Canopy-scale δ13C of photosynthetic and respiratory CO2 fluxes: observations in forest biomes across the United States

The δ¹³C values of atmospheric carbon dioxide (CO₂) can be used to partition global patterns of CO₂ source/sink relationships among terrestrial and oceanic ecosystems using the inversion technique. This approach is very sensitive to estimates of photosynthetic ¹³C discrimination by terrestrial vegetation (Δ_A), and depends on δ¹³C values of respired CO₂ fluxes (δ¹³C_R). Here we show that by combining two independent data streams – the stable isotope ratios of atmospheric CO₂ and eddy‐covariance CO₂ flux measurements – canopy scale estimates of Δ_A can be successfully derived in terrestrial ecosystems. We also present the first weekly dataset of seasonal variations in δ¹³C_R from dominant forest ecosystems in the United States between 2001 and 2003. Our observations indicate considerable summer‐time variation in the weekly value of δ¹³C_R within coniferous forests (4.0‰ and 5.4‰ at Wind River Canopy Crane Research Facility and Howland Forest, respectively, between May and September). The monthly mean values of δ¹³C_R showed a smaller range (2–3‰), which appeared to significantly correlate with soil water availability. Values of δ¹³C_R were less variable during the growing season at the deciduous forest (Harvard Forest). We suggest that the negative correlation between δ¹³C_R and soil moisture content observed in the two coniferous forests should represent a general ecosystem response to the changes in the distribution of water resources because of climate change. Shifts in δ¹³C_R and Δ_A could be of sufficient magnitude globally to impact partitioning calculations of CO₂ sinks between oceanic and terrestrial compartments.     

Comparison of ecosystem water-use efficiency among Douglas-fir forest, aspen forest and grassland using eddy covariance and carbon isotope techniques

Comparisons were made among Douglas‐fir forest, aspen (broad leaf deciduous) forest and wheatgrass (C₃) grassland for ecosystem‐level water‐use efficiency (WUE). WUE was defined as the ratio of photosynthetic CO₂ assimilation rate and evapotranspiration (ET) rate. The ET data measured by eddy covariance were screened so that they overwhelmingly represented transpiration. The three sites used in this comparison spanned a range of vegetation (plant functional) types and environmental conditions within western Canada. When compared in the relative order Douglas‐fir (located on Vancouver Island, BC), aspen (northern Saskatchewan), grassland (southern Alberta), the sites demonstrated a progressive decline in precipitation and a general increase in maximum air temperature and atmospheric saturation deficit (D_max) during the mid‐summer. The average (±SD) WUE at the grassland site was 2.6±0.7 mmol mol^?1, which was much lower than the average values observed for the two other sites (aspen: 5.4±2.3, Douglas‐fir: 8.1±2.4). The differences in WUE among sites were primarily because of variation in ET. The highest maximum ET rates were approximately 5, 3.2 and 2.7 mm day^?1 for the grassland, aspen and Douglas‐fir sites, respectively. There was a strong negative correlation between WUE and D_max for all sites. We also made seasonal measurements of the carbon isotope ratio of ecosystem respired CO₂ (δ_R) in order to test for the expected correlation between shifts in environmental conditions and changes to the ecosystem‐integrated ratio of leaf intercellular to ambient CO₂ concentration (c_i/c_a). There was a consistent increase in δ_R values in the grassland, aspen forest and Douglas‐fir forest associated with a seasonal reduction in soil moisture. Comparisons were made between WUE measured using eddy covariance with that calculated based on D and δ_R measurements. There was excellent agreement between WUE values calculated using the two techniques. Our δ_R measurements indicated that c_i/c_a values were quite similar among the Douglas‐fir, aspen and grassland sites, despite large variation in environmental conditions among sites. This implied that the shorter‐lived grass species had relatively high c_i/c_a values for the D of their habitat. By contrast, the longer‐lived Douglas‐fir trees were more conservative in water‐use with lower c_i/c_a values relative to their habitat D. This illustrates the interaction between biological and environmental characteristics influencing ecosystem‐level WUE. The strong correlation we observed between the two independent measurements of WUE, indicates that the stable isotope composition of respired CO₂ is a useful ecosystem‐scale tool to help study constraints to photosynthesis and acclimation of ecosystems to environmental stress.     

