Measurement of rhizosphere respiration and organic matter decomposition using natural 13C

Due to the limitations in methodology it has been a difficult task to measure rhizosphere respiration and original soil carbon decomposition under the influence of living roots. ¹⁴C-labeling has been widely used for this purpose in spite of numerous problems associated with the labeling method. In this paper, a natural ¹³C method was used to measure rhizosphere respiration and original soil carbon decomposition in a short-term growth chamber experiment. The main objective of the experiment was to validate a key assumption of this method: the ¹³C value of the roots represents the ¹³C value of the rhizosphere respired CO₂. Results from plants grown in inoculated carbon-free medium indicated that this assumption was valid. This natural ¹³C method was demonstrated to be advantageous for studying rhizosphere respiration and the effects of living roots on original soil carbon decomposition.     

Influence of rhizodeposition under elevated CO2 on plant nutrition and soil organic matter

Atmospheric CO₂ concentrations can influence ecosystem carbon storage through net primary production (NPP), soil carbon storage, or both. In assessing the potential for carbon storage in terrestrial ecosystems under elevated CO₂, both NPP and processing of soil organic matter (SOM), as well as the multiple links between them, must be examined. Within this context, both the quantity and quality of carbon flux from roots to soil are important, since roots produce specialized compounds that enhance nutrient acquisition (affecting NPP), and since the flux of organic compounds from roots to soil fuels soil microbial activity (affecting processing of SOM).From the perspective of root physiology, a technique is described which uses genetically engineered bacteria to detect the distribution and amount of flux of particular compounds from single roots to non-sterile soils. Other experiments from several labs are noted which explore effects of elevated CO₂ on root acid phosphatase, phosphomonoesterase, and citrate production, all associated with phosphorus nutrition. From a soil perspective, effects of elevated CO₂ on the processing of SOM developed under a C4 grassland but planted with C3 California grassland species were examined under low (unamended) and high (amended with 20 g m^–2 NPK) nutrients; measurements of soil atmosphere 13C combined with soil respiration rates show that during vegetative growth in February, elevated CO₂ decreased respiration of carbon from C4 SOM in high nutrient soils but not in unamended soils.This emphasis on the impacts of carbon loss from roots on both NPP and SOM processing will be essential to understanding terrestrial ecosystem carbon storage under changing atmospheric CO₂ concentrations.Abbreviations SOM soil organic matter - NPP net primary productivity - NEP net ecosystem productivity - PNPP p-nitrophenyl phosphate     

Diurnal cycle of carbon isotope ratio in soil CO2 in various ecosystems

Our investigations of diurnal variations of the ¹³C/¹²C ratio and CO₂ content in soil air were carried out in three environments during periods of high biosphere activity. It has been observed that diurnal variation of CO₂ concentration is negatively correlated ¹³. Particularly great variations occurred at shallow soil depths (10–30 cm) when the plant cover activity was high while the soil temperature was rather low. Under such conditions the ¹³ variations had the magnitude of 4, while the CO₂ concentration varied more than doubly. The maximum of the ¹³C/¹²C ratlo and the minimum of the CO₂ concentration in a cultivated field with winter wheat took place in the afternoon, whereas in deciduous forest similar patterns were observed at dawn. In these cases soil temperatures at 10 cm depths varied less than 2°C. Hence, under wheat the variation in root respiration rate seem to be the main reason of the recorded varations. In an uncultivated grass-field during the hottest period in summer we did not measure any distinct variations of CO₂ properties in spite of the fact that soil temperature varied up to 5°C. This might be due to dominant microbial respiration at the high soil temperature, which exceeded 20°C.     

Keeling plots for hummingbirds: a method to estimate carbon isotope ratios of respired CO<Subscript>2</Subscript> in small vertebrates

The carbon isotope composition of an animals breath reveals the composition of the nutrients that it catabolizes for energy. Here we describe the use of Keeling plots, a method widely applied in ecosystem ecology, to measure the ¹³C of respired CO₂ of small vertebrates. We measured the ¹³C of Rufous Hummingbirds (Selasphorus rufus) in the laboratory and of Mourning (Zenaida macroura) and White-winged (Z. asiatica) Doves in the field. In the laboratory, when hummingbirds were fed a sucrose based C3 diet, the ¹³C of respired CO₂ was not significantly different from that of their diet (¹³C_{C3 diet}). The ¹³C of respired CO₂ for C3 fasted birds was slightly, albeit significantly, depleted in ¹³C relative to ¹³C_{C3 diet}. Six hours after birds were shifted to a sucrose based C4 diet, the isotopic composition of their breath revealed that birds were catabolizing a mixture of nutrients derived from both the C3 and the C4 diet. In the field, the ¹³C of respired CO₂ from Mourning and White-winged Doves reflected that of their diets: the CAM saguaro cactus (Carnegeia gigantea) and C3 seeds, respectively. Keeling plots are an easy, effective and inexpensive method to measure ¹³C of respired CO₂ in the lab and the field.     

