Influence of physiological phenology on the seasonal pattern of ecosystem respiration in deciduous forests

Understanding the environmental and biotic drivers of respiration at the ecosystem level is a prerequisite to further improve scenarios of the global carbon cycle. In this study we investigated the relevance of physiological phenology, defined as seasonal changes in plant physiological properties, for explaining the temporal dynamics of ecosystem respiration (R_ECO) in deciduous forests. Previous studies showed that empirical R_ECO models can be substantially improved by considering the biotic dependency of R_ECO on the short‐term productivity (e.g., daily gross primary production, GPP) in addition to the well‐known environmental controls of temperature and water availability. Here, we use a model‐data integration approach to investigate the added value of physiological phenology, represented by the first temporal derivative of GPP, or alternatively of the fraction of absorbed photosynthetically active radiation, for modeling R_ECO at 19 deciduous broadleaved forests in the FLUXNET La Thuile database. The new data‐oriented semiempirical model leads to an 8% decrease in root mean square error (RMSE) and a 6% increase in the modeling efficiency (EF) of modeled R_ECO when compared to a version of the model that does not consider the physiological phenology. The reduction of the model‐observation bias occurred mainly at the monthly time scale, and in spring and summer, while a smaller reduction was observed at the annual time scale. The proposed approach did not improve the model performance at several sites, and we identified as potential causes the plant canopy heterogeneity and the use of air temperature as a driver of ecosystem respiration instead of soil temperature. However, in the majority of sites the model‐error remained unchanged regardless of the driving temperature. Overall, our results point toward the potential for improving current approaches for modeling R_ECO in deciduous forests by including the phenological cycle of the canopy.     

Latitudinal patterns of magnitude and interannual variability in net ecosystem exchange regulated by biological and environmental variables

Over the last two and half decades, strong evidence showed that the terrestrial ecosystems are acting as a net sink for atmospheric carbon. However the spatial and temporal patterns of variation in the sink are not well known. In this study, we examined latitudinal patterns of interannual variability (IAV) in net ecosystem exchange (NEE) of CO₂ based on 163 site-years of eddy covariance data, from 39 northern-hemisphere research sites located at latitudes ranging from ∼29°N to ∼64°N. We computed the standard deviation of annual NEE integrals at individual sites to represent absolute interannual variability (AIAV), and the corresponding coefficient of variation as a measure of relative interannual variability (RIAV). Our results showed decreased trends of annual NEE with increasing latitude for both deciduous broadleaf forests and evergreen needleleaf forests. Gross primary production (GPP) explained a significant proportion of the spatial variation of NEE across evergreen needleleaf forests, whereas, across deciduous broadleaf forests, it is ecosystem respiration (Re). In addition, AIAV in GPP and Re increased significantly with latitude in deciduous broadleaf forests, but AIAV in GPP decreased significantly with latitude in evergreen needleleaf forests. Furthermore, RIAV in NEE, GPP, and Re appeared to increase significantly with latitude in deciduous broadleaf forests, but not in evergreen needleleaf forests. Correlation analyses showed air temperature was the primary environmental factor that determined RIAV of NEE in deciduous broadleaf forest across the North American sites, and none of the chosen climatic factors could explain RIAV of NEE in evergreen needleleaf forests. Mean annual NEE significantly increased with latitude in grasslands. Precipitation was dominant environmental factor for the spatial variation of magnitude and IAV in GPP and Re in grasslands.     