The role of interannual,seasonal, and synoptic climate on the carbon isotope ratio of ecosystem respiration at a semiarid woodland

The terrestrial carbon cycle is influenced by environmental variability at scales ranging from diurnal to interannual. Here, we present 5‐years of growing season (day 131–275) observations of the carbon isotope ratio of ecosystem respiration (δ¹³C_R) from a semiarid woodland. This ecosystem has a large necromass component resulting from 97%Pinus edulis mortality in 2002, is dominated by drought‐tolerant Juniperus monosperma trees, and experiences large variability in the timing and intensity of seasonal and synoptic water availability. Mean growing season δ¹³C_R was remarkably invariant (?23.57±0.4‰), with the exception of particularly enriched δ¹³C_R in 2006 following a winter with anomalously low snowfall. δ¹³C_R was strongly coupled to climate during premonsoon periods (～May to June), including fast (≤2 days) responses to changes in crown‐level stomatal conductance (G_c) and vapor pressure deficit (vpd) following rain pulses. In contrast, δ¹³C_R was relatively decoupled from G_c and environmental drivers during monsoon and postmonsoon periods (July–August and September, respectively), exhibiting only infrequent couplings of δ¹³C_R to vpd and soil water content (SWC) with longer lags (～8 days) and variable response slopes (both positive and negative). Notably, δ¹³C_R exhibited consistent dynamics after rainfall events, with depleted δ¹³C_R occurring within 1 h, progressive hourly δ¹³C_R enrichment over the remainder of the night, and net δ¹³C_R depletions over the multiple nights postevent in monsoon and postmonsoon periods. Overall this ecosystem demonstrated strong dependence of δ¹³C_R on precipitation, with an apparent dominance by the autotrophic δ¹³C signal in premonsoon periods when deep soil moisture is abundant and surface soil moisture is low, and weaker coupling during monsoonal periods consistent with increasing heterotrophic dominance when deep soil moisture has declined and surface moisture is variable.     

Combining meteorology, eddy fluxes, isotope measurements, and modeling to understand environmental controls of carbon isotope discrimination at the canopy scale

Estimates of terrestrial carbon isotope discrimination are useful to quantify the terrestrial carbon sink. Carbon isotope discrimination by terrestrial ecosystems may vary on seasonal and interannual time frames, because it is affected by processes (e.g. photosynthesis, stomatal conductance, and respiration) that respond to variable environmental conditions (e.g. air humidity, temperature, light). In this study, we report simulations of the temporal variability of canopy‐scale C₃ photosynthetic carbon isotope discrimination obtained with an ecophysiologically based model (ISOLSM) designed for inclusion in global models. ISOLSM was driven by half‐hourly meteorology, and parameterized with eddy covariance measurements of carbon and energy fluxes and foliar carbon isotope ratios from a pine forest in Metolius (OR). Comparing simulated carbon and energy fluxes with observations provided a range of parameter values that optimized the simulated fluxes. We found that the sensitivity of photosynthetic carbon isotope discrimination to the slope of the stomatal conductance equation (m, Ball–Berry constant) provided an additional constraint to the model, reducing the wide parameter space obtained from the fluxes alone. We selected values of m that resulted in similar simulated long‐term discrimination as foliar isotope ratios measured at the site. The model was tested with ¹³C measurements of ecosystem (δ_R) and foliar (δ_f) respiration. The daily variability of simulated ¹³C values of assimilated carbon (δ_A) was similar to that of observed δ_f, and higher than that of observed and simulated δ_R. We also found similar relationships between environmental factors (i.e. vapor pressure deficit) and simulated δ_R as measured in ecosystem surveys of δ_R. Therefore, ISOLSM reasonably simulated the short‐term variability of δ_A controlled by atmospheric conditions at the canopy scale, which can be useful to estimate the variability of terrestrial isotope discrimination. Our study also shows that including the capacity to simulate carbon isotope discrimination, together with simple ecosystem isotope measurements, can provide a useful constraint to land surface and carbon balance models.     

Seasonal dynamics in the stable carbon isotope composition δ¹³C from non-leafy branch, trunk and coarse root CO₂ efflux of adult deciduous (Fagus sylvatica) and evergreen (Picea abies) trees

Respiration is a substantial driver of carbon (C) flux in forest ecosystems and stable C isotopes provide an excellent tool for its investigation. We studied seasonal dynamics in δ¹³C of CO₂ efflux (δ¹³C_E) from non‐leafy branches, upper and lower trunks and coarse roots of adult trees, comparing deciduous Fagus sylvatica (European beech) with evergreen Picea abies (Norway spruce). In both species, we observed strong and similar seasonal dynamics in the δ¹³C_E of above‐ground plant components, whereas δ¹³C_E of coarse roots was rather stable. During summer, δ¹³C_E of trunks was about ?28.2‰ (Beech) and ?26.8‰ (Spruce). During winter dormancy, δ¹³C_E increased by 5.6–9.1‰. The observed dynamics are likely related to a switch from growth to starch accumulation during fall and remobilization of starch, low TCA cycle activity and accumulation of malate by PEPc during winter. The seasonal δ¹³C_E pattern of branches of Beech and upper trunks of Spruce was less variable, probably because these organs were additionally supplied by winter photosynthesis. In view of our results and pervious studies, we conclude that the pronounced increases in δ¹³C_E of trunks during the winter results from interrupted access to recent photosynthates.     

