Carbon dioxide exchange above a Mediterranean C3/C4 grassland during two climatologically contrasting years

Eddy‐covariance measurements of net ecosystem carbon exchange (NEE) were carried out above a grazed Mediterranean C3/C4 grassland in southern Portugal, during two hydrological years, 2004–2005 and 2005–2006, of contrasting rainfall. Here, we examine the seasonal and interannual variation in NEE and its major components, gross primary production (GPP) and ecosystem respiration (R_eco), in terms of the relevant biophysical controls. The first hydrological year was dry, with total precipitation 45% below the long‐term mean (669 mm) and the second was normal, with total precipitation only 12% above the long‐term mean. The drought conditions during the winter and early spring of the dry year limited grass production and the leaf area index (LAI) was very low. Hence, during the peak of the growth period, the maximum daily rate of NEE and the light‐use and water‐use efficiencies were approximately half of those observed in the normal year. In the summer of 2006, the warm‐season C4 grass, Cynodon dactylon L., exerted an evident positive effect on NEE by converting the ecosystem into a carbon sink after strong rain events and extending the carbon sequestration for several days, after the end of senescence of the C3 grasses. On an annual basis, the GPP and NEE were 524 and 49 g C m^?2, respectively, for the dry year, and 1261 and ?190 g C m^?2 for the normal year. Therefore, the grassland was a moderate net source of carbon to the atmosphere, in the dry year, and a considerable net carbon sink, in the normal year. In these 2 years of experiment the total amount of precipitation was the main factor determining the interannual variation in NEE. In terms of relevant controls, GPP and NEE were strongly related to incident photosynthetic photon flux density on short‐term time scales. Changes in LAI explained 84% and 77% of the variation found in GPP and NEE, respectively. Variations in R_eco were mainly controlled by canopy photosynthesis. After each grazing event, the reduction in LAI affected negatively the NEE.     

Is Net Ecosystem Production Equal to Ecosystem Carbon Accumulation?

Net ecosystem production (NEP), defined as the difference between gross primary production and total ecosystem respiration, represents the total amount of organic carbon in an ecosystem available for storage, export as organic carbon, or nonbiological oxidation to carbon dioxide through fire or ultraviolet oxidation. In some of the recent literature, especially that on terrestrial ecosystems, NEP has been redefined as the rate of organic carbon accumulation in the system. Here we argue that retaining the original definition maintains the conceptual coherence between NEP and net primary production and that it is congruous with the widely accepted definitions of ecosystem autotrophy and heterotrophy. Careful evaluation of NEP highlights the various potential fates of nonrespired carbon in an ecosystem.     

森林生态系统碳循环的基本概念和野外测定方法评述

全球气候变化与森林生态系统碳循环息息相关,定量评估森林碳收支是生态系统与全球变化研究的重要任务。30年来森林生态系统碳循环研究已经取得了长足的进展,但全球和区域森林碳收支仍然存在很大的不确定性。这一方面与森林生态系统本身的复杂性有关,另一方面也与具体研究方法有关。评述了森林生态系统碳循环的基本概念和主要野外测定方法,为我国森林生态系统碳循环研究提供可参考的方法论。从生态系统碳浓度、密度、通量、分配和周转5个方面回顾了碳循环相关概念,指出碳浓度和碳储量是对碳库的静态描述,而碳通量和碳周转是对碳库的动态描述。净初级生产力是测量最普遍的碳通量组分,但大多数情况下因忽略了一些细节而被系统低估。普遍使用的净生态系统生产力,由于没有包含非CO2形式的水文、气象和干扰过程产生的碳通量,通常情况下高于生态系统净碳累积速率。在详细介绍碳通量组分的基础上,改进了森林生态系统碳循环的概念模型。重点讨论了碳通量的3种地面实测方法:测树学方法、箱法和涡度协方差法,并指出了其注意事项和不确定性来源。针对当前碳循环研究的突出问题,建议从4个方面减小碳循环测定的不确定性:(1)恰当运用生物量方程估算乔木生物量;(2)尽可能全面测定生态系统碳组分;(3)给出碳通量估算值的不确定性;(4)多种途径交互验证。     

