Spatial and temporal scaling of intercellular CO2 concentration in a temperate rain forest dominated by Dacrydium cupressinum in New Zealand

Seven methods, including measurements of photosynthesis (A) and stomatal conductance (g(s)), carbon isotope discrimination, ecosystem CO2 and water vapour exchange using eddy covariance and the use of a multilayer canopy model and ecosystem Keeling plots, were employed to derive estimates of intercellular CO2 concentration (Ci) across a range of spatial and temporal scales in a low productivity rain forest ecosystem dominated by the conifer Dacrydium cupressinum Lamb. in New Zealand. Estimates of shoot and canopy Ci across temporal scales ranging from minutes to years were remarkably similar (range of 274-294 micromol mol(-1)). The gradual increase in shoot Ci with depth in the canopy was more likely attributable to decreases in A resulting from lower irradiance (Q) than to increases in g, due to changes in air saturation deficit (D). The lack of marked vertical gradients in A and g(s) at saturating Q through the canopy and the low seasonal variability in environmental conditions contributed to the efficacy of scaling Ci. However, the canopy Ci estimate calculated from the carbon isotope composition of respired ecosystem CO2 (delta13CR; 236 micromol mol(-1)) was much lower than other estimates of canopy Ci. Partitioning delta13CR into four components (soil, roots, litter and foliage) indicated root respiration as the dominant (> 50%) contributor to delta13CR. Variable time lags and differences in isotopic composition during photosynthesis and respiration make the direct estimation of canopy Ci from delta 13CR problematic.     

Diurnal, seasonal and interannual variability of carbon isotope discrimination at the canopy level in response to environmental factors in a boreal forest ecosystem

Accurate estimation of temporal and spatial variations in photosynthetic discrimination of 13C is critical to carbon cycle research. In this study, a combined ecosystem-boundary layer isotope model, which was satisfactorily validated against intensive campaign data, was used to explore the temporal variability of carbon discrimination in response to environmental driving factors in a boreal ecosystem in the vicinity of Fraserdale Tower, Ontario, Canada (49 degrees 52'30'N, 81 degrees 34'12'W). A 14 year (1990-1996 and 1998-2004) hourly CO2 concentration and meteorological record measured on this tower was used for this purpose. The 14 year mean yearly diurnal amplitude of canopy-level discrimination Delta(canopy) was computed to be 2.8 +/- 0.5 per thousand, and the overall diurnal cycle showed that the greatest Delta(canopy) values occurred at dawn and dusk, while the minima generally appeared in mid-afternoon. The average annual Delta(canopy) varied from 18.3 to 19.7 per thousand with the 14 year average of 19 +/- 0.4 per thousand. The overall seasonality of Delta(canopy) showed a gradually increasing trend from leaf emergence in May-September and with a slight decrease at the end of the growing season in October. Delta(canopy) was negatively correlated to vapour pressure deficit and air temperature across hourly to decadal timescales. A strong climatic control on stomatal regulation of ecosystem isotope discrimination was found in this study.     

Variation in the degree of coupling between delta13C of phloem sap and ecosystem respiration in two mature Nothofagus forests

Day-to-day variability in the carbon isotope composition of phloem sap (delta13Chd) and ecosystem respiratory CO2 (delta13CR) were measured to assess the tightness of coupling between canopy photosynthesis (delta13Chd) and ecosystem respiration (delta13CR) in two mature Nothofagus solandri (Hook. f.) forests in New Zealand. Abundant phloem-tapping scale insects allowed repeated, nondestructive access to stem phloem sap 1-2 m above ground. delta13Chd was compared with delta13C predicted by an environmentally driven, process-based canopy photosynthesis model. Keeling plots of within-canopy CO2 were used to estimate delta13CR. By including a lag of 3 d, there was good agreement in the timing and direction of variation in delta13Chd and predictions by the canopy photosynthesis model, suggesting that delta13Chd represents a photosynthesis-weighted, integrative record of canopy photosynthesis and conductance. Significant day-to-day variability in delta13CR was recorded at one of the two forests. At this site, delta13CR reflected variability in delta13Chd only on days with <2 mm rain. We conclude that the degree of coupling between canopy photosynthesis and ecosystem respiration varies between sites, and with environmental conditions at a single site.     