Environmental and physiological controls on the carbon isotope composition of CO2 respired by leaves and roots of a C3 woody legume (Prosopis velutina) and a C4 perennial grass (Sporobolus wrightii)

Accurate estimates of the δ¹³C value of CO₂ respired from roots (δ¹³C_{R_root}) and leaves (δ¹³C_{R_leaf}) are important for tracing and understanding changes in C fluxes at the ecosystem scale. Yet the mechanisms underlying temporal variation in these isotopic signals are not fully resolved. We measured δ¹³C_{R_leaf}, δ¹³C_{R_root}, and the δ¹³C values and concentrations of glucose and sucrose in leaves and roots in the C₄ grass Sporobolus wrightii and the C₃ tree Prosopis velutina in a savanna ecosystem in southeastern Arizona, USA. Night‐time variation in δ¹³C_{R_leaf} of up to 4.6 ± 0.6‰ in S. wrightii and 3.0 ± 0.6‰ in P. velutina were correlated with shifts in leaf sucrose concentration, but not with changes in δ¹³C values of these respiratory substrates. Strong positive correlations between δ¹³C_{R_root} and root glucose δ¹³C values in P. velutina suggest large diel changes in δ¹³C_{R_root} (were up to 3.9‰) influenced by short‐term changes in δ¹³C of leaf‐derived phloem C. No diel variation in δ¹³C_{R_root} was observed in S. wrightii. Our findings show that short‐term changes in δ¹³C_{R_leaf} and δ¹³C_{R_root} were both related to substrate isotope composition and concentration. Changes in substrate limitation or demand for biosynthesis may largely control short‐term variation in the δ¹³C of respired CO₂ in these species.     

Contributions of evaporation, isotopic non-steady state transpiration and atmospheric mixing on the delta18O of water vapour in Pacific Northwest coniferous forests

Changes in the ²H and ¹⁸O of atmospheric water vapour provide information for integrating aspects of gas exchange within forest canopies. In this study, we show that diurnal fluctuations in the oxygen isotope ratio (δ¹⁸O) as high as 4‰ were observed for water vapour (δ¹⁸O_vp) above and within an old‐growth coniferous forest in the Pacific Northwest region of the United States. Values of δ¹⁸O_vp decreased in the morning, reached a minimum at midday, and recovered to early‐morning values in the late afternoon, creating a nearly symmetrical diurnal pattern for two consecutive summer days. A mass balance budget was derived and assessed for the ¹⁸O of canopy water vapour over a 2‐d period by considering the ¹⁸O‐isoflux of canopy transpiration, soil evaporation and the air entering the canopy column. The budget was used to address two questions: (1) do δ¹⁸O values of canopy water vapour reflect the biospheric influence, or are such signals swamped by atmospheric mixing? and (2) what mechanisms drive temporal variations of δ¹⁸O_vp? Model calculations show that the entry of air into the canopy column resulted in an isotopically depleted ¹⁸O‐isoflux in the morning of day 1, causing values of δ¹⁸O_vp to decrease. An isotopically enriched ¹⁸O‐isoflux resulting from transpiration then offset this decreased δ¹⁸O_vp later during the day. Contributions of ¹⁸O‐isoflux from soil evaporation were relatively small on day 1 but were more significant on day 2, despite the small H₂¹⁶O fluxes. From measurements of leaf water volume and sapflux, we determined the turnover time of leaf water in the needles of Douglas‐fir trees as ≈ 11 h at midday. Such an extended turnover time suggests that transpiration may not have occurred at the commonly assumed isotopic steady state. We tested a non‐steady state model for predicting δ¹⁸O of leaf water. Our model calculations show that assuming isotopic steady state increased isoflux of transpiration. The impact of this increase on the modelled δ ¹⁸O_vp was clearly detectable, suggesting the importance of considering isotopic non‐steady state of transpiration in studies of forest ¹⁸O water balance.     