Reconciling Carbon-cycle Concepts, Terminology, and Methods

Recent projections of climatic change have focused a great deal of scientific and public attention on patterns of carbon (C) cycling as well as its controls, particularly the factors that determine whether an ecosystem is a net source or sink of atmospheric carbon dioxide (CO₂). Net ecosystem production (NEP), a central concept in C-cycling research, has been used by scientists to represent two different concepts. We propose that NEP be restricted to just one of its two original definitions—the imbalance between gross primary production (GPP) and ecosystem respiration (ER). We further propose that a new term—net ecosystem carbon balance (NECB)—be applied to the net rate of C accumulation in (or loss from [negative sign]) ecosystems. Net ecosystem carbon balance differs from NEP when C fluxes other than C fixation and respiration occur, or when inorganic C enters or leaves in dissolved form. These fluxes include the leaching loss or lateral transfer of C from the ecosystem; the emission of volatile organic C, methane, and carbon monoxide; and the release of soot and CO₂ from fire. Carbon fluxes in addition to NEP are particularly important determinants of NECB over long time scales. However, even over short time scales, they are important in ecosystems such as streams, estuaries, wetlands, and cities. Recent technological advances have led to a diversity of approaches to the measurement of C fluxes at different temporal and spatial scales. These approaches frequently capture different components of NEP or NECB and can therefore be compared across scales only by carefully specifying the fluxes included in the measurements. By explicitly identifying the fluxes that comprise NECB and other components of the C cycle, such as net ecosystem exchange (NEE) and net biome production (NBP), we can provide a less ambiguous framework for understanding and communicating recent changes in the global C cycle.     

四川西北部亚高山云杉天然林生态系统碳密度、净生产量和碳贮量的初步研究

该文利用野外实际调查数据对四川西北部亚高山云杉(Picea asperata)天然林碳密度、净生产量、碳贮量及其分布进行了分析,结果表明,在调查区域,云杉天然林分平均生物量为230.37×10³ kg·hm^-2,其中乔木层为212.77×10³ kg·hm^-2,占林分生物量的92.30%。云杉天然林生态系统各组分的平均碳密度为树干57.85%,树皮47.12%,树枝51.22%,树叶48.27%和树根52.39%,灌木层平均碳密度49.91%,草本层平均碳密度46.34%,地被层平均碳密度43.21%,枯落物层平均碳密度39.44%,土壤碳密度平均值为1.41%,随土层深度增加各层次土壤碳密度逐渐减少。云杉林平均生态系统总碳贮量为273.79×10³ kg·hm^-2,其中乔木层109.30×10³ kg·hm^-2,占云杉林生态系统总碳贮量的39.92%,灌木层5.69×10³ kg·hm^-2,占2.08%,草本层1.26×10³ kg·hm^-2,占0.46%,地被物层0.60×10³ kg·hm^-2,占0.22%,枯落物层0.83×10³ kg·hm^-2,占0.30%,林内土壤(0~100 cm)碳贮量为156.11×10³ kg·hm^-2,占57.01%。云杉林的碳库分布序列为土壤(0~100 cm)>乔木层>灌木层>草本层>枯落物层>地被物层。云杉天然林分平均净生产总量为6 838.5 kg·hm^-2·a^-1,碳素年总净固量平均为3 584.98 kg·hm^-2·a^-1,其中乔木层净生产量为4 676 kg·hm^-2·a^-1,占林分总量的68.38%,碳素年平均固定量2 552.99 kg·hm^-2·a^-1,占林分总量的71.21%。     

