Stable Isotopes and Carbon Cycle Processes in Forests and Grasslands

Abstract: Scaling and partitioning are frequently two difficult challenges facing ecology today. With regard to ecosystem carbon balance studies, ecologists and atmospheric scientists are often interested in asking how fluxes of carbon dioxide scale across the landscape, region and continent. Yet at the same time, physiological ecologists and ecosystem ecologists are interested in dissecting the net ecosystem CO₂ exchange between the biosphere and the atmosphere to achieve a better understanding of the balance between photosynthesis and respiration within a forest. In both of these multiple-scale ecological questions, stable isotope analyses of carbon dioxide can play a central role in influencing our understanding of the extent to which terrestrial ecosystems are carbon sinks. In this synthesis, we review the theory and present field evidence to address isotopic scaling of CO₂ fluxes. We first show that the ¹³C isotopic signal which ecosystems impart to the atmosphere does not remain constant over time at either temporal or spatial scales. The relative balances of different biological activities and plant responses to stress result in dynamic changes in the ¹³C isotopic exchange between the biosphere and atmosphere, with both seasonal and stand-age factors playing major roles influencing the ¹³C biosphere-atmosphere exchange. We then examine how stable isotopes are used to partition net ecosystem exchange fluxes in order to calculate shifts in the balance of photosynthesis and respiration. Lastly, we explore how fundamental differences in the ¹⁸O isotopic gas exchange of forest and grassland ecosystems can be used to further partition terrestrial fluxes.     

稳定性同位素技术与Keeling曲线法在陆地生态系统碳/水交换研究中的应用

稳定性同位素技术和Keeling曲线法是现代生态学研究的重要手段和方法之一。稳定性同位素能够整合生态系统复杂的生物学、生态学和生物地球化学过程在时间和空间尺度上对环境变化的响应。Keeling曲线法是以生物过程前后物质平衡理论为基础,将CO₂或H₂O的同位素组成(δD、δ¹³C或δ¹⁸O)与其对应浓度测量结合起来,将生态系统净碳通量区分为光合固定和呼吸释放通量,或将整个生态系统水分蒸散区分为植物蒸腾和土壤蒸发。在全球尺度上,稳定性同位素技术、Keeling曲线法与全球尺度陆地生态系统模型相结合,还可区分陆地生态系统和海洋生态系统对全球碳通量的贡献以及不同植被类型(C₃或C₄)在全球CO₂同化量中所占的比例。然而,生态系统的异质性使得稳定性同位素技术和Keeling曲线法从冠层尺度外推到生态系统、区域或全球尺度时存在有一定程度的不确定性。此外,取样时间、地点的选取也会影响最终的研究结果。尽管如此,随着分析手段的不断精确和研究方法的日趋完善,稳定性同位素技术和Keeling曲线法与其它测量方法(如微气象法)的有机结合将成为未来陆地生态系统碳/水交换研究的重要手段和方法之一。     

生态系统光合与呼吸拆分的同位素理论、方法和应用进展

生态系统光合和呼吸是构成净生态系统CO₂交换量(NEE)的重要组分。涡度相关技术可直接观测生态系统NEE,并通过建立温度回归或光响应曲线等函数将NEE统计拆分为生态系统光合和呼吸,但是存在自相关和高估白天呼吸等问题。稳定同位素红外光谱技术的进步使高时间分辨率大气CO₂及其稳定碳同位素组成(δ¹³C)的连续观测成为可能,与涡度相关技术观测的NEE数据相结合,可实现昼夜和季节尺度生态系统光合和呼吸拆分。本文系统阐述了生态系统光合与呼吸的同位素通量拆分方法的基本理论与假设,阐述了同位素通量观测技术的发展及其应用进展,综述了同位素通量拆分理论解析生态系统光合与呼吸过程的新机制认识,最后总结并展望了同位素通量拆分理论的不确定性以及开展多种拆分方法综合比较的必要性。     

Using continuous stable isotope measurements to partition net ecosystem CO2 exchange