Modeling dynamics of stable carbon isotopic exchange between a boreal forest ecosystem and the atmosphere

Stable isotopes of CO₂ contain unique information on the biological and physical processes that exchange CO₂ between terrestrial ecosystems and the atmosphere. In this study, we developed an integrated modeling system to simulate dynamics of stable carbon isotope of CO₂, as well as moisture, energy, and momentum, between a boreal forest ecosystem and the atmosphere, as well as their transport/mixing processes through the convective boundary layer (CBL), using remotely sensed surface parameters to characterize the surface heterogeneity. It has the following characteristics: (i) it accounts for the influences of the CBL turbulent mixing and entrainment of the air aloft; (ii) it scales individual leaf‐level photosynthetic discrimination up to the whole canopy (Δ_canopy) through the separation of sunlit and shaded leaf groups; (iii) it has the capacity to examine the detailed interrelationships among plant water‐use efficiency, isotope discrimination, and vapor pressure deficit; and (iv) it has the potential to investigate how an ecosystem discriminates against ¹³C at various time and spatial scales. The monthly mean isotopic signatures of ecosystem respiration (i.e. δ¹³C_R) used for isotope flux calculation are retrieved from the nighttime flask data from the intensive campaigns (1998–2000) at 20 m level on Fraserdale tower, and the data from the growing season in 1999 are used for model validation. Both the simulated CO₂ mixing ratio and δ¹³C of CO₂ at the 20 m level agreed with the measurements well in different phases of the growing season. On a diurnal basis, the greatest photosynthetic discrimination at canopy level (i.e. Δ_canopy) occurred early morning and late afternoon with a varying range of 10–26‰. The diurnal variability of Δ_canopy was also associated with the phases of growing season and meteorological variables. The annual mean Δ_canopy in 1999 was computed to be 19.58‰. The monthly averages of Δ_canopy varied between 18.55‰ and 20.84‰ with a seasonal peak during the middle growing season. Because of the strong opposing influences of respired and photosynthetic fluxes on forest air (both CO₂ and ¹³CO₂) on both the diurnal and seasonal time scales, CO₂ was consistently enriched with the heavier ¹³C isotope (less negative δ¹³C) from July to October and depleted during the remaining months, whereas on a diurnal basis, CO₂ was enriched with the heavier ¹³C in the late afternoon and depleted in early morning. For the year 1999, the model results reveal that the boreal ecosystem in the vicinity of Fraserdale tower was a small sink with net uptake of 29.07 g ¹²C m^?2 yr^?1 and 0.34 g ¹³C m^?2 yr^?1.     

Carbon isotopic composition of lipid biomarkers from an endoevaporitic gypsum crust microbial mat reveals cycling of mineralized organic carbon

Microbial mats that inhabit gypsum deposits in ponds at Guerrero Negro, Baja California Sur, Mexico, developed distinct pigmented horizons that provided an opportunity to examine the fixation and flow of carbon through a trophic structure and, in conjunction with previous phylogenetic analyses, to assess the diagenetic fates of molecular δ¹³C biosignatures. The δ¹³C values of individual biomarker lipids, total carbon, and total organic carbon (TOC) were determined for each of the following horizons: tan‐orange (TO) at the surface, green (G), purple (P), and olive‐black (OB) at the bottom. δ¹³C of individual fatty acids from intact polar lipids (IPFA) in TO were similar to δ¹³C of dissolved inorganic carbon (DIC) in the overlying water column, indicating limited discrimination by cyanobacteria during CO₂ fixation. δ¹³C_TOC of the underlying G was 3‰ greater than that of TO. The most δ¹³C‐depleted acetogenic lipids in the upper horizons were the cyanobacterial biomarkers C₁₇ n‐alkanes and polyunsaturated fatty acids. Bishomohopanol was 4 to 7‰ enriched, relative to alkanes and intact polar fatty acids (IPFA), respectively. Acyclic C₂₀ isoprenoids were depleted by 14‰ relative to bishomohopanol. Significantly, ?[δ¹³C_TOC ? δ¹³C_∑IPFA] increased from 6.9‰ in TO to 14.7‰ in OB. This major trend might indicate that ¹³C‐enriched residual organic matter accumulated at depth. The permanently anoxic P horizon was dominated by anoxygenic phototrophs and sulfate‐reducing bacteria. P hosted an active sulfur‐dependent microbial community. IPFA and bishomohopanol were ¹³C‐depleted relative to upper crust by 7 and 4‰, respectively, and C₂₀ isoprenoids were somewhat ¹³C‐enriched. Synthesis of alkanes in P was evidenced only by ¹³C‐depleted n‐octadecane and 8‐methylhexadecane. In OB, the marked increase of total inorganic carbon δ¹³C (δ¹³C_TIC) of >6‰ perhaps indicated terminal mineralization. This δ¹³C_TIC increase is consistent with degradation of the osmolyte glycine betaine by methylotrophic methanogens and loss of ¹³C‐depleted methane from the mat.     