Carbon-isotope discrimination by leaves of Flaveria species exhibiting different amounts of C3-and C4-cycle co-function

Carbon-isotope ratios were examined as ¹³C values in several C₃, C₄, and C₃–C₄ Flaveria species, and compared to predicted ¹³C, values generated from theoretical models. The measured ¹³C values were within 4 of those predicted from the models. The models were used to identify factors that contribute to C₃-like ¹³C values in C₃–C₄ species that exhibit considerable C₄-cycle activity. Two of the factors contributing to C₃-like ¹³C values are high CO₂ leakiness from the C₄ pathway and pi/pa values that were higher than C₄ congeners. A marked break occurred in the relationship between the percentage of atmospheric CO₂ assimilated through the C₄ cycle and the ¹³C value. Below 50% C₄-cycle assimialtion there was no significant relationship between the variables, but above 50% the ¹³C values became less negative. These results demonstrate that the level of C₄-cycle expression can increase from, 0 to 50% with little integration of carbon transfer from the C₄ to the C₃ cycle. As expression increaces above 50%, however, increased integration of C₃- and C₄-cycle co-function occurs.Abbreviations and symbols RuBP carboxylase ribulose-1,5-bisphosphate carboxylase (EC 4.1.1.39) - PEP carboxylase phosphoenolpyruvate carboxylase (EC 4.1.1.31) - pa atmospheric CO₂ partial pressure - pi intercellular CO₂ partial pressure - isotope ratio - quantum yield for CO₂ uptake     

Spatial and temporal distribution of carbon isotopes in soil organic matter at the Dinghushan Biosphere Reserve, South China

The spatial and temporal distribution of carbon isotopes (¹³C, ¹⁴C) in soil organic matter (SOM) were studied based on SOM content, SOM ¹⁴C and SOM ¹³C of thinly layered soil samples for six soil profiles with different elevations at the Dinghushan Biosphere Reserve (DHSBR), South China. The results indicate that variations of SOM ¹³C with depth of the soil profiles at different elevations are controlled by soil development, and correlate well with SOM composition in terms of SOM compartments with different turnover rates, and SOM turnover processes at the DHSBR. The effect of carbon isotope fractionation was obvious during transformation of organic matter (OM) from plant debris to SOM in topsoil and SOM turnover processes after the topsoil was buried, which resulted in great increments of OM ¹³C, respectively. Increments of SOM ¹³C of topsoil from ¹³C of plant debris were controlled by SOM turnover rates. Both topsoil SOM ¹³C and plant debris ¹³C increase with elevation, indicating regular changes in vegetation species and composition with elevation, which is consistent with the vertical distribution of vegetation at the DHSBR. The six soil profiles at different elevations had similar characteristics in variations of SOM ¹³C with depth, alterations of SOM contents with depth and that SOM ¹⁴C apparent ages increasing with depth, respectively. These are presumably attributed to the regular distribution of different SOM compartments with depth because of their regular turnover during soil development. Depth with the maximal SOM ¹³C value is different in mechanism and magnitude with penetrating depth of ¹⁴C produced by nuclear explosion into atmosphere from 1952 to 1962, and both indicate controls of topography and vegetation on the distribution of SOM carbon isotopes with depth. Elevation exerts indirect controls on the spatial and temporal distribution of SOM carbon isotopes of the studied mountainous soil profiles at the DHSBR. This study shows that mountainous soil profiles at different elevations and with distinctive aboveground vegetation are presumably ideal sites for studies on soil carbon dynamics in different climatic-vegetation zones.     