An analysis of soil respiration across northern hemisphere temperate ecosystems

Over two-thirds of terrestrial carbon is stored belowground and a significant amount of atmospheric CO₂ is respired by roots and microbes in soils. For this analysis, soil respiration (Rs) data were assembled from 31 AmeriFlux and CarboEurope sites representing deciduous broadleaf, evergreen needleleaf, grasslands, mixed deciduous/evergreen and woodland/savanna ecosystem types. Lowest to highest rates of soil respiration averaged over the growing season were grassland and woodland/savanna < deciduous broadleaf forests < evergreen needleleaf, mixed deciduous/evergreen forests with growing season soil respiration significantly different between forested and non-forested biomes (p < 0.001). Timing of peak respiration rates during the growing season varied from March/April in grasslands to July–September for all other biomes. Biomes with overall strongest relationship between soil respiration and soil temperature were from the deciduous and mixed forests (R² ≥ 0.65). Maximum soil respiration was weakly related to maximum fine root biomass (R² = 0.28) and positively related to the previous years’ annual litterfall (R² = 0.46). Published rates of annual soil respiration were linearly related to LAI and fine root carbon (R² = 0.48, 0.47), as well as net primary production (NPP) (R² = 0.44). At 10 sites, maximum growing season Rs was weakly correlated with annual GPP estimated from eddy covariance towersites (R² = 0.29; p < 0.05), and annual soil respiration and total growing season Rs were not correlated with annual GPP (p > 0.1). Yet, previous studies indicate correlations on shorter time scales within site (e.g., weekly, monthly). Estimates of annual GPP from the Biome-BGC model were strongly correlated with observed annual estimates of soil respiration for six sites (R² = 0.84; p < 0.01). Correlations from observations of Rs with NPP, LAI, fine root biomass and litterfall relate above and belowground inputs to labile pools that are available for decomposition. Our results suggest that simple empirical relationships with temperature and/or moisture that may be robust at individual sites may not be adequate to characterize soil CO₂ effluxes across space and time, agreeing with other multi-site studies. Information is needed on the timing and phenological controls of substrate availability (e.g., fine roots, LAI) and inputs (e.g., root turnover, litterfall) to improve our ability to accurately quantify the relationships between soil CO₂ effluxes and carbon substrate storage.For this study, these authors received significant contributions from: M. Aubinet, D. Baldocchi, C. Bernhofer, P. Bolstad, A. Bosc, J.L. Campbell, Y. Cheng, J. Curiel Yuste, P. Curtis, E.A. Davidson, D. Epron, A. Granier, T. Grünwald, D. Hollinger, I.A. Janssens, B. Longdoz, D. Loustau, J. Martin, R. Monson, W. Oechel, J. Pippen, F. Ponti, R. Ryel, K. Savage, L. Scott-Denton, J.-A. Subke, J. Tang, J. Tenhunen, V. Turcu, C. S. Vogel.     

Functional changes in the control of carbon fluxes after 3 years of increased drought in a Mediterranean evergreen forest?

Our objective was to test how a long‐term increased water limitation affects structural and functional properties of a Mediterranean ecosystem, and how these changes modify the response of the main carbon fluxes to climatic controls. In 2003, a 27% throughfall exclusion experiment was installed in a Quercus ilex L. forest in France. Gross primary production (GPP), ecosystem respiration (R_ECO) and net ecosystem exchange (NEE) were estimated in a control and a dry treatment. Decreasing throughfall decreased GPP by 14% and had a smaller effect on R_ECO (?12%), especially soil respiration R_S (?11%). Interannual variability of GPP (29%) was higher than for R_ECO (12%). Error propagation was used to estimates uncertainties in the NEE fluxes, which ranged from 3% to 10% in the control treatment but up to 167% for NEE in the dry treatment because more steps and data types were involved in the scaling. After 3 years of throughfall exclusion, we found no acclimation of R_S to climatic drivers. Functional properties of the response of R_S to soil water, temperature and rain pulse remained similar in the control and the dry treatments. A diurnal clockwise hysteresis in R_S was probably controlled by canopy photosynthesis with a 3 h lag. The proportion of diurnal variation of respiration due to photosynthesis was similar in all treatments (4–5%). Because of the characteristic of rain in Mediterranean climates, a continuous decrease of water input in these environments have an effect on topsoil water and consequently on R_S only during short periods when rainfall is characterized by infrequent and small events that does not allow the topsoil to reach field capacity and does not allow to dry completely. However, in the longer term, we expect a stronger decrease in R_S in the dry treatment driven by the decrease in GPP.     

新疆喀纳斯国家自然保护区植被叶面积指数观测与遥感估算

叶面积指数(Leaf Area Index,LAI)是重要的植被结构参数,调控着植被与大气之间的物质与能量交换,在生态环境脆弱的我国西北部开展植被LAI的研究对阐明该地区植被对气候变化和人类活动的响应特征具有重要的科学意义.利用LAI-2200和TRAC仪器观测了新疆喀纳斯国家级自然保护区森林和草地的有效叶面积指数(LAIe)和真实LAI,构建了其遥感估算模型,生成了研究区LAIe和LAI的空间分布图.在此基础上,分析了LAI随地形因子(海拔、坡度、坡向)的变化特征,探讨了将其应用于估算研究区森林生物量密度的可行性,并评估了研究区MODIS LAI产品的精度.结果表明:研究区阔叶林、针阔混交林、针叶林、草地LAIe的平均值分别为4.40、3.18、2.57、1.76,LAI的平均值分别为4.76、3.93、3.27、2.30.LAIe和LAI的高值主要集中分布在湖泊和河流附近;植被LAI随海拔、坡度和坡向的变化表现出明显的垂直地带性的特点.LAI随海拔和坡度的增加呈现先增加后减小的变化趋势,坡向对针叶林和草地LAI的影响明显,但对阔叶林和针阔混交林LAI的影响较弱;森林生物量密度(BD)随LAI增加而线性增加(BD=44.396LAI-25.946,R2=0.83),研究区森林生物量密度平均值为120.3 t/hm2,估算的总生物量为5.0×l06 t;MODIS LAI产品与利用TM数据生成的LAI之间具有一定的相似性(森林R2=0.42,草地R2=0.53),但森林和草地的MODIS LAI产品分别比利用TM数据生成的LAI偏低16.5％和24.4％.     