The intramolecular ¹³C-distribution in ethanol reveals the influence of the CO₂ -fixation pathway and environmental conditions on the site-specific ¹³C variation in glucose

Efforts to understand the cause of ¹²C versus ¹³C isotope fractionation in plants during photosynthesis and post‐photosynthetic metabolism are frustrated by the lack of data on the intramolecular ¹³C‐distribution in metabolites and its variation with environmental conditions. We have exploited isotopic carbon‐13 nuclear magnetic resonance (¹³C NMR) spectrometry to measure the positional isotope composition (δ¹³C_i, ‰) in ethanol samples from different origins: European wines, liquors and sugars from C₃, C₄ and crassulacean acid metabolism (CAM) plants. In C₃‐ethanol samples, the methylene group was always ¹³C‐enriched (～2‰) relative to the methyl group. In wines, this pattern was correlated with both air temperature and δ¹⁸O of wine water, indicating that water vapour deficit may be a critical defining factor. Furthermore, in C₄‐ethanol, the reverse relationship was observed (methylene‐C relatively ¹³C‐depleted), supporting the concept that photorespiration is the key metabolic process leading to the ¹³C distribution in C₃‐ethanol. By contrast, in CAM‐ethanol, the isotopic pattern was similar to but stronger than C₃‐ethanol, with a relative ¹³C‐enrichment in the methylene‐C of up to 13‰. Plausible causes of this ¹³C‐pattern are briefly discussed. As the intramolecular δ¹³C_i‐values in ethanol reflect that in source glucose, our data point out the crucial impact on the ratio of metabolic pathways sustaining glucose synthesis.     

Elemental turnover rates and trophic discrimination in juvenile Lebranche mullet Mugil liza under experimental conditions

The aim of this study was to determine the isotopic‐turnover rate (R_IT) and trophic‐discrimination factor (F_TD) in muscle tissues of Lebranche mullet Mugil liza fed an experimental diet (δ¹³C = ?27·1‰; δ¹⁵N = 1·0‰). Juvenile M. liza exhibited a relatively fast R_IT, with a half‐life (t₅₀) of only 16 and 14 days for δ¹³C and δ¹⁵N respectively and a nearly complete isotopic turnover (t₉₅) of 68 and 60 days for δ¹³C and δ¹⁵N.     

Seasonal variation in δ13C and δ18O of cellulose from growth rings of Pinus radiata

Seasonal variation in δ¹³C and δ¹⁸O of cellulose (δ¹³C_c and δ¹⁸O_c) was measured within two annual rings of Pinus radiata growing at three sites in New Zealand. In general, both δ¹³C_c and δ¹⁸O_c increased to a peak over summer. The three sites differed markedly in annual water balance, and these differences were reflected in δ¹³C_c and δ¹⁸O_c. Average δ¹³C_c and δ¹⁸O_c from each site were positively related, so that the driest site had the most enriched cellulose. δ¹³C_c and δ¹⁸O_c were also related within each site, although both the slope and the closeness of fit of the relationship varied between sites. Supporting the theory, the site with the lowest average relative humidity also had the greatest change in δ¹⁸O_c‰ change in δ¹³C_c. Specific climatic events, such as drought or high rainfall, were recorded as a peak or a trough in enrichment, respectively. These results suggest that seasonal and between‐site variation in δ¹³C_c and δ¹⁸O_c are driven by the interaction between variation in climatic conditions and soil water availability, and plant response to this variation.     

δ 13C variation of C3 and C4 plants across an Asian monsoon rainfall gradient in arid northwestern China

We have investigated carbon isotopic compositions of four plant genus/species, Bothriochloa ischaemum (C₄), Stipa bungeana (C₃), Lespedeza sp. (C₃) and Heteropappus less (C₃), along a precipitation gradient in northwest China in order to assess the impact of water availability on the carbon isotopic discrimination against ¹³C during carbon assimilation in this area. This information is necessary for reconstruction of paleovegetation, particularly paleo‐C₃/C₄ plant ratios using δ¹³C value of organic matter in loess and paleosols in the Chinese Loess Plateau. The δ¹³C of C₃ plants, as a group, exhibits a negative correlation with the annual precipitation amount with a total change and sensitivity of 5‰ and ?1.1‰/100 mm, respectively, for the precipitation range from 200 to 700 mm. The C₄ grass, B. ischaemum responds to aridity by decreasing 1.7‰ for over the precipitation range from 350 to 700 mm; the plant δ¹³C is significantly correlated with annual precipitation with a slope ?0.61‰/100 mm. This result implies that without considering the effect of water availability on the plant δ¹³C values, reconstruction of percent C₄ vegetation during the last glaciation can be overestimated by about a factor of two.     