小兴安岭谷地云冷杉林的碳密度与生产力

采用样地清查和异速生长方程法,量化了处于衰退状态的小兴安岭谷地云冷杉林的森林碳密度和生产力.结果表明： 2011年森林碳密度总量为268.14 t C·hm^-2,其中植被碳密度、碎屑碳密度和土壤碳密度分别为74.25、16.86和177.03 t C·hm^-2.2006—2011年,乔木层碳密度从80.86 t C·hm^-2减少到71.73 t C·hm^-2,主要树种冷杉、白桦、云杉和兴安落叶松的碳密度年均减少比例分别为0.5%、1.2%、2.7%和3.7%,毛赤杨、红松和花楷槭的碳密度年均增加比例分别为2.9%、3.9%和7.2%.森林净初级生产力(NPP)为4.69 t C·hm^-2·a^-1,地下部和地上部NPP比值为0.56,凋落物损失部分是总NPP的最大组分,所占比例为34.5%.森林生态系统中2个主要碳输出途径异养呼吸和粗木质残体分解的年通量分别为293.67和119.29 g C·m^-2·a^-1.森林净生态系统生产力(NEP)为55.90 g C·m^-2·a^-1.研究结果表明,处于衰退状态的谷地云冷杉林仍具有一定的碳汇功能.     

Patterns of Production and Precipitation-Use Efficiency of Winter Wheat and Native Grasslands in the Central Great Plains of the United States

The Great Plains of the United States is characterized by a large west–east gradient in annual precipitation and a similar large north–south gradient in annual temperature. Native grasslands and winter wheat are found over a large portion of the precipitation and temperature gradients. In this article, we use long-term data to analyze the differences in the patterns in aboveground net primary production and precipitation-use efficiency between wheat and native grassland ecosystems in the central portion of Great Plains, and their relationships to potential water availability (precipitation). Aboveground net primary production of native grasslands shows a large response to precipitation. Aboveground net primary production of winter wheat has a smaller response to changing precipitation. Annual precipitation-use efficiency of native grasslands is unaffected by increases in average annual precipitation, but precipitation-use efficiency of summer-fallow wheat ecosystems decreases substantially with increased average precipitation. Our results suggest that in the wetter portion of the central Great Plains, summer-fallow wheat management is relatively inefficient, because increased water availability results in diminishing returns. Comparisons with data from continuously cropped wheat confirmed this result. Shifts across the region to continuous cropping of wheat potentially could have significant impacts on regional wheat yield, carbon balance, and economic status. Received 15 October 1999; accepted 10 March 2000.     

Net primary production in three bioenergy crop systems following land conversion

Aims Identifying the amount of production and the partitioning to above- and belowground biomass is generally the first step toward selecting bioenergy systems. There are very few existing studies on the dynamics of production following land conversion. The objectives of this study were to (i) determine the differences in aboveground net primary production (ANPP), belowground net primary production (BNPP), shoot-to-root ratio (S:R) and leaf area index in three bioenergy crop systems and (ii) evaluate the production of these three systems in two different land use conversions.Methods This investigation included biometric analysis of NPP on three agricultural sites converted from conservation reserve program (CRP) management to bioenergy crop production (corn, switchgrass and prairie mix) and three sites converted from traditional agriculture production to bioenergy crop production.Important findings The site converted from conventional agriculture produced smaller ANPP in corn (19.03±1.90 standard error [SE] Mg ha^-1 year^-1) than the site converted from CRP to corn (24.54±1.43 SE Mg ha^-1 year^-1). The two land conversions were similar in terms of ANPP for switchgrass (4.88±0.43 SE for CRP and 2.04±0.23 SE Mg ha^-1 year^-1 for agriculture) and ANPP for prairie mix (4.70±0.50 SE for CRP and 3.38±0.33 SE Mg ha^-1 year^-1 for agriculture). The BNPP at the end of the growing season in all the bioenergy crop systems was not significantly different (P = 0.75, N = 8).     

Plant respiration: Controlled by photosynthesis or biomass?