The response of soil respiration to precipitation change is asymmetric and differs between grasslands and forests

Intensification of the Earth's hydrological cycle amplifies the interannual variability of precipitation, which will significantly impact the terrestrial carbon (C) cycle. However, it is still unknown whether previously observed relationship between soil respiration (R_s) and precipitation remains applicable under extreme precipitation change. By analyzing the observations from a much larger dataset of field experiments (248 published papers including 151 grassland studies and 97 forest studies) across a wider range of precipitation manipulation than previous studies, we found that the relationship of R_s response with precipitation change was highly nonlinear or asymmetric, and differed significantly between grasslands and forests, between moderate and extreme precipitation changes. Response of R_s to precipitation change was negatively asymmetric (concave‐down) in grasslands, and double‐asymmetric in forests with a positive asymmetry (concave‐up) under moderate precipitation changes and a negative asymmetry (concave‐down) under extreme precipitation changes. In grasslands, the negative asymmetry in R_s response was attributed to the higher sensitivities of soil moisture, microbial and root activities to decreased precipitation (DPPT) than to increased precipitation (IPPT). In forests, the positive asymmetry was predominantly driven by the significant increase in microbial respiration under moderate IPPT, while the negative asymmetry was caused by the reductions in root biomass and respiration under extreme DPPT. The different asymmetric responses of R_s between grasslands and forests will greatly improve our ability to forecast the C cycle consequences of increased precipitation variability. Specifically, the negative asymmetry of R_s response under extreme precipitation change suggests that the soil C efflux will decrease across grasslands and forests under future precipitation regime with more wet and dry extremes.     

Carbon dynamics of mature and regrowth tropical forests derived from a pantropical database (TropForC‐db)

Tropical forests play a critical role in the global carbon (C) cycle, storing ~45% of terrestrial C and constituting the largest component of the terrestrial C sink. Despite their central importance to the global C cycle, their ecosystem‐level C cycles are not as well‐characterized as those of extra‐tropical forests, and knowledge gaps hamper efforts to quantify C budgets across the tropics and to model tropical forest‐climate interactions. To advance understanding of C dynamics of pantropical forests, we compiled a new database, the Tropical Forest C database (TropForC‐db), which contains data on ground‐based measurements of ecosystem‐level C stocks and annual fluxes along with disturbance history. This database currently contains 3568 records from 845 plots in 178 geographically distinct areas, making it the largest and most comprehensive database of its type. Using TropForC‐db, we characterized C stocks and fluxes for young, intermediate‐aged, and mature forests. Relative to existing C budgets of extra‐tropical forests, mature tropical broadleaf evergreen forests had substantially higher gross primary productivity (GPP) and ecosystem respiration (R_eco), their autotropic respiration (R_a) consumed a larger proportion (~67%) of GPP, and their woody stem growth (ANPP_stem) represented a smaller proportion of net primary productivity (NPP, ~32%) or GPP (~9%). In regrowth stands, aboveground biomass increased rapidly during the first 20 years following stand‐clearing disturbance, with slower accumulation following agriculture and in deciduous forests, and continued to accumulate at a slower pace in forests aged 20–100 years. Most other C stocks likewise increased with stand age, while potential to describe age trends in C fluxes was generally data‐limited. We expect that TropForC‐db will prove useful for model evaluation and for quantifying the contribution of forests to the global C cycle. The database version associated with this publication is archived in Dryad (DOI: 10.5061/dryad.t516f ) and a dynamic version is maintained at https://github.com/forc-db .     