Hair of grazing cattle provides an integrated measure of the effects of site conditions and interannual weather variability on δ13C of temperate humid grassland

The carbon isotope composition (δ¹³C) of C3 ecosystems is sensitive to water availability, and provides important information for the assessment of terrestrial carbon (C) sink/source activity. Here, we report the effects of plant available soil water (PAW) on community ¹³C signatures of temperate humid grassland. The 5‐year study was conducted on pastures exhibiting a large range of PAW capacity that were located on two site types: peat and mineral soils. The data set included the centennial drought year 2003, and data from wet years (2000 and 2002). Seasonal variation of PAW was modeled using PAW capacity of each pasture, precipitation inputs and evapotranspiration estimates. Community ¹³C signatures were derived from the δ¹³C of vegetation and segments of tail switch hair of cattle grown while grazing pastures. Hair ¹³C signatures provided an assimilation‐weighted ¹³C signal that integrated both spatial (paddock‐scale) and temporal (grazing season) variation of ¹³C signatures on a pasture. The δ¹³C of hair and vegetation increased with decreasing modeled PAW in the same way on mineral and peat soils. But, at a given PAW, the δ¹³C of hair was 2.6‰ less negative than that of vegetation, reflecting the diet‐hair isotopic shift. Furthermore, the δ¹³C of hair and vegetation on peat soil pastures was 0.5‰ more negative than on pastures situated on mineral soil. This may have resulted from a ～10 ppm CO₂ enrichment of canopy air derived from ongoing peat mineralization. Community‐scale season‐mean ¹³C discrimination (Δ) exhibited a saturation‐type response towards season‐mean modeled PAW (r²=0.78), and ranged between 19.8‰ on soils with low PAW capacity during the drought year of 2003, and 21.4‰ on soils with high PAW capacity in a wet year. This indicated relatively small variation in season‐mean assimilation‐weighted p_i/p_a (0.68–0.75) between contrasting sites and years. However, this range is similar to that reported in other studies, which encompass the range from subtropical arid to humid temperate grassland. Furthermore, the tight relationship between season‐mean Δ and modeled mean PAW suggests that PAW may be used as proxy for Δ.     

Site‐specific responses to short‐term environmental variation are reflected in leaf and phloem‐sap carbon isotopic abundance of field grown Eucalyptus globulus

The carbon isotopic composition (δ¹³C) of plant material has been used extensively as an indirect measure of carbon fixation per volume of water used. More recently, the δ¹³C of phloem sap (δ¹³C_phl) has been used as a surrogate measure of short‐term, canopy scale δ¹³C. Using a combination of δ¹³C physiological, structural and chemical indices from leaves and phloem sap of Eucalyptus globulus at sites of contrasting water availability, we sought to identify short‐term, canopy scale resource limitations. Results illustrate that δ¹³C_phl offers valid reflections of short‐term, canopy scale values of leaf δ¹³C and tree water status. Under conditions limited by water, leaf and phloem sap photoassimilates differ in ¹³C abundance of a magnitude large enough to significantly influence predictions of water use efficiency. This pattern was not detected among trees with adequate water supply indicating fractionation into heterotrophic tissues that may be sensitive to plant water status. Trees employed a range of physiological, biochemical and structural adaptations to acclimate to resource limitation that differed among sites providing a useful context upon which to interpret patterns in δ¹³C. Our results highlight that such easily characterized properties are ideal for use as minimally invasive tools to monitor growth and resilience of plants to variations in resource availability.     

Carbon isotopic composition of forest soil respiration in the decade following bark beetle and stem girdling disturbances in the Rocky Mountains

Bark beetle outbreaks are widespread in western North American forests, reducing primary productivity and transpiration, leading to forest mortality across large areas and altering ecosystem carbon cycling. Here the carbon isotope composition (δ¹³C) of soil respiration (δ_J) was monitored in the decade after disturbance for forests affected naturally by mountain pine beetle infestation and artificially by stem girdling. The seasonal mean δ_J changed along both chronosequences. We found (a) enrichment of δ_J relative to controls (<1 ‰) in near‐surface soils in the first 2 years after disturbance; (b) depletion (1‰ or no change) during years 3–7; and (c) a second period of enrichment (1–2‰) in years 8–10. Results were consistent with isotopic patterns associated with the gradual death and decomposition of rhizosphere organisms, fine roots, conifer needles and woody roots and debris over the course of a decade after mortality. Finally, δ_J was progressively more ¹³C‐depleted deeper in the soil than near the surface, while the bulk soil followed the well‐established pattern of ¹³C‐enrichment at depth. Overall, differences in δ_J between mortality classes (<1‰) and soil depths (<3‰) were smaller than variability within a class or depth over a season (up to 6‰).     

