flux densities. Bulletin of the American Meteorological Society, 82(11), 347 2415–2435. 348 Bunn, A. G., & Goetz, S. J. (2006). Trends in satellite-observed circumpolar photosynthetic 349 activity from 1982 to 2003: the influence of seasonality, cover type, and vegetation density. 350 Earth Interactions, 10(12), 1–19. 351 Bousquet, P., Peylin, P., Ciais, P., Le Quéré, C., Friedlingstein, P. & Tans, P. P. (2000). Regional 352 changes in carbon dioxide fluxes of land and ocean since 1980. Science, 290, 1342–1345. 353 Beer, C., Reichstein, M., Tomelleri, E., Ciais, P., Jung, M., Carvalhais, N., et al. (2010). Terrestrial 354 gross carbon dioxide uptake: global distribution and covariation with climate. Science, 329, 355 834–838. 356 357 358 Braswell, B. H., Schimel, D. S., Linder, E. & Moore, B. (1997). The response of global terrestrial ecosystems to interannual temperature variability. Science, 278, 870–872. Cramer, W., Bondeau, A., Moore, B., Churkina, C., Nemry, B., Ruimy, A., et al. (1999).Comparing 359 global models of terrestrial net primary productivity (NPP): overview and key results. Global 360 Change Biology, 5(S1), 1−15. 361 362 Chen, W., Chen, J. M., Price D. T. & Cihlar J. (2002). Effects of stand age on net primary productivity of boreal black spruce forests in Ontario, Canada. Canadian Journal for Forest 17 363 364 Research, 32(5), 833–842. Canadell, J. G., Kirschbaum, M. U. F., Kurz, W. A., Sanz, M. J., Schlamadinger, B., & Yamagata, 365 Y. (2007). Factoring out natural and indirect human effects on terrestrial carbon sources and 366 sinks. Environmental Science & Policy, 10(4), 370–384. 367 368 369 Chapin III, F. S., Matson, P. A., Mooney, H. A. (2012). Principles of terrestrial ecosystem ecology. Springer, 2nd edition. Ciais, P., P. P. Tans, M. Trolier, J. W. C. White, & R. J. Francey. (1995). A large northern 370 hemisphere terrestrial CO2 sink indicated by the 13C/12C ratio of atmospheric CO2, Science, 371 269, 1098–1102. 372 Coops, N. C., Waring, R. H., Law, B. E. (2005). Assessing the past and future distribution and 373 productivity of ponderosa pine in the Pacific Northwest using a process model, 3-PG. 374 Ecological Modelling, 183(1), 107–124. 375 376 377 DeLucia, E. & Thomas, R. (2000). Photosynthetic responses to CO2 enrichment of four hardwood species in a forest understory. Oecologia, 122(1), 11–19. Dietze, M. C., Vargas, R., Richardson, A. D., Stoy, P. C., Barr, A. G., Anderson, R. S., et al. (2011). 378 Characterizing the performance of ecosystem models across time scales: a spectral analysis of 379 the North American Carbon Program site-level synthesis. Journal of Geophysical Research: 380 Biogeosciences, 116(G4), 2005–2012. 381 Field, C. B., Jackson, R. B., & Mooney, H. A. (1995). Stomatal responses to increased CO2: 382 implications from the plant to the global scale. Plant, Cell & Environment, 18(10), 1214–1225. 383 Friedl, M. A., Sulla-Menashe, D., Tan, B., Schneider, A., Ramankutty, N., Sibley, A., et al. (2010). 384 MODIS Collection 5 global land cover: algorithm refinements and characterization of new 18 385 386 datasets. Remote Sensing of Environment, 114(1), 168–182. Gu, L., Baldocchi, D., Verma, S. B., Black, T. A., Vesala, T., Falge, E. M., et al. (2002). 387 Advantages of diffuse radiation for terrestrial ecosystem productivity. Journal of geophysical 388 research: Atmospheres, 107(D6), ACL 2-1–ACL 2-23. 389 Gu, L. H., Baldocchi, D., Wofsy, S. C., Munger, J. W., Michalsky, J. J., Urbanski, S. P., et al. 390 (2003). Response of a deciduous forest to the mount Pinatubo eruption: enhanced 391 photosynthesis. Science, 299, 2035–2038. 