全球陆地生态系统光合作用与呼吸作用的温度敏感性

基于全球647套通量数据,定量分析了全球尺度下生态系统光合作用和呼吸作用的温度敏感性(Q10)随纬度、气候和植被的分布规律。结果表明:在全球尺度下,光合作用和呼吸过程的温度敏感性(Q10,G和Q10,R)都随纬度的升高而增加,其中Q10,G和Q10,R的均值分别为3.99±0.21和2.28±0.074。除热带多树草原、常绿落叶林外,Q10,G均大于Q10,R值。不同植被类型的温度敏感性存在显著性差异,表现为:针叶林阔叶林;落叶林常绿林,其中生态系统的季节性变异是造成差异的主要原因。当植被类型和纬度区域共同影响Q10值时,植被类型对Q10值的总变异贡献更大。气候类型对Q10,G和Q10,R都有显著影响。在气候带上,干旱带的Q10,G最小,而冷温带的Q10,G最高。不同气候类型下(除温带草原气候外)的Q10,G都大于Q10,R。在极端条件下,温度可能不在是主导因素,而水分对温度敏感性的影响不可忽略,今后的研究需要更多的关注生态系统温度敏感性对水分变化的响应。     

Sensitivity of CO<Subscript>2</Subscript> Exchange of Fen Ecosystem Components to Water Level Variation

Abstract Climate change is predicted to bring about a water level (WL) draw-down in boreal peatlands. This study aimed to assess the effect of WL on the carbon dioxide (CO₂) dynamics of a boreal oligotrophic fen ecosystem and its components; Sphagnum mosses, sedges, dwarf shrubs and the underlying peat. We measured CO₂ exchange with closed chambers during four growing seasons in a study site that comprised different vegetation treatments. WL gradient in the site was broadened by surrounding half of the site with a shallow ditch that lowered the WL by 10–25 cm. We modeled gross photosynthesis (P _G) and ecosystem respiration (R _ECO) and simulated the CO₂ exchange in three WL conditions: prevailing and WL draw-down scenarios of 14 and 22 cm. WL draw-down both reduced the P _G and increased the R _ECO, thus leading to a smaller net CO₂ uptake in the ecosystem. Of the different components, Sphagnum mosses were most sensitive to WL draw-down and their physiological activities almost ceased. Vascular plant CO₂ exchange, en bloc, hardly changed but whereas sedges contributed twice as much to the CO₂ exchange as shrubs in the prevailing conditions, the situation was reversed in the WL draw-down scenarios. Peat respiration was the biggest component in R _ECO in all WL conditions and the increase in R _ECO following the WL draw-down was due to the increase in peat respiration. The results imply that functional diversity buffers the ecosystem against environmental variability and that in the long term, WL draw-down changes the vegetation composition of boreal fens.     

Respiration acclimation contributes to high carbon-use efficiency in a seasonally dry pine forest

Predictions of warming and drying in the Mediterranean and other regions require quantifying of such effects on ecosystem carbon dynamics and respiration. Long‐term effects can only be obtained from forests in which seasonal drought is a regular feature. We carried out measurements in a semiarid Pinus halepensis (Aleppo pine) forest of aboveground respiration rates of foliage, R_f, and stem, R_t over 3 years. Component respiration combined with ongoing biometric, net CO₂ flux [net ecosystem productivity (NEP)] and soil respiration measurements were scaled to the ecosystem level to estimate gross and net primary productivity (GPP, NPP) and carbon‐use efficiency (CUE=NPP/GPP) using 6 years data. GPP, NPP and NEP were, on average, 880, 350 and 211 g C m^?2 yr^?1, respectively. The above ground respiration made up half of total ecosystem respiration but CUE remained high at 0.4. Large seasonal variations in both R_f and R_t were not consistently correlated with seasonal temperature trends. Seasonal adjustments of respiration were observed in both the normalized rate (R₂₀) and short‐term temperature sensitivity (Q₁₀), resulting in low respiration rates during the hot, dry period. R_f in fully developed needles was highest over winter–spring, and foliage R₂₀ was correlated with photosynthesis over the year. Needle growth occurred over summer, with respiration rates in developing needles higher than the fully developed foliage at most times. R_t showed a distinct seasonal maximum in May irrespective of year, which was not correlated to the winter stem growth, but could be associated with phenological drivers such as carbohydrate re‐mobilization and cambial activity. We show that in a semiarid pine forest photosynthesis and stem growth peak in (wet) winter and leaf growth in (dry) summer, and associated adjustments of component respiration, dominated by those in R₂₀, minimize annual respiratory losses. This is likely a key for maintaining high CUE and ecosystem productivity similar to much wetter sites, and could lead to different predictions of the effect of warming and drying climate on productivity of pine forests than based on short‐term droughts.     