Large daily variation in 13C-enrichment of leaf-respired CO2 in two Quercus forest canopies

The use of the 13C : 12C isotopic ratio (delta13C) of leaf-respired CO2 to trace carbon fluxes in plants and ecosystems is limited by little information on temporal variations in delta13C of leaf dark-respired CO2 (delta13Cr) under field conditions. Here, we explored variability in delta13Cr and its relationship to key respiratory substrates from collections of leaf dark-respired CO2, carbohydrate extractions and gas exchange measurements over 24-h periods in two Quercus canopies. Throughout both canopies, delta13Cr became progressively 13C-enriched during the photoperiod, by up to 7%, then 13C-depleted at night relative to the photoperiod. This cycle could not be reconciled with delta13C of soluble sugars (delta13Css), starch (delta13Cst), lipids (delta13Cl), cellulose (delta13Cc) or with calculated photosynthetic discrimination (Delta). However, photoperiod progressive enrichment in delta13Cr was correlated with cumulative carbon assimilation (r2 = 0.91). We concluded that there is considerable short-term variation in delta13Cr in forest canopies, that it is consistent with current hypotheses for 13C fractionation during leaf respiration, that leaf carbohydrates cannot be used as surrogates for delta13Cr, and that diel changes in leaf carbohydrate status could be used to predict changes in delta13Cr empirically.     

Carbon isotopes in terrestrial ecosystem pools and CO2 fluxes

Stable carbon isotopes are used extensively to examine physiological, ecological, and biogeochemical processes related to ecosystem, regional, and global carbon cycles and provide information at a variety of temporal and spatial scales. Much is known about the processes that regulate the carbon isotopic composition (delta(13)C) of leaf, plant, and ecosystem carbon pools and of photosynthetic and respiratory carbon dioxide (CO(2)) fluxes. In this review, systematic patterns and mechanisms underlying variation in delta(13)C of plant and ecosystem carbon pools and fluxes are described. We examine the hypothesis that the delta(13)C of leaf biomass can be used as a reference point for other carbon pools and fluxes, which differ from the leaf in delta(13)C in a systematic fashion. Plant organs are typically enriched in (13)C relative to leaves, and most ecosystem pools and respiratory fluxes are enriched relative to sun leaves of dominant plants, with the notable exception of root respiration. Analysis of the chemical and isotopic composition of leaves and leaf respiration suggests that growth respiration has the potential to contribute substantially to the observed offset between the delta(13)C values of ecosystem respiration and the bulk leaf. We discuss the implications of systematic variations in delta(13)C of ecosystem pools and CO(2) fluxes for studies of carbon cycling within ecosystems, as well as for studies that use the delta(13)C of atmospheric CO(2) to diagnose changes in the terrestrial biosphere over annual to millennial time scales.     

Correlations Between Foliar Stable Carbon Isotope Composition and Environ-mental Factors in Desert Plant Reaumuria soongorica (Pall.) Maxim.

Leaves of 407 individuals of Reaumuria soongorica (Pall.) Maxim. collected from the major distribution areas were measured to investigate the distribution characteristics of the stable carbon isotope in this desert plant, as well as correlations between δ13C values and environmental factors. Results showed that δ13C values in R. soongorica ranged from-22.77‰. to-29.85‰. and that the mean δ13C value (-26.52‰)was higher than a previously reported δ13C value for a different desert ecosystem. This indicates that R.soongorica belongs to the C3 photosynthetic pathway and has higher water use efficiency than other species. The correlations between δ13C values and environmental factors demonstrated that the foliar δ13C values in R. soongorica increased significantly with decreasing mean annual precipitation and mean relative humidity, and decreased with decreasing duration of sunshine and evaporation. The spatial distribution trend of δ13C values in R. soongorica was not obvious and there was no significant correlation between the δ13C values and mean annual temperature. We conclude that different distribution trends in δ13C values for R. soongorica were likely caused by stomatal limitation rather than by nutrient-related changes in photosynthetic efficiency and that precipitation played an important role in the wide distribution range of R.soongorica. This pattern of δ13C values for R. soongorica reinforced that it is a super-xerophil in terms of its adaptive strategies to a desert environment.     