Sucrose application,soil microbial respiration and evolved carbon dioxide isotope enrichment under contrasting land uses

Heterotrophic decomposition of organic matter dictates that substrate supply rate, including energy and nutrients, can limit soil microbial activity. In New Zealand, soils are naturally deficient in nitrogen and phosphorus. Fertiliser application is a part of pastoral agriculture, the countrys most widespread land use. We postulated that organic soils under grazed pasture and pristine forest would be at the extremes of substrate quality and supply rate, and thus potential microbial response to food opportunities. Soil microbial responses to the addition of fresh energy (sucrose) were determined by laboratory experiments with root-free samples and intact cores including roots. Responses were quantified by respiration and respired carbon (C) isotope (¹³C) enrichment measurements. A supra-trace sucrose dose (0.002 mol kg^–1 (soil)) caused the forest soils microbial respiration rate to nearly double within 2 h. The peak response took 20 h, and saturation occurred beyond a sucrose dose of 0.05 mol kg^–1 (soil). Intact soil cores from the forest had similar respiration rates and responses. For root-free soil samples from the grazed pasture, respiration response to sucrose was nearly immediate, dose dependent, and there was up to a 9-fold increase in the rate. Intact cores from the pasture had much higher respiration rates, but a similar response to sucrose. For both soils, the similarity of sucrose application effects on respiration and relative ¹³C enrichment of the respired carbon was striking.     

Forest- and pasture-derived carbon contributions to carbon stocks and microbial respiration of tropical pasture soils

The clearing of tropical forest for pasture leads to important changes in soil organic carbon (C) stocks and cycling patterns. We used the naturally occurring distribution of¹³C in soil organic matter (SOM) to examine the roles of forest- and pasture-derived organic matter in the carbon balance in the soils of 3- to 81-year-old pastures created following deforestation in the western Brazilian Amazon Basin state of Rondônia. Different ¹³C values of C3 forest-derived C (-28) and C4 pasture-derived C (-13) allowed determination of the origin of total soil C and soil respiration. The ¹³C of total soil increased steadily across ecosystems from -27.8 in the forest to -15.8 in the 81-year-old pasture and indicated a replacement of forest-derived C with pasture-derived C. The ¹³C of respired CO₂ increased more rapidly from -26.5 in the forest to -17 in the 3- to 13-year-old pastures and indicated a faster shift in the origin of more labile SOM. In 3-year-old pasture, soil C derived from pasture grasses made up 69% of respired C but only 17% of total soil C in the top 10 cm. Soils of pastures 5 years old and older had higher total C stocks to 30 cm than the original forest. This occurred because pasture-derived C in soil organic matter increased more rapidly than forest-derived C was lost. The increase of pasture-derived C in soils of young pastures suggests that C inputs derived from pasture grasses play a critical role in development of soil C stocks in addition to fueling microbial respiration. Management practices that promote high grass production will likely result in greater inputs of grass-derived C to pasture soils and will be important for maintaining tropical pasture soil C stocks.     

δ13C-variations of leaves in forests as an indication of reassimilated CO2 from the soil

Summary An attempt has been made to evaluate the contribution of soil respired CO₂ to the total assimilation of a forest tree, by heeding the ¹³C-concentrations of CO₂ from the free atmosphere and from mineralization processes within the soil respectively. An expression has been derived, according to which the assimilated fraction of CO₂ from the soil at a particular height of a tree is given by the ¹³C-value of the corresponding leaves, ¹³C of atmospheric CO₂, ¹³C of soil respired CO₂ and the physiological state of the leaves expressed as the ratio of total respiration over gross photosynthesis and internal over external CO₂-concentration. In the particular case investigated, a ¹³C-difference of 5 has been determined from bottom to top of a beech tree which results in a CO₂ contribution from the soil of about 22% for the lower forest strata, while the total contribution of soil respired CO₂ accounts for about 5% of the overall assimilation.     

The carbon isotope ratio of plant organic material reflects temporal and spatial variations in CO2 within tropical forest formations in Trinidad

Summary A method of monitoring and collecting CO₂ samples in the field has been developed which has been used to study both temporal and spatial variations in canopy CO₂ isotopic signatures in two contrasting tropical forest formations in Trinidad. These have been related to vertical gradients in the carbon isotope ratio (¹³C) of organic material in conjunction with measurements of other environmental parameters. The ¹³C of leaf material from two canopies showed a gradient with respect to height, more negative values being found low in the understorey. The deciduous secondary forest, (Simla) showed a difference of 4.6 and the semi-evergreen seasonal canopy (Aripo), 2.8. The range of ¹³C values at Simla was 4 less negative than those at Aripo. In order to relate these measurements to the interaction between diffusion or carboxylation limitation, and source CO₂ effects, variations in environmental parameters through the canopy have been compared with changes in CO₂ partial pressure (P _a) and isotopic composition ¹³C throughout the day during the dry season. Values of P _a20 m above the ground at Aripo varied from 380 vpm at dawn to 340 vpm at midday, at which time the partial pressure 15 cm above the ground was 375 vpm. The CO₂ partial pressure did not stabilise during the course of the day, and there was good correlation (r ²=0.82) between _a and P _a, with more negative values of _a occuring in the understorey. Diuraal changes of 2 were evident at all canopy positions. In the more open canopy at Simla, these gradients were similar, but less marked. Leaf-air vapour pressure deficit (VPD) showed no relationship with height, possibly as a result of minimal water flux from both the soil and the canopy due to low soil water content; VPD was 1.5 kPa higher at midday than dawn. A 3° C temperature gradient between the understorey and upper canopy was observed at Aripo but not in the more open Simla canopy. CO₂ partial pressure stabilised for only 4 h in the middle of the day, while other parameters showed no stable period. The proportion of floor respired CO₂ reassimilated at Aripo has been calculated as 26%, 19%, and 8% for the periods 0600–1000, 1000–1400, and 1400–1800 hours. In order to quantify source CO₂ effects, measurements of the environmental parameters and assimilation rate must be made at all canopy positions and throughout the day.     