Pan-European δ13C values of air and organic matter from forest ecosystems

We present carbon stable isotope, δ¹³C, results from air and organic matter samples collected during 98 individual field campaigns across a network of Carboeuroflux forest sites in 2001 (14 sites) and 2002 (16 sites). Using these data, we tested the hypothesis that δ¹³C values derived from large‐scale atmospheric measurements and models, which are routinely used to partition carbon fluxes between land and ocean, and potentially between respiration and photosynthesis on land, are consistent with directly measured ecosystem‐scale δ¹³C values. In this framework, we also tested the potential of δ¹³C in canopy air and plant organic matter to record regional‐scale ecophysiological patterns. Our network estimates for the mean δ¹³C of ecosystem respired CO₂ and the related ‘discrimination’ of ecosystem respiration, δ_er and Δ_er, respectively, were ?25.6±1.9‰ and 17.8 ±2.0‰ in 2001 and ?26.6±1.5‰ and 19.0±1.6‰ in 2002. The results were in close agreement with δ¹³C values derived from regional‐scale atmospheric measurement programs for 2001, but less so in 2002, which had an unusual precipitation pattern. This suggests that regional‐scale atmospheric sampling programs generally capture ecosystem δ¹³C signals over Europe, but may be limited in capturing some of the interannual variations. In 2001, but less so in 2002, there were discernable longitudinal and seasonal trends in δ_er. From west to east, across the network, there was a general enrichment in ¹³C (～3‰ and ～1‰ for the 2 years, respectively) consistent with increasing Gorczynski continentality index for warmer and drier conditions. In 2001 only, seasonal ¹³C enrichment between July and September, followed by depletion in November (from about ?26.0‰ to ?24.5‰ to ?30.0‰), was also observed. In 2001, July and August δ_er values across the network were significantly related to average daytime vapor pressure deficit (VPD), relative humidity (RH), and, to a lesser degree, air temperature (T_a), but not significantly with monthly average precipitation (P_m). In contrast, in 2002 (a much wetter peak season), δ_er was significantly related with T_a, but not significantly with VPD and RH. The important role of plant physiological processes on δ_er in 2001 was emphasized by a relatively rapid turnover (between 1 and 6 days) of assimilated carbon inferred from time‐lag analyses of δ_er vs. meteorological parameters. However, this was not evident in 2002. These analyses also noted corresponding diurnal cycles of δ_er and meteorological parameters in 2001, indicating a rapid transmission of daytime meteorology, via physiological responses, to the δ_er signal during this season. Organic matter δ¹³C results showed progressive ¹³C enrichment from leaves, through stems and roots to soil organic matter, which may be explained by ¹³C fractionation during respiration. This enrichment was species dependent and was prominent in angiosperms but not in gymnosperms. δ¹³C values of organic matter of any of the plant components did not well represent short‐term δ_er values during the seasonal cycle, and could not be used to partition ecosystem respiration into autotrophic and heterotrophic components.     

Carbon isotope discrimination and growth response of old Pinus ponderosa trees to stand density reductions