Temporal dynamics of the carbon isotope composition in a Pinus sylvestris stand: from newly assimilated organic carbon to respired carbon dioxide

The (13)C isotopic signature (C stable isotope ratio; delta(13)C) of CO(2) respired from forest ecosystems and their particular compartments are known to be influenced by temporal changes in environmental conditions affecting C isotope fractionation during photosynthesis. Whereas most studies have assessed temporal variation in delta(13)C of ecosystem-respired CO(2) on a day-to-day scale, not much information is available on its diel dynamics. We investigated environmental and physiological controls over potential temporal changes in delta(13)C of respired CO(2) by following the short-term dynamics of the (13)C signature from newly assimilated organic matter pools in the needles, via phloem-transported organic matter in twigs and trunks, to trunk-, soil- and ecosystem-respired CO(2). We found a strong 24-h periodicity in delta(13)C of organic matter in leaf and twig phloem sap, which was strongly dampened as carbohydrates were transported down the trunk. Periodicity reappeared in the delta(13)C of trunk-respired CO(2), which seemed to originate from apparent respiratory fractionation rather than from changes in delta(13)C of the organic substrate. The diel patterns of delta(13)C in soil-respired CO(2) are partly explained by soil temperature and moisture and are probably due to changes in the relative contribution of heterotrophic and autotrophic CO(2) fluxes to total soil efflux in response to environmental conditions. Our study shows that direct relations between delta(13)C of recent assimilates and respired CO(2) may not be present on a diel time scale, and other factors lead to short-term variations in delta(13)C of ecosystem-emitted CO(2). On the one hand, these variations complicate ecosystem CO(2) flux partitioning, but on the other hand they provide new insights into metabolic processes underlying respiratory CO(2) emission.     

Non-steady state effects in diurnal 180 discrimination by Picea sitchensis branches in the field

We report diurnal variations in 18O discrimination (18 delta) during photosynthesis (18 delta A) and respiration (18 delta R) of Picea sitchensis branches measured in branch chambers in the field. These observations were compared with predicted 18 delta (18 delta pred) based on concurrent measurements of branch gas exchange to evaluate steady state and non-steady state (NSS) models of foliage water 18O enrichment for predicting the impact of this ecosystem on the Delta 18O of atmospheric CO2. The non-steady state approach substantially improved the agreement between 18 delta pred and observed 18 delta (18 delta obs) compared with the assumption of isotopic steady state (ISS) for the Delta 18O signature of foliage water. In addition, we found direct observational evidence for NSS effects: extremely high apparent 18 delta values at dusk, dawn and during nocturnal respiration. Our experiments also show the importance of bidirectional foliage gas exchange at night (isotopic equilibration in addition to the net flux). Taken together, neglecting these effects leads to an underestimation of daily net canopy isofluxes from this forest by up to 30%. We expect NSS effects to be most pronounced in species with high specific leaf water content such as conifers and when stomata are open at night or when there is high relative humidity, and we suggest modifications to ecosystem and global models of delta 18O of CO2.     

Short-term variations in δ<Superscript>13</Superscript>C of ecosystem respiration reveals link between assimilation and respiration in a deciduous forest