392 Gu, L. H., Fuentes, J. D., Shugart, H. H., Staebler, R. M., & Black, T. A. (1999). Responses of net 393 ecosystem exchanges of carbon dioxide to changes in cloudiness: results from two North 394 American deciduous forests. Journal of Geophysical Research: Atmospheres, 104(D24), 395 31421–31434. 396 Goulden, M. L., Munger, J. W., Fan, S. M., Daube, B. C., & Wofsy, S.C. (1996). Exchange of 397 carbon dioxide by a deciduous forest: response to interannual climate variability. Science, 271, 398 1576–1578. 399 He MZ, Ju WM, Zhou YL, Chen JM, He HL, Wang SQ, Wang HM, Guan DX, Yan JH, Li YN, 400 Hao YB, Zhao FH. (2013). Development of a two-leaf light use efficiency model for 401 improving the calculation of terrestrial gross primary productivity. Agricultural and Forest 402 Meteorology, 173, 28-39. 403 404 405 406 Hui, D., Luo, Y., & Katul, G. (2003). Partitioning interannual variability in net ecosystem exchange between climatic variability and functional change. Tree physiology, 23(7), 433–442. Huntzinger, D. N., Post, W. M., Wei, Y., Michalak, A. M., West, T. O., Jacobson, A. R., et al. (2012). North American Carbon Program (NACP) regional interim synthesis: terrestrial 19 407 408 biospheric model intercomparison. Ecological Modelling, 232, 144–157. Heinsch, F. A., Zhao, M., Running, S. W., Kimball, J. S., Nemani, R. R., Davis, et al. (2006). 409 Evaluation of remote sensing based terrestrial productivity from MODIS using regional tower 410 eddy flux network observations. IEEE Transactions on Geoscience and Remote Sensing, 44(7), 411 1908–1925. 412 Keenan, T. F., Baker, I., Barr, A., Ciais, P., Davis, K., Dietze, M., et al. (2012). Terrestrial 413 biosphere model performance for inter-annual variability of land-atmosphere CO2 exchange. 414 Global Change Biology, 18(6), 1971-1987. 415 Knapp, A. K., Fay, P. A., Blair, J. M., Collins, S. L., Smith, M. D., Carlisle, J. D., et al. (2002). 416 Rainfall variability, carbon cycling, and plant species diversity in a mesic grassland. Science, 417 298, 2202–2205. 418 Kositsup, B., Montpied, P., Kasemsap, P., Thaler, P., Améglio, T., & Dreyer, E. (2009). 419 Photosynthetic capacity and temperature responses of photosynthesis of rubber trees (Hevea 420 brasiliensis Müll. Arg.) acclimate to changes in ambient temperatures. Trees, 23(2), 357–365. 421 Kurc, S. A., & Small, E. E. (2004). Dynamics of evapotranspiration in semiarid grassland and 422 shrubland ecosystems during the summer monsoon season, central New Mexico. Water 423 Resources Research, 40(9), 305, doi:10.1029/2004WR003068. 424 425 426 427 428 King, D. A., Turner, D. P., & Ritts, W. D. (2011). Parameterization of a diagnostic carbon cycle model for continental scale application. Remote Sensing of Environment, 115(7), 1653-1664. Landsberg, J.J. (1986). Physiological ecology of forest production. Academic Press, London, 165-178. Landsberg, J. J., & Sands, P. (2010). Physiological ecology of forest production: principles, 20 429 processes and models (Vol. 4). Academic Press, London, 165–178. 430 Landsberg, J. J., & Waring, R. H. (1997). A generalised model of forest productivity using 431 simplified concepts of radiation-use efficiency, carbon balance and partitioning. Forest 432 Ecology and Management, 95(3), 209–228. 433 Melillo, J. M., Kicklighter, D. W., McGuire, A. D., Peterjohn, W. T., & Newkirk, K. M. (1995). 434 Global change and its effects on soil organic carbon stocks. In Dahlem Conference 435 Proceedings. (pp. 175–189). New York: John Wiley and Sons. 436 Matsuda, R., Ohashi-Kaneko, K., Fujiwara, K., Goto, E., & Kurata, K. (2004). Photosynthetic 437 characteristics of rice leaves grown under red light with or without supplemental blue light. 