Climatic controls on the carbon and water balances of a boreal aspen forest, 1994–2003

The carbon and water budgets of boreal and temperate broadleaf forests are sensitive to interannual climatic variability and are likely to respond to climate change. This study analyses 9 years of eddy‐covariance data from the Boreal Ecosystem Research and Monitoring Sites (BERMS) Southern Old Aspen site in central Saskatchewan, Canada and characterizes the primary climatic controls on evapotranspiration, net ecosystem production (F_NEP), gross ecosystem photosynthesis (P) and ecosystem respiration (R). The study period was dominated by two climatic extremes: extreme warm and cool springs, which produced marked contrasts in the canopy duration, and a severe, 3‐year drought. Annual F_NEP varied among years from 55 to 367 g C m⁻² (mean 172, SD 94). Interannual variability in F_NEP was controlled primarily by factors that affected the R/P ratio, which varied between 0.74 and 0.96 (mean 0.87, SD 0.06). Canopy duration enhanced P and F_NEP with no apparent effect on R. The fraction of annual photosynthetically active radiation (PAR) that was absorbed by the canopy foliage varied from 38% in late leaf‐emergence years to 51% in early leaf‐emergence years. Photosynthetic light‐use efficiency (mean 0.0275, SD 0.026 mol C mol⁻¹ photons) was relatively constant during nondrought years but declined with drought intensity to a minimum of 0.0228 mol C mol⁻¹ photons during the most severe drought year. The impact of drought on F_NEP varied with drought intensity. Years of mild‐to‐moderate drought suppressed R while having little effect on P, so that F_NEP was enhanced. Years of severe drought suppressed both R and P, causing either little change or a subtle reduction in F_NEP. The analysis produced new insights into the dominance of canopy duration as the most important biophysical control on F_NEP. The results suggested a simple conceptual model for annual F_NEP in boreal deciduous forests. When water is not limiting, annual P is controlled by canopy duration via its influence on absorbed PAR at constant light‐use efficiency. Water stress suppresses P, by reducing light‐use efficiency, and R, by limiting growth and/or suppressing microbial respiration. The high photosynthetic light‐use efficiency showed this site to be a highly productive boreal deciduous forest, with properties similar to many temperate deciduous forests.     

Assessment of ecosystem CO2 efflux and its components in a Pinus canariensis forest at the treeline

We estimated respiratory fluxes in a treeline-associated Pinus canariensis forest in Tenerife, Canary Islands, an ecotone with strong seasonal changes in soil water availability. CO₂ efflux rates from the foliage, above ground woody tissue and the soil were measured by chamber techniques. Site-specific models obtained from these chamber measurements were then combined with half-hourly measurements of canopy, stem and soil temperature as well as with soil water potential, leaf area and stem sapwood volume data for scaling up component-specific CO₂ efflux to ecosystem respiration (R _ECO). Integrated over an entire year R _ECO was 550 g C m^?2 ground surface area (average of 2008 and 2009) and comprised the following component fluxes: 57 % from the soil, 10 % from above ground woody tissue and 33 % from the foliage. Between year differences in R _ECO and its components were <3 %. R _ECO varied markedly throughout an entire year generally following the seasonal trends in temperature during most of the year. During the dry summer, however, R _ECO was significantly reduced due to limited soil water availability in the rooting horizon. Thus, in treeline-associated forests under predicted scenarios of increasing aridity in Mediterranean regions, it is likely that there will be a shift in the contribution of CO₂ efflux from the soil (R _S), the foliage (R _F) and above ground woody tissue (R _W) to R _ECO from predominately below ground to increasingly above ground. Such changes should be taken into account for predicting the response of treeline forests to a changing climate at their upper distribution limit.     