We present a comprehensive dataset of hourly, daily, and monthly measurements of carbon isotope measurements of CO₂ in canopy air from a temperate deciduous forest with the aim to identify the relevance of short-term variations in the isotopic signature of ecosystem respiration (¹³C_R) and to understand its underlying physiological processes. We show that during daytime low vertical mixing inside the canopy can lead to decoupling of the air in the lower and upper canopy layer resulting in large spatial variation of ¹³C in CO₂ of canopy air. Intercept of Keeling Plots also showed large temporal variation (3.8) over the course of the day demonstrating that intercepts can differ between day and night and suggesting that choosing the right time for sampling is essential to capture the isotopic signature of ecosystem respiration (¹³C_R). ¹³C_R as obtained from night-time measurements showed large variation of up to 2.65 on a day-to-day basis, which was similar to the observed variation of ¹³C_R over the seasonal cycle (3.08). This highlights the importance of short-term physiological processes within ecosystems for the isotopic composition of CO₂ in the atmosphere, not reflected by bulk plant and soil organic samples. At daily and monthly time scales, ¹³C_R increased with increasing ratio of vapour pressure deficit to photosynthetically active radiation, measured 4–5 days before. This suggests that ecosystem respiration was isotopically linked to assimilation. Furthermore, assimilates recently fixed in the canopy seem to form a labile carbon pool with a short mean residence time that is respired back to the atmosphere after 4–5 days.     

Continuous monitoring reveals multiple controls on ecosystem metabolism in a suburban stream

1. Primary production and respiration in streams, collectively referred to as stream ecosystem metabolism, are fundamental processes that determine trophic structure, biomass and nutrient cycling. Few studies have used high‐frequency measurements of gross primary production (GPP) and ecosystem respiration (ER) over extended periods to characterise the factors that control stream ecosystem metabolism at hourly, daily, seasonal and annual scales. 2. We measured ecosystem metabolism at 5‐min intervals for 23 months in Shepherd Creek, a small suburban stream in Cincinnati, Ohio (U.S.A.). 3. Daily GPP was best predicted by a model containing light and its synergistic interaction with water temperature. Water temperature alone was not significantly related to daily GPP, rather high temperatures enhanced the capacity of autotrophs to use available light. 4. The relationship between GPP and light was further explored using photosynthesis–irradiance curves (P–I curves). Light saturation of GPP was evident throughout the winter and spring and the P–I curve frequently exhibited strong counterclockwise hysteresis. Hysteresis occurred when water temperatures were greater in the afternoon than in the morning, although light was similar, further suggesting that light availability interacts synergistically with water temperature. 5. Storm flows strongly depressed GPP in the spring while desiccation arrested aquatic GPP and ER in late summer and autumn. 6. Ecosystem respiration was best predicted by GPP, water temperature and the rate of water exchange between the surface channel and transient storage zones. We estimate that c. 70% of newly fixed carbon was immediately respired by autotrophs and closely associated heterotrophs. 7. Interannual, seasonal, daily and hourly variability in ecosystem metabolism was attributable to a combination of light availability, water temperature, storm flow dynamics and desiccation. Human activities affect all these factors in urban and suburban streams, suggesting stream ecosystem processes are likely to respond in complex ways to changing land use and climate.     

Short-term dynamics of the carbon isotope composition of CO2 emitted from a wheat agroecosystem--physiological and environmental controls

Understanding environmental and physiological controls of the variations in δ(13) C of CO(2) respired (δ(13) C(R)) from different compartments of an ecosystem is important for separation of CO(2) fluxes and to assess coupling between assimilation and respiration. In a wheat field, over 3 days we characterised the temporal dynamics of δ(13) C(R) from shoots and roots, from the soil and from the whole agroecosystem. To evaluate the basis of potential variations in δ(13) C(R), we also measured δ(13) C in different organic matter pools, as well as meteorological and gas exchange parameters. We observed strong diel variations up to ca. 6% in shoot, root and soil δ(13) C(R), but not in δ(13) C of the putative organic substrates for respiration, which varied by not more than ca. 1% within 24 h. Whole ecosystem-respired CO(2) was least depleted in (13) C in the afternoon and most negative in the early morning. We assume that temporally variable respiratory carbon isotope fractionation and changes in fluxes through metabolic pathways, rather than photosynthetic carbon isotope fractionation, governs the δ(13) C of respired CO(2) at the diel scale, and thus provides insights into the metabolic processes related to respiration under field conditions.     