Two simplifying hypotheses have been proposed for whole‐plant respiration. One links respiration to photosynthesis; the other to biomass. Using a first‐principles carbon balance model with a prescribed live woody biomass turnover, applied at a forest research site where multidecadal measurements are available for comparison, we show that if turnover is fast the accumulation of respiring biomass is low and respiration depends primarily on photosynthesis; while if turnover is slow the accumulation of respiring biomass is high and respiration depends primarily on biomass. But the first scenario is inconsistent with evidence for substantial carry‐over of fixed carbon between years, while the second implies far too great an increase in respiration during stand development—leading to depleted carbohydrate reserves and an unrealistically high mortality risk. These two mutually incompatible hypotheses are thus both incorrect. Respiration is not linearly related either to photosynthesis or to biomass, but it is more strongly controlled by recent photosynthates (and reserve availability) than by total biomass.     

Site-level evaluation of satellite-based global terrestrial gross primary production and net primary production monitoring

Operational monitoring of global terrestrial gross primary production (GPP) and net primary production (NPP) is now underway using imagery from the satellite‐borne Moderate Resolution Imaging Spectroradiometer (MODIS) sensor. Evaluation of MODIS GPP and NPP products will require site‐level studies across a range of biomes, with close attention to numerous scaling issues that must be addressed to link ground measurements to the satellite‐based carbon flux estimates. Here, we report results of a study aimed at evaluating MODIS NPP/GPP products at six sites varying widely in climate, land use, and vegetation physiognomy. Comparisons were made for twenty‐five 1 km² cells at each site, with 8‐day averages for GPP and an annual value for NPP. The validation data layers were made with a combination of ground measurements, relatively high resolution satellite data (Landsat Enhanced Thematic Mapper Plus at ～30 m resolution), and process‐based modeling. There was strong seasonality in the MODIS GPP at all sites, and mean NPP ranged from 80 g C m^?2 yr^?1 at an arctic tundra site to 550 g C m^?2 yr^?1 at a temperate deciduous forest site. There was not a consistent over‐ or underprediction of NPP across sites relative to the validation estimates. The closest agreements in NPP and GPP were at the temperate deciduous forest, arctic tundra, and boreal forest sites. There was moderate underestimation in the MODIS products at the agricultural field site, and strong overestimation at the desert grassland and at the dry coniferous forest sites. Analyses of specific inputs to the MODIS NPP/GPP algorithm – notably the fraction of photosynthetically active radiation absorbed by the vegetation canopy, the maximum light use efficiency (LUE), and the climate data – revealed the causes of the over‐ and underestimates. Suggestions for algorithm improvement include selectively altering values for maximum LUE (based on observations at eddy covariance flux towers) and parameters regulating autotrophic respiration.     

Response of understory vegetation to variable tree mortality following a mountain pine beetle epidemic in lodgepole pine stands in northern Utah

We analyzed the long-term dynamics of aboveground biomass ofLeymus chinense steppe in relation to interannual variation of precipitation and temperature during 1980–1989 at levels of community, growth form and species in the Xilin river basin, Inner Mongolia Autonomous Region, China. Annual aboveground net primary production (ANPP) varied from 154.00 g m^-2 yr^-1 in 1980 to 318.59 g m^-2 yr^-1 in 1988, with a mean of 248.63 g m^-2 yr^-1 and the coefficient of variation of 25%. ANPP was not significantly correlated to annual precipitation and total precipitation during April–September atp0.05 level, but precipitation in May and August accounted for 69% of interannual variation of ANPP. The means of rain use efficiency and water use efficiency ofL. chinense steppe were 8.1 kg DM ha^-1 mm^-1 yr^-1 and 0.89 mg DM g^-1 H₂O respectively. Aboveground biomass of various growth forms and species had different response patterns to interannual variation of precipitation and temperature. Monthly and seasonal distribution of precipitation and temperature were the key controls of aboveground biomass of species.     

Does climate directly influence NPP globally?