438 Plant & cell physiology, 45(12), 1870-1874. 439 Mahadevan, P., Wofsy, S. C., Matross, D. M., Xiao, X. M., Dunn, A. L., Lin, J. C., et al. (2008). A 440 satellite-based biosphere parameterization for net ecosystem CO2 exchange: Vegetation 441 Photosynthesis and Respiration Model (VPRM). Global Biogeochemical Cycles, 22(2), 1–17. 442 Poulter, B., Ciais, P., Hodson, E., Lischke, H., Maignan, F., Plummer, S., et al. (2011). Plant 443 functional type mapping for earth system models. Geoscientific Model Development, 4(4), 444 993–1010. 445 446 447 Prince, S. D., & Goward, S. N. (1995). Global primary production: a remote sensing approach. Journal of biogeography, 22(4/5), 815–835. Potter, C. S., Randerson, J. T., Field, C. B., Matson, P. A., Vitousek, P. M., Mooney, H. A., et al. 448 (1993). Terrestrial ecosystem production: a process model based on global satellite and surface 449 data. Global Biogeochemical Cycles, 7(4), 811–841. 450 Qi, Y., Xu, M., & Wu, J., (2002). Temperature sensitivity of soil respiration and its effects on 21 451 452 ecosystem carbon budget: nonlinearity begets surprise. Ecological Modelling. 153, 131–142. Raupach, M. R., Canadell, J. G., & Le Quéré, C. (2008). Anthropogenic and biophysical 453 contributions to increasing atmospheric CO2 growth rate and airborne fraction. Biogeosciences 454 Discuss, 5(4), 2867–2896. 455 Richardson, A., Hollinger, D., Aber, J., Ollinger, S., and Braswell, B. (2007). Environmental 456 variation is directly responsible for short but not long term variation in forest atmosphere 457 carbon exchange. Global Change Biol., 13, 788-803. 458 Rossini, M., Cogliati, S., Meroni, M., Migliavacca, M., Galvagno, M., Busetto, L., et al. (2012). 459 Remote sensing-based estimation of gross primary production in a subalpine grassland. 460 Biogeosciences, 9(7), 2565–2584. 461 Ruimy, A., Kergoat, L., & Bondeau, A. (1999). Comparing global models of terrestrial net primary 462 productivity (NPP): analysis of differences in light absorption and light-use efficiency. Global 463 Change Biology, 5(S1), 56–64. 464 Running, S. W., Nemani, R. R., Heinsch, F. A., Zhao, M., Reeves, M., & Hashimoto, H. (2004). A 465 continuous satellite-derived measure of global terrestrial primary production. Bioscience, 466 54(6), 547–560. 467 Ruimy, A., Saugier, B., & Dedieu, G. (1994). Methodology for the estimation of terrestrial net 468 primary production from remotely sensed data. Journal of Geophysical Research: 469 Atmospheres, 99(D3), 5263–5283. 470 Running, S. W., Thornton, P. E., Nemani, R., & Glassy, J. M. (2000). Global terrestrial gross and 471 net primary productivity from the Earth Observing System. Methods in ecosystem science, 472 44–57. 22 473 474 475 476 Schimel, D. (2007). Carbon cycle conundrums. Proceedings of the National Academy of Sciences, 104(47), 18353–18354. Suleiman, A., Crago, R., (2004). Hourly and daytime evapotranspiration from grassland using radiometric surface temperatures. Agronomy Journal. 96, 384–390. 477 Schaefer, K., Schwalm, C. R., Williams, C., Arain, M. A., Barr, A., Chen, J. M., et al. (2012). A 478 model-data comparison of gross primary productivity: Results from the North American 479 Carbon Program site synthesis. Journal of Geophysical Research: Biogeosciences, 117(G3), 480 1–15. 481 Schwalm, C. R., Williams, C. A., Schaefer, K., Arneth, A., Bonal, D., Buchmann, N., et al. (2010). 482 Assimilation exceeds respiration sensitivity to drought: A FLUXNET synthesis. Global 483 Change Biology, 16(2), 657–670. 