Comparisons of δ<Superscript>13</Superscript>C of photosynthetic products and ecosystem respiratory CO<Subscript>2</Subscript> and their responses to seasonal climate variability

This study investigated the relationship between ¹³C of ecosystem components, soluble plant carbohydrates and the isotopic signature of ecosystem respired CO₂ (¹³C_R) during seasonal changes in soil and atmospheric moisture in a beech (Fagus sylvatica L.) forest in the central Apennine mountains, Italy. Decrease in soil moisture and increase in air vapour pressure deficit during summer correlated with substantial increase in ¹³C of leaf and phloem sap soluble sugars. Increases in ¹³C of ecosystem respired CO₂ were linearly related to increases in phloem sugar ¹³C (r²=0.99, P0.001) and leaf sugar ¹³C (r²=0.981, P0.01), indicating that a major proportion of ecosystem respired CO₂ was derived from recent assimilates. The slopes of the best-fit lines differed significantly (P0.05), however, and were about 0.86 (SE=0.04) for phloem sugars and about 1.63 (SE=0.16) for leaf sugars. Hence, changes in isotopic signature in phloem sugars were transferred to ecosystem respiration in the beech forest, while leaf sugars, with relatively small seasonal changes in ¹³C, must have a slower turnover rate or a significant storage component. No significant variation in ¹³C was observed in bulk dry matter of various plant and ecosystem components (including leaves, bark, wood, litter and soil organics). The apparent coupling between the ¹³C of soluble sugars and ecosystem respiration was associated with large apparent isotopic disequilibria. Values of ¹³C_R were consistently more depleted by about 4 relative to phloem sugars, and by about 2 compared to leaf sugars. Since no combination of the measured pools could produce the observed ¹³C_R signal over the entire season, a significant isotopic discrimination against ¹³C might be associated with short-term ecosystem respiration. However, these differences might also be explained by substantial contributions of other not measured carbon pools (e.g., lipids) to ecosystem respiration or contributions linked to differences in footprint area between Keeling plots and carbohydrate sampling. Linking the seasonal and inter-annual variations in carbon isotope composition of carbohydrates and respiratory CO₂ should be applicable in carbon cycle models and help the understanding of inter-annual variation in biospheric sink strength.     

Uptake of organic matter by the roots of sweet potato: Analysis of the δ14C value of plants

Summary To determine whether sweet potato (Ipomoea batatas (L.) Poir.) takes up organic matter through the roots from the medium, the concentrations of natural¹⁴C (¹⁴C) in plant organic matter, atmospheric CO₂ and compost applied to media were examined under soil and sand culture conditions. In these experiments, three kinds of composts of different ¹⁴C were used. CO₂ derived from the mineralization of compost was continuously pumped out from the pots and its direct uptake by the leaves was prevented.¹⁴C of plant parts harvested after the 43 days experimental period were affected by the ¹⁴C of the compost in the treatments where the compost of rice straw was applied, and which suggested that a significant amount of plant carbon was derived from the compost.     

Influence of vegetation and soil CO2 exchange on the concentration and stable oxygen isotope ratio of atmospheric CO2 within a Pinus resinosa canopy