Ecosystem-scale estimation of photosynthesis and respiration using micrometeorological techniques remains an important, yet difficult, challenge. In this study, we combined micrometeorological and stable isotope methods to partition net ecosystem CO2 exchange (FN) into photosynthesis (F(A)) and respiration (F(R)) in a corn-soybean rotation ecosystem during the summer 2003 corn phase. Mixing ratios of (12)CO2 and (13)CO2 were measured continuously using tunable diode laser (TDL) absorption spectroscopy. The dynamics of the isotope ratio of ecosystem respiration (R), net ecosystem CO2 exchange (deltaN) and photosynthetic discrimination at the canopy scale (delta canopy) were examined. During the period of full canopy closure, F(N) was partitioned into photosynthesis and respiration using both the isotopic approach and the conventional night-time-derived regression methodology. Results showed that deltaR had significant seasonal variation (-32 to -11% per hundred) corresponding closely with canopy phenology. Daytime deltaN typically varied from -12 to -4% per hundred, while delta canopy remained relatively constant in the vicinity of 3% per hundred. Compared with the regression approach, the isotopic flux partitioning showed more short-term variations and was considerably more symmetric about F(N). In this experiment, the isotopic partitioning resulted in larger uncertainties, most of which were caused by the uncertainties in deltaN. and the daytime estimate of deltaR. By sufficiently reducing these uncertainties, the tunable diode laser (TDL)-micrometeorological technique should yield a better understanding of the processes controlling photosynthesis, respiration and ecosystem-scale discrimination.     

基于稳定同位素的SPAC水碳拆分及耦合研究进展

土壤-植被-大气连续体(SPAC)是陆地水文学、生态学和全球变化领域的重要研究对象,其水碳循环过程及耦合机制是前沿性问题.稳定同位素技术示踪、整合和指示的特征有助于评估分析生态系统固碳和耗水情况.本文在简述稳定同位素应用原理和技术的基础上,重点阐释了基于稳定同位素光学技术的SPAC系统水碳交换研究进展,包括:在净碳通量中拆分光合与呼吸量,在蒸散通量中拆分蒸腾与蒸发量,以及在系统尺度上的水碳耦合研究.新兴的技术和方法实现了生态系统尺度上长期高频的同位素观测,但在测量精准度、生态系统呼吸拆分、非稳态模型适应性、尺度转换和水碳耦合机制等方面存在挑战.本文探讨了现有主要研究成果、局限性以及未来研究展望,以期对稳定同位素生态学领域的新研究和技术发展有所帮助.     

The dependence of respiration on photosynthetic substrate supply and temperature: integrating leaf, soil and ecosystem measurements

Interactions between photosynthetic substrate supply and temperature in determining the rate of three respiration components (leaf, belowground and ecosystem respiration) were investigated within three environmentally controlled, Populus deltoides forest bays at Biosphere 2, Arizona. Over 2 months, the atmospheric CO₂ concentration and air temperature were manipulated to test the following hypotheses: (1) the responses of the three respiration components to changes in the rate of photosynthesis would differ both in speed and magnitude; (2) the temperature sensitivity of leaf and belowground respiration would increase in response to a rise in substrate availability; and, (3) at the ecosystem level, the ratio of respiration to photosynthesis would be conserved despite week‐to‐week changes in temperature. All three respiration rates responded to the CO₂ concentration‐induced changes in photosynthesis. However, the proportional change in the rate of leaf respiration was more than twice that of belowground respiration and, when photosynthesis was reduced, was also more rapid. The results suggest that aboveground respiration plays a key role in the overall response of ecosystem respiration to short‐term changes in canopy photosynthesis. The short‐term temperature sensitivity of leaf respiration, measured within a single night, was found to be affected more by developmental conditions than photosynthetic substrate availability, as the Q₁₀ was lower in leaves that developed at high CO₂, irrespective of substrate availability. However, the temperature sensitivity of belowground respiration, calculated between periods of differing air temperature, appeared to be positively correlated with photosynthetic substrate availability. At the ecosystem level, respiration and photosynthesis were positively correlated but the relationship was affected by temperature; for a given rate of daytime photosynthesis, the rate of respiration the following night was greater at 25 than 20°C. This result suggests that net ecosystem exchange did not acclimate to temperature changes lasting up to 3 weeks. Overall, the results of this study demonstrate that the three respiration terms differ in their dependence on photosynthesis and that, short‐ and medium‐term changes in temperature may affect net carbon storage in terrestrial ecosystems.     