Carbon balance of a European mountain bog at contrasting stages of regeneration

Carbon dioxide and methane (CH4) fluxes were measured in a cutover bog of the Jura Mountains (France) together with biotic and abiotic variables for two entire vegetation periods in order to compare the carbon balance of the bog at three stages of regeneration. Among all factors, air temperature and vegetation index (including leaf area of vascular plants, bryophyte density and bryophyte desiccation) were the two main determinants of ecosystem respiration and gross photosynthesis at light saturation. During 2004 and 2005, the vegetated plots acted as carbon sinks. Net carbon exchange ranged between 67 and 166 g C m(-2) yr(-1) for the Eriophorum-dominated plots and between 93 and 183 g C m(-2) yr(-1) for the Sphagnum-dominated plots. The bare peat plots represented a net carbon source (between -19 and -32 g C m(-2) yr(-1)). Methane fluxes accounted for a very small part of the total carbon efflux (< 2%). The recovery of vegetation in our naturally regenerating bog was beneficial for the carbon sequestration after the relatively short period of 20 yr.     

Correlations Between Foliar Stable Carbon Isotope Composition and Environmental Factors in Desert Plant Reaumuria soongorica （Pall.） Maxim.

Leaves of 407 individuals of Reaumuria soongorica （Pall.） Maxim. collected from the major distribution areas were measured to investigate the distribution characteristics of the stable carbon isotope in this desert plant, as well as correlations between δ^13C values and environmental factors. Results showed that δ^13C values in R. soongorica ranged from -22.77‰ to -29.85‰ and that the mean δ^13C value （-26.52‰） was higher than a previously reported δ^13C value for a different desert ecosystem. This indicates that R. soongorica belongs to the C3 photosynthetic pathway and has higher water use efficiency than other species. The correlations between δ^13C values and environmental factors demonstrated that the foliar δ^13C values in R. soongorica increased significantly with decreasing mean annual precipitation and mean relative humidity, and decreased with decreasing duration of sunshine and evaporation. The spatial distribution trend of δ^13C values in R. soongorica was not obvious and there was no significant correlation between the δ^13C values and mean annual temperature. We conclude that different distribution trends in δ^13C values for R. soongorica were likely caused by stomatal limitation rather than by nutrient-related changes in photosynthetic efficiency and that precipitation played an important role in the wide distribution range of R. soongorica. This pattern of δ^13C values for R. soongorica reinforced that it is a super-xerophil in terms of its adaptive strategies to a desert environment.     

Temporal variation in δ<Superscript>13</Superscript>C of ecosystem respiration in the Pacific Northwest: links to moisture stress

We measured seasonal and interannual variations in delta(13)C values within the carbon reservoirs (leaves and soil) and CO(2) fluxes (soil and ecosystem respired CO(2)) of an old growth coniferous forest in the Pacific Northwest USA with relation to local meteorological conditions. There were significant intra-annual and interannual differences in the carbon isotope ratios of CO(2) respired at both the ecosystem (delta(13)C(R)) and the soil levels (delta(13)C(R-soil)), but only limited variations in the carbon isotope ratios of carbon stocks. The delta(13)C(R) values varied by as much as 4.4 per thousand over a growing season, while delta(13)C(R-soil )values changed as much as 6.2 per thousand. The delta(13)C of soil organic carbon (delta(13)C(SOC)) and needle organic carbon (delta(13)C(P)) exhibited little or no significant changes over the course of this study. Carbon isotope discrimination within leaves (Delta(p)) showed systematic decreases with increased canopy height, but remained fairly constant throughout the year (Delta(p)=17.9 per thousand -19.2 per thousand at the top of the canopy, Delta(p)=19.6 per thousand -20.9 per thousand at mid-canopy, Delta(p)=23.3 per thousand -25.1 per thousand at the canopy base). The temporal variation in the delta(13)C of soil and ecosystem respired CO(2) was correlated ( r=0.93, P<0.001) with soil moisture levels, with dry summer months having the most (13)C-enriched values. The dynamic seasonal changes in delta(13)C of respired CO(2) are hypothesized to be the result of fast cycling of recently fixed carbon back to the atmosphere. One scaling consequence of the seasonal and interannual variations in delta(13)C(R) is that inversion-based carbon-cycle models dependent on observed atmospheric CO(2) concentration and isotope values may be improved by incorporating dynamic delta(13)C(R) values to interpret regional carbon sink strength.     