484 Turner, D. P., Ritts, W. D., Styles, J. M., Yang, Z., Cohen, W. B., Law, B. E., et al. (2006). A 485 diagnostic carbon flux model to monitor the effects of disturbance and interannual variation in 486 climate on regional NEP. Tellus B, 58(5), 476−490. 487 Turner, D. P., Urbanski, S., Bremer, D., Wofsy, S. C., Meyers, T., Gower, S. T., et al. (2003). A 488 cross-biome comparison of daily light use efficiency for gross primary production. Global 489 Change Biology, 9(3), 383–395. 490 Urban, O., JANOUŠ, D., Acosta, M., CZERNÝ, R., MarkovA, I., NavrATil, M.,et al. (2007). 491 Ecophysiological controls over the net ecosystem exchange of mountain spruce stand: 492 comparison of the response in direct vs. diffuse solar radiation. Global Change Biology, 13(1), 493 157–168. 494 Veroustraete, F., Sabbe, H., & Eerens, H. (2002). Estimation of carbon mass fluxes over Europe 23 495 496 using the C-Fix model and Euroflux data. Remote Sensing of Environment. 83(3), 376–399. Viña, A., & Gitelson, A. A. (2005). New developments in the remote estimation of the fraction of 497 absorbed photosynthetically active radiationin crops, Geophysical Research Letters, 32(17), 498 L17403, doi:10.1029/2005GL023647. 499 Walter-Shea, E. A., Privette, J., Cornell, D., Mesarch, M. A., & Hays, C. J. (1997). Relations 500 between directional spectral vegetation indices and leaf area and absorbed radiation in alfalfa. 501 Remote Sensing of Environment, 61(1), 162–177. 502 White, M. A., Running, S. W. & Thornton, P. E. (1999). The impact of growing-season length 503 variability on carbon assimilation and evapotranspiration over 88 years in the eastern US 504 deciduous forest. International Journal of Biometeorology, 42(3), 139–145. 505 Xiao, X. M., Zhang, Q. Y., Braswell, B., Urbanski, S., Boles, S., Wofsy, S., et al. (2004). Modeling 506 gross primary production of temperate deciduous broadleaf forest using satellite images and 507 climate data. Remote Sensing of Environment. 91(2), 256–270. 508 Yuan, W. P., Liu, S. G., Yu, G. R., Bonnefond, J. M., Chen, J. Q., Davis, K., et al. (2010). Global 509 estimates of evapotranspiration and gross primary production based on MODIS and global 510 meteorology data. Remote Sensing of Environment. 114(7), 1416–1431. 511 Yuan, W. P., Liu, S. G., Zhou, G. S., Zhou, G. Y., Tieszen, L. L., Baldocchi, D., et al. (2007). 512 Deriving a light use efficiency model from eddy covariance flux data for predicting daily gross 513 primary production across biomes. Agricultural and Forest Meteorology, 143(3), 189–207. 514 Zhang, Y. Q., Liu, C. M., Yu, Q., Shen, Y. J., Kendy, E., Kondoh, A., et al. (2004). Energy fluxes 515 and the Priestley-Taylor parameter over winter wheat and maize in the North China Plain. 516 Hydrological Processes, 18(12), 2235–2246. 24 517 518 Zhao, M., & Running, S. W. (2010). Drought-induced reduction in global terrestrial net primary production from 2000 through 2009. Science, 329, 940–943. 25 519 Table 1 Calibrated model parameter values for seven models at the two parameterization 520 schemes. Vegetation Type Parameter CSH DBF EBF ENF GRA MIF CASA ɛ0 0.62±0.20 1.22±0.43 0.87±0.15 0.85±0.18 0.78±0.17 1.04±0.36 0.55±0.26 1.27±0.07 1.33±0.15 1.09±0.06 1.14±0.09 1.16±0.13 ɛmax 1.12±0.28 3.07±0.25 3.02±0.18 2.29±0.12 2.53±0.15 2.53±0.25 ɛcs 0.66±0.05 1.17±0.07 1.12±0.10 0.95±0.05 1.08±0.08 1.05±0.13 Tmin -13.76±6.54 -4.35±2.13 -14.46±4.35 -14.20±2.40 -20.12±7.06 -14.26±4.17 Tmax 13.04±8.79 13.08±3.23 20.00±0.00 8.25±1.82 9.55±5.87 13.79±3.74 VPDmin 0.33±0.34 0.11±0.01 0.12±0.07 0.11±0.00 0.12±0.01 0.12±0.02 VPDmax 3.42±0.58 2.99±0.32 2.56±0.32 2.79±0.13 3.23±0.47 