Effects of Experimental Water Table and Temperature Manipulations on Ecosystem CO2 Fluxes in an Alaskan Rich Fen

Peatlands store 30% of the world’s terrestrial soil carbon (C) and those located at northern latitudes are expected to experience rapid climate warming. We monitored growing season carbon dioxide (CO₂) fluxes across a factorial design of in situ water table (control, drought, and flooded plots) and soil warming (control vs. warming via open top chambers) treatments for 2 years in a rich fen located just outside the Bonanza Creek Experimental Forest in interior Alaska. The drought (lowered water table position) treatment was a weak sink or small source of atmospheric CO₂ compared to the moderate atmospheric CO₂ sink at our control. This change in net ecosystem exchange was due to lower gross primary production and light-saturated photosynthesis rather than increased ecosystem respiration. The flooded (raised water table position) treatment was a greater CO₂ sink in 2006 due largely to increased early season gross primary production and higher light-saturated photosynthesis. Although flooding did not have substantial effects on rates of ecosystem respiration, this water table treatment had lower maximum respiration rates and a higher temperature sensitivity of ecosystem respiration than the control plot. Surface soil warming increased both ecosystem respiration and gross primary production by approximately 16% compared to control (ambient temperature) plots, with no net effect on net ecosystem exchange. Results from this rich fen manipulation suggest that fast responses to drought will include reduced ecosystem C storage driven by plant stress, whereas inundation will increase ecosystem C storage by stimulating plant growth.     

Spatial and Temporal Variability in Growing-Season Net Ecosystem Carbon Dioxide Exchange at a Large Peatland in Ontario,Canada

We measured net ecosystem exchange of carbon dioxide (CO2) (NEE) during wet and dry summers (2000 and 2001) across a range of plant communities at Mer Bleue, a large peatland near Ottawa, southern Ontario, Canada. Wetland types included ombrotrophic bog hummocks and hollows, mineral-poor fen, and beaver pond margins. NEE was significantly different among the sites in both years, but rates of gross photosynthesis did not vary spatially even though species composition at the sites was variable. Soil respiration rates were very different across sites and dominated interannual variability in summer NEE within sites. During the dry summer of 2001, net CO2 uptake was significantly smaller, and most locations switched from a net sink to a source of CO2 under a range of levels of photosynthetically active radiation (PAR). The wetter areas--poor fen and beaver pond margin--had the largest rates of CO2 uptake and smallest rates of respiratory loss during the dry summer. Communities dominated by ericaceous shrubs (bog sites) maintained similar rates of gross photosynthesis between years; by contrast, the sedge-dominated areas (fen sites) showed signs of early senescence under drought conditions. Water table position was the strongest control on respiration in the drier summer, whereas surface peat temperature explained most of the variability in the wetter summer. Q 10 temperature-respiration quotients averaged 1.6 to 2.2. The ratio between maximum photosynthesis and respiration ranged from 3.7:1 in the poor fen to 1.2:1 at some bog sites; it declined at all sites in the drier summer owing to greater respiration rates relative to photosynthesis in evergreen shrub sites and a change in both processes in sedge sites. Our ability to predict ecosystem responses to changing climate depends on a more complete understanding of the factors that control NEE across a range of peatland plant communities.     