Analyzing the Ecosystem Carbon Dynamics of Four European Coniferous Forests Using a Biogeochemistry Model

This paper provides the first steps toward a regional-scale analysis of carbon (C) budgets. We explore the ability of the ecosystem model BIOME-BGC to estimate the daily and annual C dynamics of four European coniferous forests and shifts in these dynamics in response to changing environmental conditions. We estimate uncertainties in the model results that arise from incomplete knowledge of site management history (for example, successional stage of forest). These uncertainties are especially relevant in regional-scale simulations, because this type of information is difficult to obtain. Although the model predicted daily C and water fluxes reasonably well at all sites, it seemed to have a better predictive capacity for the photosynthesis-related processes than for respiration. Leaf area index (LAI) was modeled accurately at two sites but overestimated at two others (as a result of poor long-term climate drivers and uncertainties in model parameterization). The overestimation of LAI (and consequently gross photosynthetic production (GPP)), in combination with reasonable estimates of the daily net ecosystem productivity (NEP) of those forests, also illustrates the problem with modeled respiration. The model results suggest that all four European forests have been net sinks of C at the rate of 100–300 gC/m²/y and that this C sequestration capacity would be 30%–70% lower without increasing nitrogen (N) deposition and carbon dioxide (CO₂) concentrations. The magnitude of the forest responses was dependent not only on the rate of changes in environmental factors, but also on site-specific conditions such as climate and soil depth. We estimated that the modeled C exchange at the study sites was reduced by 50%–100% when model simulations were performed for climax forests rather than regrowing forests. The estimates of water fluxes were less sensitive to different initializations of state variables or environmental change scenarios than C fluxes.     

Disentangling drought-induced variation in ecosystem and soil respiration using stable carbon isotopes

Combining C flux measurements with information on their isotopic composition can yield a process-based understanding of ecosystem C dynamics. We studied the variations in both respiratory fluxes and their stable C isotopic compositions (δ¹³C) for all major components (trees, understory, roots and soil microorganisms) in a Mediterranean oak savannah during a period with increasing drought. We found large drought-induced and diurnal dynamics in isotopic compositions of soil, root and foliage respiration (δ¹³C_res). Soil respiration was the largest contributor to ecosystem respiration (R _eco), exhibiting a depleted isotopic signature and no marked variations with increasing drought, similar to ecosystem respired δ¹³CO₂, providing evidence for a stable C-source and minor influence of recent photosynthate from plants. Short-term and diurnal variations in δ¹³C_res of foliage and roots (up to 8 and 4‰, respectively) were in agreement with: (1) recent hypotheses on post-photosynthetic fractionation processes, (2) substrate changes with decreasing assimilation rates in combination with increased respiratory demand, and (3) decreased phosphoenolpyruvate carboxylase activity in drying roots, while altered photosynthetic discrimination was not responsible for the observed changes in δ¹³C_res. We applied a flux-based and an isotopic flux-based mass balance, yielding good agreement at the soil scale, while the isotopic mass balance at the ecosystem scale was not conserved. This was mainly caused by uncertainties in Keeling plot intercepts at the ecosystem scale due to small CO₂ gradients and large differences in δ¹³C_res of the different component fluxes. Overall, stable isotopes provided valuable new insights into the drought-related variations of ecosystem C dynamics, encouraging future studies but also highlighting the need of improved methodology to disentangle short-term dynamics of isotopic composition of R _eco.     