Seasonal variation in leaf properties and ecosystem carbon budget in a cool-temperate deciduous broad-leaved forest: simulation analysis at Takayama site, Japan

Seasonal changes in gross primary production (GPP) and net ecosystem production (NEP) in temperate deciduous forests are mostly driven by environmental conditions and the phenology of leaf demography. This study addresses another factor, temporal changes in leaf properties, i.e., leaf aging from emergence to senescence. A process-based model was used to link the ecosystem-scale carbon budget with leaf-level properties on the basis of field observation and scaling procedures; temporal variations in leaf thickness (leaf mass per area, LMA), photosynthetic rubisco (V_cmax) and electron-transport (J_max) capacity, and dark respiration (R_d) were empirically parameterized. The model was applied to a cool-temperate deciduous broad-leaved forest at Takayama, in central Japan, and validated with data of net ecosystem CO₂ exchange (NEE=–NEP) measured using the eddy-covariance method. NEP of the Takayama site varied seasonally from 3 g C m^–2 day^–1 net source in late winter to 5 g C m^–2 day^–1 net sink in early to mid-summer. A sensitivity experiment showed that removing the leaf-aging effect changed the seasonal CO₂ exchange pattern, and led to overestimation of annual GPP by 6% and annual NEP by 38%. We found that seasonal variation in V_cmax affected the seasonal pattern and annual budget of CO₂ exchange most strongly; LMA and R_d had moderate influences. The rapid change in V_cmax and R_d during leaf emergence and senescence was important in evaluating GPP and NEP of the temperate deciduous forest.     

An improved approach for remotely sensing water stress impacts on forest C uptake

Given that forests represent the primary terrestrial sink for atmospheric CO₂, projections of future carbon (C) storage hinge on forest responses to climate variation. Models of gross primary production (GPP) responses to water stress are commonly based on remotely sensed changes in canopy ‘greenness’ (e.g., normalized difference vegetation index; NDVI). However, many forests have low spectral sensitivity to water stress (SSWS) – defined here as drought‐induced decline in GPP without a change in greenness. Current satellite‐derived estimates of GPP use a vapor pressure deficit (VPD) scalar to account for the low SWSS of forests, but fail to capture their responses to water stress. Our objectives were to characterize differences in SSWS among forested and nonforested ecosystems, and to develop an improved framework for predicting the impacts of water stress on GPP in forests with low SSWS. First, we paired two independent drought indices with NDVI data for the conterminous US from 2000 to 2011, and examined the relationship between water stress and NDVI. We found that forests had lower SSWS than nonforests regardless of drought index or duration. We then compared satellite‐derived estimates of GPP with eddy‐covariance observations of GPP in two deciduous broadleaf forests with low SSWS: the Missouri Ozark (MO) and Morgan Monroe State Forest (MMSF) AmeriFlux sites. Model estimates of GPP that used VPD scalars were poorly correlated with observations of GPP at MO (r² = 0.09) and MMSF (r² = 0.38). When we included the NDVI responses to water stress of adjacent ecosystems with high SSWS into a model based solely on temperature and greenness, we substantially improved predictions of GPP at MO (r² = 0.83) and for a severe drought year at the MMSF (r² = 0.82). Collectively, our results suggest that large‐scale estimates of GPP that capture variation in SSWS among ecosystems could improve predictions of C uptake by forests under drought.     

Effects of seasonal and interannual variations in leaf photosynthesis and canopy leaf area index on gross primary production of a cool-temperate deciduous broadleaf forest in Takayama, Japan

Revealing the seasonal and interannual variations in forest canopy photosynthesis is a critical issue in understanding the ecological mechanisms underlying the dynamics of carbon dioxide exchange between the atmosphere and deciduous forests. This study examined the effects of temporal variations of canopy leaf area index (LAI) and leaf photosynthetic capacity [the maximum velocity of carboxylation (V _cmax)] on gross primary production (GPP) of a cool-temperate deciduous broadleaf forest for 5 years in Takayama AsiaFlux site, central Japan. We made two estimations to examine the effects of canopy properties on GPP; one is to incorporate the in situ observation of V _cmax and LAI throughout the growing season, and another considers seasonality of LAI but constantly high V _cmax. The simulations indicated that variation in V _cmax and LAI, especially in the leaf expansion period, had remarkable effects on GPP, and if V _cmax was assumed constant GPP will be overestimated by 15%. Monthly examination of air temperature, radiation, LAI and GPP suggested that spring temperature could affect canopy phenology, and also that GPP in summer was determined mainly by incoming radiation. However, the consequences among these factors responsible for interannual changes of GPP are not straightforward since leaf expansion and senescence patterns and summer meteorological conditions influence GPP independently. This simulation based on in situ ecophysiological research suggests the importance of intensive consideration and understanding of the phenology of leaf photosynthetic capacity and LAI to analyze and predict carbon fixation in forest ecosystems.     