马占相思人工林净二氧化碳交换和碳同位素通量

应用稳定碳同位素技术,对马占相思人工林冠层受光和遮荫叶片的碳同化率(A_net)和叶面积指数(L)进行加权,将叶片水平的¹³C甄别率(Δ_i)扩展至冠层光合甄别率(Δ_canopy),测定光合固定和呼吸释放的碳同位素通量及其净交换通量.结果表明：Δ_canopy的日变化明显,日出前和中午出现较低值(18.47‰和19.87‰),而日落前达到最大(21.21‰);秋季末期(11月)至翌年夏季,Δ_canopy逐步升高,年平均为(20.37±0.29)‰.不同季节自养呼吸(日间叶片呼吸除外)和异养呼吸释放CO₂的碳同位素比率(δ¹³C)平均值分别为(-28.70±0.75)‰和(-26.75±1.3)‰,春季林冠夜间呼吸CO₂的δ¹³C最低(-30.14‰),秋季末期最高(-28.01‰).马占相思林与大气的CO₂碳同位素通量在春季和夏季中午时峰值分别为178.5和217 μmol·m^-2 ·s^-1·‰,日均值分别为638.4 和873.2 μmol·m^-2·s^-1·‰.冠层叶片吸收CO₂的碳同位素通量较呼吸释出CO₂的碳同位素通量高1.6～2.5倍,表明马占相思林日间吸收大量CO₂,降低空气CO₂浓度,具有改善环境的良好生态服务功能.     

Response of respiration of soybean leaves grown at ambient and elevated carbon dioxide concentrations to day-to-day variation in light and temperature under field conditions

BACKGROUND AND AIMS: Respiration is an important component of plant carbon balance, but it remains uncertain how respiration will respond to increases in atmospheric carbon dioxide concentration, and there are few measurements of respiration for crop plants grown at elevated [CO(2)] under field conditions. The hypothesis that respiration of leaves of soybeans grown at elevated [CO(2)] is increased is tested; and the effects of photosynthesis and acclimation to temperature examined. METHODS: Net rates of carbon dioxide exchange were recorded every 10 min, 24 h per day for mature upper canopy leaves of soybeans grown in field plots at the current ambient [CO(2)] and at ambient plus 350 micromol mol(-1) [CO(2)] in open top chambers. Measurements were made on pairs of leaves from both [CO(2)] treatments on a total of 16 d during the middle of the growing seasons of two years. KEY RESULTS: Elevated [CO(2)] increased daytime net carbon dioxide fixation rates per unit of leaf area by an average of 48 %, but had no effect on night-time respiration expressed per unit of area, which averaged 53 mmol m(-2) d(-1) (1.4 micromol m(-2) s(-1)) for both the ambient and elevated [CO(2)] treatments. Leaf dry mass per unit of area was increased on average by 23 % by elevated [CO(2)], and respiration per unit of mass was significantly lower at elevated [CO(2)]. Respiration increased by a factor of 2.5 between 18 and 26 degrees C average night temperature, for both [CO(2)] treatments. CONCLUSIONS: These results do not support predictions that elevated [CO(2)] would increase respiration per unit of area by increasing photosynthesis or by increasing leaf mass per unit of area, nor the idea that acclimation of respiration to temperature would be rapid enough to make dark respiration insensitive to variation in temperature between nights.     

Distinct impacts of different mammalian herbivore assemblages on arctic tundra CO2 exchange during the peak of the growing season

Herbivores play a key role in the carbon (C) cycle of arctic ecosystems, but these effects are currently poorly represented within models predicting land–atmosphere interactions under future climate change. Although some studies have examined the influence of various individual species of herbivores on tundra C sequestration, few studies have directly compared the effects of different herbivore assemblages. We measured peak growing season instantaneous ecosystem carbon dioxide (CO₂) exchange (photosynthesis, respiration and net ecosystem exchange) on replicated plots in arctic tundra which, for 14 years, have excluded different portions of the herbivore population (grazed controls, large mammals excluded, both small and large mammals excluded). Herbivory suppressed photosynthetic CO₂ uptake, but caused little change in ecosystem respiration. Despite evidence that small mammals consume a greater portion of plant biomass in these ecosystems, the effect of excluding only large herbivores was indistinguishable from that of excluding both large and small mammals. The herbivory‐induced decline in photosynthesis was not entirely attributable to a decline in leaf area but also likely reflects shifts in plant community composition and/or species physiology. One shrub species – Betula nana – accounted for only around 13% of total aboveground vascular plant biomass but played a central role in controlling ecosystem CO₂ uptake and release, and was suppressed by herbivory. We conclude that herbivores can have large effects on ecosystem C cycling due to shifts in plant aboveground biomass and community composition. An improved understanding of the mechanisms underlying the distinct ecosystem impacts of different herbivore groups will help to more accurately predict the net impacts of diverse herbivore communities on arctic C fluxes.     