Stand density reductions have been proposed as a method by which old‐growth ponderosa pine (Pinus ponderosa) forests of North America can be converted back to pre‐1900 conditions, thereby reducing the danger of catastrophic forest fires and insect attacks while increasing the productivity of the remaining old‐growth individuals. However, the duration of productivity response of individual trees and the physiological mechanisms underlying such a response remain speculative issues, particularly in old trees. Tree‐ring measurements of carbon isotope ratios (δ¹³C) and basal area increment (BAI) were used to assess the response of intrinsic water‐use efficiency (the ratio of photosynthesis, A to stomatal conductance, g) and growth of individual> 250‐year‐old‐ponderosa pine trees to stand density reductions. It was hypothesized that reductions in stand density would increase soil moisture availability, thus decreasing canopy A/g and increasing carbon isotope discrimination (Δ). Cellulose‐δ¹³C of annual tree rings, soil water availability (estimated from pre‐dawn leaf water potential), photosynthetic capacity, stem basal growth and xylem anatomy were measured in individual trees within three pairs of thinned and un‐thinned stands. The thinned stands were treated 7 to 15 years prior to measurement. The values of δ¹³C and BAI were assessed for 20 consecutive years overlapping the date of thinning in a single intensively studied stand, and was measured for 3 years on either side of the date of thinning for the two other stands to assess the generality of the response. After thinning, Δ increased by 0.89‰ (± 0.15‰). The trees in the un‐thinned stands showed no change in Δ (0.00‰ ± 0.04‰). In the intensively studied trees, significant differences were expressed in the first growing season after the thinning took place but it took 6 years before the full 0.89‰ difference was observed. BAI doubled or tripled after disturbance, depending on the stand, and the increased BAI lasted up to 15 years after thinning. In the intensively studied trees, the BAI response did not begin until 3 years after the Δ response, peaked 1 year after the Δ peak, and then BAI and Δ oscillated in unison. The lag between BAI and Δ was not due to slow changes in anatomical properties of the sapwood, because tracheid dimensions and sapwood‐specific conductivity remained unchanged after disturbance. The Δ response of thinned trees indicated that A/g decreased after thinning. Photosynthetic capacity, as indexed by foliar nitrogen ([N]) and by the relationship between photosynthesis and internal CO₂ (A–C_i curves), was unchanged by thinning, confirming our suspicion that the decline in A/g was due to a relatively greater increase in g in comparison with A. Model estimates agreed with this conclusion, predicting that g increased by nearly 25% after thinning relative to a 15% increase in A. Pre‐dawn leaf water potential averaged 0.11 MPa (± 0.03 MPa) less negative for the thinned compared with the un‐thinned trees in all stands, and was strongly correlated with Δ post‐thinning (R² = 0.91). There was a strong relationship between BAI and modelled A, suggesting that changes in water availability and g have a significant effect on carbon assimilation and growth of these old trees. These results confirm that stand density reductions result in increased growth of individual trees via increased stomatal conductance. Furthermore, they show that a physiological response to stand density reductions can last for up to 15 years in old ponderosa pines if stand leaf area is not fully re‐established.     

Partitioning net ecosystem carbon exchange with isotopic fluxes of CO2

Because biological and physical processes alter the stable isotopic composition of atmospheric CO₂, variations in isotopic content can be used to investigate those processes. Isotopic flux measurements of ¹³CO₂ above terrestrial ecosystems can potentially be used to separate net ecosystem CO₂ exchange (NEE) into its component fluxes, net photosynthetic assimilation (F_A) and ecosystem respiration (F_R). In this paper theory is developed to partition measured NEE into F_A and F_R, using measurements of fluxes of CO₂ and ¹³CO₂, and isotopic composition of respired CO₂ and forest air. The theory is then applied to fluxes measured (or estimated, for ¹³CO₂) in a temperate deciduous forest in eastern Tennessee (Walker Branch Watershed). It appears that there is indeed enough additional information in ¹³CO₂ fluxes to partition NEE into its photosynthetic and respiratory components. Diurnal patterns in F_A and F_R were obtained, which are consistent in magnitude and shape with patterns obtained from NEE measurements and an exponential regression between night‐time NEE and temperature (a standard technique which provides alternate estimates of F_R and F_A). The light response curve for photosynthesis (F_A vs. PAR) was weakly nonlinear, indicating potential for saturation at high light intensities. Assimilation‐weighted discrimination against ¹³CO₂ for this forest during July 1999 was 16.8–17.1‰, depending on canopy conductance. The greatest uncertainties in this approach lie in the evaluation of canopy conductance and its effect on whole‐canopy photosynthetic discrimination, and thus the indirect methods used to estimate isotopic fluxes. Direct eddy covariance measurements of ¹³CO₂ flux are needed to assess the validity of the assumptions used and provide defensible isotope‐based estimates of the component fluxes of net ecosystem exchange.     

Estimating the contribution of leaf litter decomposition to soil CO2 efflux in a beech forest using 13C-depleted litter

The contribution of leaf litter decomposition to total soil CO₂ efflux (F_L/F) was evaluated in a beech (Fagus sylvatica L.) forest in eastern France. The Keeling‐plot approach was applied to estimate the isotopic composition of respired soil CO₂ from soil covered with either control (?30.32‰) or ¹³C‐depleted leaf litter (?49.96‰). The δ¹³C of respired soil CO₂ ranged from ?25.50‰ to ?22.60‰ and from ?24.95‰ to ?20.77‰, respectively, with depleted or control litter above the soil. The F_L/F ratio was calculated by a single isotope linear mixing model based on mass conservation equations. It showed seasonal variations, increasing from 2.8% in early spring to about 11.4% in mid summer, and decreasing to 4.2% just after leaf fall. Between December 2001 and December 2002, cumulated F and F_L reached 0.98 and 0.08 kg_C m^?2, respectively. On an annual basis, decomposition of fresh leaf litter accounted for 8% of soil respiration and 80% of total C loss from fresh leaf litter. The other fraction of carbon loss during leaf litter decomposition that is assumed to have entered the soil organic matter pool (i.e. 20%) represents only 0.02 kg_C m^?2.