Estimates of CO2 uptake and release among European forests based on eddy covariance data

The net ecosystem exchange (NEE) of forests represents the balance of gross primary productivity (GPP) and respiration (R). Methods to estimate these two components from eddy covariance flux measurements are usually based on a functional relationship between respiration and temperature that is calibrated for night‐time (respiration) fluxes and subsequently extrapolated using daytime temperature measurements. However, respiration fluxes originate from different parts of the ecosystem, each of which experiences its own course of temperature. Moreover, if the temperature–respiration function is fitted to combined data from different stages of biological development or seasons, a spurious temperature effect may be included that will lead to overestimation of the direct effect of temperature and therefore to overestimates of daytime respiration. We used the EUROFLUX eddy covariance data set for 15 European forests and pooled data per site, month and for conditions of low and sufficient soil moisture, respectively. We found that using air temperature (measured above the canopy) rather than soil temperature (measured 5 cm below the surface) yielded the most reliable and consistent exponential (Q₁₀) temperature–respiration relationship. A fundamental difference in air temperature‐based Q₁₀ values for different sites, times of year or soil moisture conditions could not be established; all were in the range 1.6–2.5. However, base respiration (R₀, i.e. respiration rate scaled to 0°C) did vary significantly among sites and over the course of the year, with increased base respiration rates during the growing season. We used the overall mean Q₁₀ of 2.0 to estimate annual GPP and R. Testing suggested that the uncertainty in total GPP and R associated with the method of separation was generally well within 15%. For the sites investigated, we found a positive relationship between GPP and R, indicating that there is a latitudinal trend in NEE because the absolute decrease in GPP towards the pole is greater than in R.     

Enhanced CO2 uptake is marginally offset by altered fluxes of non-CO2 greenhouse gases in global forests and grasslands under N deposition

Despite the increasing impact of atmospheric nitrogen (N) deposition on terrestrial greenhouse gas (GHG) budget, through driving both the net atmospheric CO₂ exchange and the emission or uptake of non-CO₂ GHGs (CH₄ and N₂O), few studies have assessed the climatic impact of forests and grasslands under N deposition globally based on different bottom-up approaches. Here, we quantify the effects of N deposition on biomass C increment, soil organic C (SOC), CH₄ and N₂O fluxes and, ultimately, the net ecosystem GHG balance of forests and grasslands using a global comprehensive dataset. We showed that N addition significantly increased plant C uptake (net primary production) in forests and grasslands, to a larger extent for the aboveground C (aboveground net primary production), whereas it only caused a small or insignificant enhancement of SOC pool in both upland systems. Nitrogen addition had no significant effect on soil heterotrophic respiration (R_H) in both forests and grasslands, while a significant N-induced increase in soil CO₂ fluxes (R_S, soil respiration) was observed in grasslands. Nitrogen addition significantly stimulated soil N₂O fluxes in forests (76%), to a larger extent in grasslands (87%), but showed a consistent trend to decrease soil uptake of CH₄, suggesting a declined sink capacity of forests and grasslands for atmospheric CH₄ under N enrichment. Overall, the net GHG balance estimated by the net ecosystem production-based method (forest, 1.28 Pg CO₂-eq year⁻¹ vs. grassland, 0.58 Pg CO₂-eq year⁻¹) was greater than those estimated using the SOC-based method (forest, 0.32 Pg CO₂-eq year⁻¹ vs. grassland, 0.18 Pg CO₂-eq year⁻¹) caused by N addition. Our findings revealed that the enhanced soil C sequestration by N addition in global forests and grasslands could be only marginally offset (1.5%–4.8%) by the combined effects of its stimulation of N₂O emissions together with the reduced soil uptake of CH₄.     

Seasonal controls on interannual variability in carbon dioxide exchange of a near-end-of rotation Douglas-fir stand in the Pacific Northwest, 1997–2006