The contribution of C<Subscript>3</Subscript> and C<Subscript>4</Subscript> plants to the carbon cycle of a tallgrass prairie: an isotopic approach

The photosynthetic pathway composition (C₃:C₄ mixture) of an ecosystem is an important controller of carbon exchanges and surface energy flux partitioning, and therefore represents a fundamental ecophysiological distinction. To assess photosynthetic mixtures at a tallgrass prairie pasture in Oklahoma, we collected nighttime above-canopy air samples along concentration and isotopic gradients throughout the 1999 and 2000 growing seasons. We analyzed these samples for their CO₂ concentration and carbon isotopic composition and calculated C₃:C₄ proportions with a two-source mixing model. In 1999, the C₄ percentage increased from 38% in spring (late April) to 86% in early fall (mid-September). The C₄ percentages inferred from ecosystem respiration measurements in 2000 indicate a smaller shift, from 67% in spring (early May) to 77% in mid-summer (late July). We also sampled daytime CO₂ concentration and carbon isotope gradients above the canopy to determine ecosystem discrimination against ¹³CO₂ during net uptake. These discrimination values were always lower than corresponding nighttime ecosystem respiration isotopic signatures would suggest. After accounting for the isotopic disequilibria between respiration and photosynthesis resulting from seasonal variations in the C₃:C₄ mixture, we estimated canopy photosynthetic discrimination. The C₄ percentage calculated from this approach agrees with the percentage determined from nighttime respiration for sampling periods in both growing seasons. Isotopic imbalances between photosynthesis and respiration are likely to be common in mixed C₃:C₄ ecosystems and must be considered when using daytime isotopic measurements to constrain ecosystem physiology. Given the global extent of such ecosystems, isotopic imbalances likely contribute to global variations in the carbon isotopic composition of atmospheric CO₂.     

Pan-Arctic soil moisture control on tundra carbon sequestration and plant productivity

Long-term atmospheric CO₂ concentration records have suggested a reduction in the positive effect of warming on high-latitude carbon uptake since the 1990s. A variety of mechanisms have been proposed to explain the reduced net carbon sink of northern ecosystems with increased air temperature, including water stress on vegetation and increased respiration over recent decades. However, the lack of consistent long-term carbon flux and in situ soil moisture data has severely limited our ability to identify the mechanisms responsible for the recent reduced carbon sink strength. In this study, we used a record of nearly 100 site-years of eddy covariance data from 11 continuous permafrost tundra sites distributed across the circumpolar Arctic to test the temperature (expressed as growing degree days, GDD) responses of gross primary production (GPP), net ecosystem exchange (NEE), and ecosystem respiration (ER) at different periods of the summer (early, peak, and late summer) including dominant tundra vegetation classes (graminoids and mosses, and shrubs). We further tested GPP, NEE, and ER relationships with soil moisture and vapor pressure deficit to identify potential moisture limitations on plant productivity and net carbon exchange. Our results show a decrease in GPP with rising GDD during the peak summer (July) for both vegetation classes, and a significant relationship between the peak summer GPP and soil moisture after statistically controlling for GDD in a partial correlation analysis. These results suggest that tundra ecosystems might not benefit from increased temperature as much as suggested by several terrestrial biosphere models, if decreased soil moisture limits the peak summer plant productivity, reducing the ability of these ecosystems to sequester carbon during the summer.     

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.     