2.44±0.32 1.28±0.40 1.71±0.19 1.70±0.11 1.85±0.20 1.59±0.41 1.72±0.31 ɛ0 0.66±0.28 1.77±0.19 1.68±0.10 1.36±0.08 1.52±0.16 1.64±0.22 Tmin -13.76±6.54 -4.35±2.13 -14.46±4.35 -14.20±2.40 -20.12±7.06 -14.26±4.17 Tmax 13.04±8.79 13.08±3.23 20.00±0.00 8.25±1.82 9.55±5.87 13.79±3.74 VPDmin 0.33±0.34 0.11±0.01 0.12±0.07 0.11±0.02 0.12±0.01 0.12±0.02 VPDmax 3.42±0.58 2.99±0.32 2.56±0.32 2.79±0.13 3.23±0.47 2.44±0.32 1.25±0.43 2.11±0.11 2.17±0.16 2.17±0.10 1.92±0.12 2.03±0.24 ɛ0 4.42±1.88 8.63±1.14 10.88±2.17 14.89±2.10 7.87±1.08 10.16±3.04 PAR0 4.47±1.22 3.15±0.44 2.37±0.75 1.61±0.24 3.07±0.52 2.50±0.81 CFix ɛ0 CFlux EC-LUE ɛ0 MODIS VPM ɛ0 VPRM 521 522 523 CSH, DBF, EBF, ENF, GRA, MIF: calibrated parameter values within shrubland, deciduous broadleaf forest, evergreen broadleaf forest, evergreen needleleaf forest, grassland, mixed forest respectively. Parameters were introduced at the Supplemental Online Material. 26 Table 2 Comparison of water stress levels derived from seven LUE models CASA CFix/EC-LUE CFlux MODIS VPRM/VPM - 0.49±0.12 0.15±0.28 0.57±0.15 0.56±0.13 CFix/EC-LUE 0.09±0.04 - 0.52±0.12 0.65±0.11 0.58±0.11 CFlux 0.12±0.05 0.06±0.06 - 0.50±0.15 0.54±0.12 MODIS 0.16±0.09 0.19±0.09 0.18±0.09 - 0.61±0.12 VPRM/VPM 0.16±0.10 0.22±0.14 0.13±0.09 0.17±0.10 - CASA 27 524 Figure caption 525 Figure 1 Model performance of seven Light Use Efficiency Models with calibrated parameters at 526 various vegetation types. 527 Figure 2 Observed vs. the simulated GPP over the 155 EC sites with calibrated parameters. The 528 long dash line is 1:1 line and the solid line is linear regression line. 529 Figure 3 Percentage of eddy covariance sites with higher domination coefficient and lower RMSE 530 for individual model compared the mean values of seven models. 531 Figure 4 The model performance at clear, cloudy and overcast days for seven models. 532 Figure 5 Correlation between standard deviations of simulated interannual variability of GPP. 533 Figure 6 Pairwise comparisons of correlations of GPP simulations and potential light energy use 534 (PLUE) among seven LUE models. 535 Figure 7 Pairwise comparisons of correlations of realized light energy use only considering 536 temperature stress (RLUEtem) and water stress (RLUEwater) among seven LUE modes. 537 Figure 8 Comparison of water stress curves among seven LUE models. 538 Figure 9 The model performance at drought, normal and wet days for seven models. 539 28 540 541 Figure 1 Model performance of seven Light Use Efficiency Models with calibrated parameters at 542 various vegetation types. 29 543 544 Figure 2 Observed vs. the simulated GPP over the 155 EC sites with calibrated parameters. The 545 long dash line is 1:1 line and the solid line is linear regression line. 30 546 547 Figure 3 Percentage of eddy covariance sites with higher domination coefficient and lower RMSE 548 for individual model compared the mean values of seven models. 31 549 550 Figure 4 The model performance at clear, cloudy and overcast days for seven models. 32 551 552 Figure 5 Correlation between standard deviations of simulated interannual variability of GPP. 33 553 554 Figure 6 Pairwise comparisons of correlations of GPP simulations and potential light energy use 555 (PLUE) among seven LUE models. 34 556 557 Figure 7 Pairwise comparisons of correlations of realized light energy use only considering 558 temperature stress (RLUEtem) and water stress (RLUEwater) among seven LUE modes. 35 559 560 Figure 8 Comparison of water stress curves among seven LUE models. 36 561 562 Figure 9 The model performance at drought, normal and wet days for seven models. 37