This study analyzes 9 years of eddy‐covariance (EC) data carried out in a Pacific Northwest Douglas‐fir (Pseudotsuga menzesii) forest (58‐year old in 2007) on the east coast of Vancouver Island, Canada, and characterizes the seasonal and interannual variability in net ecosystem productivity (NEP), gross primary productivity (GPP), and ecosystem respiration (R_e) and primary climatic controls on these fluxes. The annual values (± SD) of NEP, GPP and R_e were 357 ± 51, 2124 ± 125, and 1767 ± 146 g C m^?2 yr^?1, respectively, with ranges of 267–410, 1592–2338, and 1642–2071 g C m^?2 yr^?1, respectively. Spring to early summer (March–June) accounted for more than 80% of annual NEP while late spring to early autumn (May–August) was mainly responsible for its interannual variability (～80%). The major drivers of interannual variability in annual carbon (C) fluxes were annual and spring mean air temperatures (T_a) and water deficiency during late summer and autumn (July–October) when this Douglas‐fir forest growth was often water‐limited. Photosynthetically active radiation (Q), and the combination of Q and soil water content (θ) explained 85% and 91% of the variance of monthly GPP, respectively; and 91% and 96% of the variance of monthly R_e was explained by T_a and the combination of T_a and θ, respectively. Annual net C sequestration was high during optimally warm and normal precipitation years, but low in unusually warm or severely dry years. Excluding 1998 and 1999, the 2 years strongly affected by an El Niño/La Niña cycle, annual NEP significantly decreased with increasing annual mean T_a. Annual NEP will likely decrease whereas both annual GPP and R_e will likely increase if the future climate at the site follows a trend similar to that of the past 40 years.     

Scaling relationships for woody tissue respiration in two tropical rain forests

The relationship between gross primary productivity (GPP) and net primary productivity (NPP) is not fully understood. One of the uncertainties relevant to this issue is the magnitude of woody tissue respiration. Although some data exist for temperate and boreal zones, measurements of woody tissue respiration in tropical forests are sparse. We made in situ chamber measurements of woody tissue respiration in two tropical rain forests, one in the Brazilian Amazon (Reserva Jarú) and one in Central Cameroon (Mbalmayo Reserve). We made measurements on a wide range of species at each site and over a range of stem diameters from 0·02 to 1·4 m. The rate of efflux of carbon dioxide (CO₂) from bark at 25 °C, R_t, varied from 0·1 to 5·2 µmol m^?2 s^?1 across the two sites, and the efflux was related to both volume and surface area components of the measured stem sections. The temperature response in R_t was slightly higher at Jarú than at Mbalmayo, with Q₁₀ values of 1·8 (± 0·1 SE) and 1·6 (± 0·1 SE), respectively. A log–log regression showed that R_t was significantly related to stem diameter, D (P < 0·001; r² = 0·58–0·62) and was significantly higher at Mbalmayo than at Jarú (P < 0·001), but that the rate of increase in R_t with stem diameter, D, was similar between sites. At the Mbalmayo site, tree growth measurements made over a 4 month period were used to make two estimates of the maintenance (R_m) and construction (R_c) components of respiration embedded in R_t. The two methods agreed closely, suggesting that R_m was approximately 80% of R_c at this site. R_m could be strongly related to D using a sigmoidal relationship that described both surface area and volume components as sources of respiratory CO₂ (r² = 0·71). This functional model was combined with inventory, growth and climate data for the Mbalmayo site to make a first estimate of annual above‐ground woody tissue respiration, R_A, which was 257 (± 18 SE) g C m^?2 year^?1. This value corresponds to approximately 10% of GPP, slightly lower than that found for another tropical rain forest, but higher than for temperate forests. When combined with data from six other sites in tropical, temperate and boreal settings, a very strong relationship was found between R_A and leaf area index (LAI), and between R_A/GPP and LAI (P < 0·001, r² = 0·98). This indicates that R_A exerts an appreciable constraint on NPP and that this constraint varies closely with LAI across widely differing types of woody vegetation.     

Composition,dynamics and disturbance regime of temperate deciduous forests in Monsoon Asia

Temperate deciduous forests in Monsoon Asia are classified into three forest types which differ in floristic composition, dynamics and disturbance regime. The cool temperate mixed deciduous broadleaf/conifer forest, dominated by Quercus spp. (mainly Q. mongolica or Q. crispula) and conifers, is distributed in northern parts of the temperate zone. The cool temperate deciduous forest, dominated by Fagus crenata, is distributed in Honshu, Japan under a humid climate through the year. The warm temperate deciduous forest dominated by Quercus spp. (mainly Q. acutissima or Q. serrata) occurs in the continental areas, the Korean Peninsula, and the Pacific Ocean side of Japan. The species diversity of cool temperate deciduous forest was lower than the other two types because of the intensive dominance of Fagus crenata. The disturbance regimes also varies among the three types; small scale treefall gaps are prevailing in the cool temperate deciduous forest, while larger scaled disturbances are important in the other two forest types. Fire seems to be important in the warm temperate deciduous forest, and both fire and large scale blowdowm seems important in the cool temperate mixed broadleaf/conifer forest. These differences in forest composition and disturbance regimes associated with climatic conditions and ancient human impacts have a close analogy with the Northeastern Hardwood forests in North America.