Increased high‐latitude photosynthetic carbon gain offset by respiration carbon loss during an anomalous warm winter to spring transition

Arctic and boreal ecosystems play an important role in the global carbon (C) budget, and whether they act as a future net C sink or source depends on climate and environmental change. Here, we used complementary in situ measurements, model simulations, and satellite observations to investigate the net carbon dioxide (CO₂) seasonal cycle and its climatic and environmental controls across Alaska and northwestern Canada during the anomalously warm winter to spring conditions of 2015 and 2016 (relative to 2010–2014). In the warm spring, we found that photosynthesis was enhanced more than respiration, leading to greater CO₂ uptake. However, photosynthetic enhancement from spring warming was partially offset by greater ecosystem respiration during the preceding anomalously warm winter, resulting in nearly neutral effects on the annual net CO₂ balance. Eddy covariance CO₂ flux measurements showed that air temperature has a primary influence on net CO₂ exchange in winter and spring, while soil moisture has a primary control on net CO₂ exchange in the fall. The net CO₂ exchange was generally more moisture limited in the boreal region than in the Arctic tundra. Our analysis indicates complex seasonal interactions of underlying C cycle processes in response to changing climate and hydrology that may not manifest in changes in net annual CO₂ exchange. Therefore, a better understanding of the seasonal response of C cycle processes may provide important insights for predicting future carbon–climate feedbacks and their consequences on atmospheric CO₂ dynamics in the northern high latitudes.     

Carbon dioxide concentration at night affects translocation from soybean leaves

Studies have indicated that the concentration of carbon dioxide [CO2] during the dark period may influence plant dry matter accumulation. It is often suggested that these effects on growth result from effects of [CO2] on rates of respiration, but responses of respiration to [CO2] remain controversial, and connections between changes in respiration rate and altered growth rate have not always been clear. The present experiments tested whether translocation, a major consumer of energy from respiration in exporting leaves, was sensitive to [CO2]. Nineteen-day-old soybean plants grown initially at a constant [CO2] of 350 micromol mol(-1) were exposed to three consecutive nights with a [CO2] of 220-1400 micromol mol(-1), with a daytime [CO2] of 350 micromol mol(-1). Change in dry mass of the individual second, third and fourth trifoliate leaves over the 3-d period was determined, along with rates of respiration and photosynthesis of second leaves, measured by net CO2 exchange. Translocation was determined from mass balance for second leaves. Additional experiments were conducted where the [CO2] around individual leaves was controlled separately from that of the rest of the plant. Results indicated that low [CO2] at night increased both respiration and translocation and elevated [CO2] decreased both processes, to similar relative extents. The effect of [CO2] during the dark on the change in leaf mass over 3 d was largest in second leaves, where the change in mass was about 50% greater at 1400 micromol mol(-1) CO2 than at 220 micromol mol(-1) CO2. The response of translocation to [CO2] was localized in individual leaves. Results indicated that effects of [CO2] on net carbon dioxide exchange rate in the dark either caused or reflected a change in a physiologically important process which is known to depend on energy supplied by respiration. Thus, it is unlikely that the observed effects of [CO2] on respiration were artefacts of the measurement process in this case.     

DROUGHT EFFECTS ON CARBON EXCHANGE IN A SUBTROPICAL CONIFEROUS PLANTATION IN CHINA

 干旱对陆地生态系统的影响已成为全球变化研究的焦点问题之一。该研究基于生态系统过程模型——CEVSA2, 结合涡度相关通量观测, 分析了不同程度干旱对亚热带人工针叶林碳交换的影响及其关键控制因素。结果表明: 1)干旱使生态系统碳交换显著下降, 2003和2004年的干旱使得年净生态系统生产力(Net ecosystem production, NEP)相比无干旱影响情景的模拟结果分别减少了63%和47%; 2)光合和呼吸对干旱具有不同的响应, 干旱时光合的下降比呼吸更为显著, 这导致了NEP的显著下降; 3)当饱和水气压差(Vapor pressure deficit, VPD)达到1.5 kPa以上时, 生态系统的光合、呼吸和净碳吸收均开始下降, 当VPD大于2.5 kPa、土壤相对含水量(土壤含水量/土壤饱和含水量)(Relative soil water content, RSW)低于40%时, 生态系统的碳收支由碳汇转为碳源; 4)土壤干旱是造成碳交换下降的主要驱动因素, 对年NEP下降的平均贡献率为46%, 而大气干旱的贡献率仅为4%。     