The need for rigorous analyses of climate impacts has never been more crucial. Current textbooks state that climate directly influences ecosystem annual net primary productivity (NPP), emphasizing the urgent need to monitor the impacts of climate change. A recent paper challenged this consensus, arguing, based on an analysis of NPP for 1247 woody plant communities across global climate gradients, that temperature and precipitation have negligible direct effects on NPP and only perhaps have indirect effects by constraining total stand biomass (M_tot) and stand age (a). The authors of that study concluded that the length of the growing season (l_gs) might have a minor influence on NPP, an effect they considered not to be directly related to climate. In this article, we describe flaws that affected that study's conclusions and present novel analyses to disentangle the effects of stand variables and climate in determining NPP. We re‐analyzed the same database to partition the direct and indirect effects of climate on NPP, using three approaches: maximum‐likelihood model selection, independent‐effects analysis, and structural equation modeling. These new analyses showed that about half of the global variation in NPP could be explained by M_tot combined with climate variables and supported strong and direct influences of climate independently of M_tot, both for NPP and for net biomass change averaged across the known lifetime of the stands (ABC = average biomass change). We show that l_gs is an important climate variable, intrinsically correlated with, and contributing to mean annual temperature and precipitation (T_ann and P_ann), all important climatic drivers of NPP. Our analyses provide guidance for statistical and mechanistic analyses of climate drivers of ecosystem processes for predictive modeling and provide novel evidence supporting the strong, direct role of climate in determining vegetation productivity at the global scale.     

The Carbon Balance of an Old-growth Forest: Building Across Approaches

The carbon budget of the Wind River old-growth forest is being addressed from a variety of perspectives and with a range of approaches. The goal of this comprehensive analysis is developing a thorough, general, and validated understanding of the carbon balance, as well as the processes controlling it. The initial results from studies addressing annual carbon (C) balance with ground-based methods, eddy flux, leaf-based models, and ecosystem models are consistent in some, but not all, respects. Net primary production is 500–600 g C m^–2 y^–1 (5–6 Mg C ha^–1 y^–1), consistent with estimates based on climate alone. The site appears to be close to carbon equilibrium, as a multiyear average, using ground-based methods but a sink of approximately 150–190 g C m^–2 y^–1 from eddy flux for a single year. An overview of the mechanisms that can drive forest carbon sinks illustrates why methods emphasizing different temporal and spatial scales, as well as different processes, can come to different conclusions, and it highlights opportunities in moving toward a truly integrated approach.     

Production,Respiration, and Overall Carbon Balance in an Old-growth <Emphasis Type="Italic">Pseudotsuga</Emphasis>-<Emphasis Type="Italic">Tsuga</Emphasis> Forest Ecosystem

Ground-based measurements of stores, growth, mortality, litterfall, respiration, and decomposition were conducted in an old-growth forest at Wind River Experimental Forest, Washington, USA. These measurements were used to estimate gross primary production (GPP) and net primary production (NPP); autotrophic respiration (R_a) and heterotrophic (R_h) respiration; and net ecosystem production (NEP). Monte Carlo methods were used to calculate uncertainty (expressed as ± 2 standard deviations of 200–400 calculations). Live carbon (C) stores were 39,800 g C m^–2 (34,800–44,800 g C m^–2). The store of C in detritus and mineral soil was 22,092 g C m^–2 (20,600–23,600 g C m^–2), and the total C stores were 61,899 g C m^–2 (56,600–67,700 g C m^–2). Total NPP was 597 g C m^–2 y^–1 (453 to 741 g C m^–2 y^–1). R_a was 1309 g C m^–2 y^–1 (845–1773 g C m^–2 y^–1), indicating a GPP of 1906 g C m^–2 y^–1 (1444–2368 g C m^–2 y^–1). R_h, including the respiration of heart rots in tree boles, was 577 g C m^–2 y^–1 (479–675 g C m^–2 y^–1). Long-term NEP was estimated to be +20 g C m^–2 y^–1 (–116 to +156 g C m^–2 y^–1), indicating this stand might be a small sink. These estimates contrast with the larger sink estimated at the same site using eddy-flux methods. Several hypotheses to explain this discrepancy were explored, including (a) undetected biomass increases, (b) underestimates of NPP, (c) unmeasured losses, and (d) a temporal mismatch between the two sets of measurements. The last hypothesis appears the most likely.