放牧与围栏羊草草原生态系统土壤呼吸作用比较

采用静态箱式法,比较分析了内蒙古典型羊草草原放牧与围栏生态系统土壤呼吸作用及其与环境因子的关系．结果表明,围栏和放牧样地的土壤呼吸作用日动态均呈单峰型曲线,高峰值一般出现在13：00～15：00,围栏样地土壤呼吸作用日动态与地表温度相关性最好,而放牧样地与地下5cm温度相关性最好．在整个观测期内,6、7月份是植物生长的旺盛期,围栏样地土壤呼吸作用明显大于放牧样地约2．7倍;到植物生长后期的8、9月份,二者差异不大,与地下生物量的变化相似,可能与牲畜的采食对不同物候期的植物影响不同及周围环境因子的改变有关,说明人类活动的干扰不一定增加土壤呼吸作用．围栏样地和放牧样地土壤呼吸作用季节动态都与0～10cm的土壤含水量相关性最好,相关系数分别是0．853和0．741,而围栏样地土壤含水量与土壤呼吸作用季节动态的相关性大于放牧样地;围栏和放牧样地不同层次土温、土壤含水量与土壤呼吸作用日、季动态的关系均表现出浅层的相关性普遍大于深层．     

Assessing environmental and physiological controls over water relations in a Scots pine (Pinus sylvestris L.) stand through analyses of stable isotope composition of water and organic matter

This study investigated the influence of meteorological, pedospheric and physiological factors on the water relations of Scots pine, as characterized by the origin of water taken up, by xylem transport as well as by carbon isotope discrimination (Delta13C) and oxygen isotope enrichment (Delta18O) of newly assimilated organic matter. For more than 1 year, we quantified delta2H and delta18O of potential water sources and xylem water as well as Delta13C and Delta18O in twig and trunk phloem organic matter biweekly, and related these values to continuously measured or modelled meteorological parameters, soil water content, stand transpiration (ST) and canopy stomatal conductance (G(s)). During the growing season, delta18O and delta2H of xylem water were generally in a range comparable to soil water from a depth of 2-20 cm. Long residence time of water in the tracheids uncoupled the isotopic signals of xylem and soil water in winter. Delta18O but not Delta13C in phloem organic matter was directly indicative of recent environmental conditions during the whole year. Delta18O could be described applying a model that included 18O fractionation associated with water exchange between leaf and atmosphere, and with the production of organic matter as well as the influence of transpiration. Phloem Delta13C was assumed to be concertedly influenced by G(s) and photosynthetically active radiation (PAR) (as a proxy for photosynthetic capacity). We conclude that isotope signatures can be used as effective tools (1) to characterize the seasonal dynamics in source and xylem water, and (2) to assess environmental effects on transpiration and G(s) of Scots pine, thus helping to understand and predict potential impacts of climate change on trees and forest ecosystems.     

Seasonal and daily time course of the 13C composition in soil CO2 efflux recorded with a tunable diode laser spectrophotometer (TDLS)

Temporal variations of carbon isotope composition of soil CO₂ efflux (F_S and δ¹³C_FS) at different time scales should reflect both temporal variations of the climate conditions that affect canopy functioning and temporal changes in the relative contribution of autotrophic respiration to total F_S. A tunable diode laser spectrophotometer (TDLS) was installed in the Hesse forest (northeast of France) early during the 2007 growing season to determine the seasonal and daily variability in δ¹³C_FS. This method, based on the measurement of the absorption of an infrared laser emission at specific wave lengths of the ¹³CO₂ and ¹²CO₂, allows the continuous monitoring of the two isotopologues. The concentrations of the two isotopologues in F_S were continuously monitored from June to November 2007 using chamber method and Keeling plots drawn from nocturnal accumulation of CO₂ below the canopy. These TDLS measurements and isotope ratio mass spectrometer based Keeling plots gave very similar values of δ¹³C_FS, showing the reliability of the TDLS system in this context. Results were analysed with regard to seasonal and daily changes in climatic and edaphic variables and compared with the δ¹³C of CO₂ respired by roots, litter and soil incubated under controlled conditions. Pronounced daily as well as seasonal variations in δ¹³C_FS were recorded (up to 1.5‰). The range of variation of δ¹³C_FS was of the same order of magnitude at both diurnal and seasonal scales. δ¹³C_FS observed in the field fluctuated between values of litter and of root respiration recorded during incubation, suggesting that temporal (and probably spatial) variations were associated with changes in the relative contribution of the two compartments during the day and during the season.     

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.