Carbon Dioxide Exchange Between an Old-growth Forest and the Atmosphere

Eddy-covariance and biometeorological methods show significant net annual carbon uptake in an old-growth Douglas-fir forest in southwestern Washington, USA. These results contrast with previous assumptions that old-growth forest ecosystems are in carbon equilibrium. The basis for differences between conventional biomass-based carbon sequestration estimates and the biometeorologic estimates are discussed. Annual net ecosystem exchange was comparable to younger ecosystems at the same latitude, as quantified in the AmeriFlux program. Net ecosystem carbon uptake was significantly correlated with photosynthetically active radiation and air temperature, as well as soil moisture and precipitation. Optimum ecosystem photosynthesis occurred at relatively cool temperatures (5°–10°C). Understory and soil carbon exchange always represented a source of carbon to the atmosphere, with a strong seasonal cycle in source strength. Understory and soil carbon exchange showed a Q₁₀ temperature dependence and represented a substantial portion of the ecosystem carbon budget. The period of main carbon uptake and the period of soil and ecosystem respiration are out of phase, however, and driven by different climatic boundary conditions. The period of strongest ecosystem carbon uptake coincides with the lowest observed values of soil and ecosystem respiration. Despite the substantial contribution of soil, the overall strength of the photosynthetic sink resulted in the net annual uptake. The net uptake estimates here included two correction methods, one for advection and the other for low levels of turbulence.     

北京山区典型暖温带落叶阔叶林大气CO_2浓度、草本层光合作用及土壤释放CO_2变化特点(英文)

为探讨森林生态系统植被、土壤等不同组分与大气CO_2交换特点,利用中型同化箱(40cm×40cm×2Ocm)及红外CO_2分析仪装置对北京山区典型暖温带森林生态系统辽东栎(Quercus liaotungensisKoidz.)林草本层净光合作用、土壤释放CO_2及林外(高出林冠2m)与林内(低于林冠2m)大气CO_2变化进行测定。结果表明:夏季及秋季大气CO_2浓度分别为(323±10)μmol·mol~(-1)和(330±1)μmol·mol~(-1);在一天内连续24h的测定中,大气与林内CO_2浓度的差值最大时可分别达-46和-61μmol·mol~(-1)。夏季草本层净光合强度为(2.59±1.05)μmol CO_2·m~(-2)·s~(-1),是秋季((1.31±0.39)μmol CO_2·m~(-2)·s~(-1))的2倍;夏季土壤呼吸释放CO_2的强度明显高于秋季,分别为(5.18±0.75)μmol CO_2·m~(-2)·s~(-1)和(1.96±0.57)μmol CO_2·m~(-2)·s~(-1)。土壤释放CO_2强度与地面温度之间存在显著相关,其关系式为Y=-0.8642 0.3101X(r=0.7164,P<0.001,n=117)。大气CO_2浓度的低值及草本层光合强度高值约出现在14:00左右;而在夜间土壤释放CO_2强度增加,表现为大气CO_2浓度升高。     

Significance of cold-season respiration and photosynthesis in a subarctic heath ecosystem in Northern Sweden

While substantial cold-season respiration has been documented in most arctic and alpine ecosystems in recent years, the significance of cold-season photosynthesis in these biomes is still believed to be small. In a mesic, subartic heath during both the cold and warm season, we measured in situ ecosystem respiration and photosynthesis with a chamber technique at ambient conditions and at artificially increased frequency of freeze–thaw (FT) cycles during fall and spring. We fitted the measured ecosystem exchange rates to respiration and photosynthesis models with R²-values ranging from 0.81 to 0.85. As expected, estimated cold-season (October, November, April and May) respiration was significant and accounted for at least 22% of the annual respiratory CO₂ flux. More surprisingly, estimated photosynthesis during this period accounted for up to 19% of the annual gross CO₂ uptake, suggesting that cold-season photosynthesis partly balanced the cold-season respiratory carbon losses and can be significant for the annual cycle of carbon. Still, during the full year the ecosystem was a significant net source of 120 ± 12 g C m⁻² to the atmosphere. Neither respiration nor photosynthetic rates were much affected by the extra FT cycles, although the mean rate of net ecosystem loss decreased slightly, but significantly, in May. The results suggest only a small response of net carbon fluxes to increased frequency of FT cycles in this ecosystem.