Measurements were made of the concentration and stable oxygen isotopic ratio of carbon dioxide in air samples collected on a diurnal basis at two heights within a Pinus resinosa canopy. Large changes in CO₂ concentration and isotopic composition were observed during diurnal time courses on all three symple dates. In addition, there was strong vertical stratification in the forest canopy, with higher CO₂ concentrations and more negative ¹⁸O values observed closer to the soil surface. The observed daily increases in ¹⁸O values of forest CO₂ were dependent on relative humidity consistent with the modelled predictions of isotopic fractionation during photosynthetic gas exchange. During photosynthetic gas exchange, a portion of the CO₂ that enters the leaf and equilibrates with leaf water is not fixed and diffuses back out of the leaf with an altered oxygen isotopic ratio. The oxygen isotope ratio of CO₂ diffusing out of a leaf depends primarily on the ¹⁸O content of leaf water which changes in response to relative humidity. In contrast, soil respiration caused a decline in the ¹⁸O values of forest CO₂ at night, because CO₂ released from the soil has equilibrated with soil water which has a lower ¹⁸O content than leaf water. The observed relationship between diurnal changes in CO₂ concentration and oxygen isotopic composition in the forest environment were consistent with a gas mixing model that considered the relative magnitudes of CO₂ fluxes associated with photosynthesis, respiration and turbulent exchange between the forest and the bulk atmosphere.     

Hydrogen isotope discrimination in higher plants: Correlations with photosynthetic pathway and environment

Summary The ratio of deuterium to hydrogen (expressed as D) in hydrogen released as water during the combustion of dried plant material was examined. The D value (metabolic hydrogen) determined on plant materials grown under controlled conditions is correlated with pathways of photosynthetic carbon metabolism. C₃ plants show mean D values of-132 for shoots and -117 for roots; C₄ plants show mean D values of -91 for shoots and-77 for roots and CAM plants a D value of-75 for roots and shoots. The difference between the D value of shoot material from C₃ and C₄ plants was confirmed in species growing under a range of glasshouse conditions. This difference in D value between C₃ and C₄ species does not appear to be due to differences in the D value (tissue water) in the plants as a result of physical fractionation of hydrogen isotopes during transpiration. In C₃ and C₄ plants the hydrogen isotope discrimination is in the same direction as the carbon isotope discrimination and factors contributing to the difference in D values are discussed. In CAM plants grown in the laboratory or collected from the field D values range from-75 to +50 and are correlated with ¹³C values. When deprived of water, the D value (metabolic hydrogen) in both soluble and insoluble material in leaves of Kalanchoe daigremontiana Hamet et Perr., becomes less negative. These changes may reflect the deuterium enrichment of tissue water during transpiration, or in field conditions, may reflect the different D value of available water in areas of increasing aridity. Whatever the origin of the variable D value in CAM plants, this parameter may be a useful index of the water relations of these plants under natural conditions.     

Origin and fate of organic carbon in the freshwater part of the Scheldt Estuary as traced by stable carbon isotope composition

We investigated the seasonal and geographical variation in the stable carbon isotope ratios of total dissolved inorganic carbon (¹³C_POC) and suspended matter (¹³C_POC) in the freshwater part of the River Scheldt. Two major sources of particulate organic matter (POM) occur in this riverine system: riverine phytoplankton and terrestrial detritus. In winter the lowest ¹³C_DIC values are observed due to enhanced input of CO₂ from decomposition of ¹³C-depleted terrestrial plant detritus (average ¹³C_DIC = –/14.3). During summer, when litter input from terrestrial flora is the lowest, water column respiration on POM of terrestrial origin is also the lowest as evidenced by less negative ¹³C_DIC values (average ¹³C_DIC = –9.9). In winter the phytoplankton biomass is low, as indicated by low chlorophyll a concentrations (Chl a < 4.5 gl^–1), compared to summer when chlorophyll a concentrations can rise to a maximum of 54 gl^–1. Furthermore, in winter the very narrow range of ¹³C_POC (from –26.5 to –27.6) is associated with relatively high C/N ratios (C/N > 9) suggesting that in winter a major fraction of POC is derived from allochthonous matter. In summer ¹³C_POC exhibits a very wide range of values, with the most negative values coinciding with high Chl a concentrations and low C/N ratios (C/N < 8). This suggests predominance of phytoplankton carbon in the total particulate carbon pool, utilising a dissolved inorganic carbon reservoir, which is already significantly depleted in ¹³C. Using a simple two source mixing approach a reconstruction of the relative importance of phytoplankton to the total POC pool and of ¹³C/¹²C fractionation by phytoplankton is attempted.     

Effects of competition and N and P supply on carbon isotope discrimination and <Superscript>15</Superscript>N-natural abundance in four grassland species

The effect of interspecific competition and element additions (N and P) on four grassland species (Poa pratensis, Lolium perenne, Festuca valida, Taraxacum officinale) grown under field conditions was studied. Two grasses (L. perenne, F. valida) grown in monoculture (absence of competition) showed lower carbon isotope discrimination (¹³C) and enriched ¹⁵N values. Nitrogen addition (as urea) had inconsistent effects on species ¹³C while caused enrichment of ¹⁵N of P. pratensis and F. valida but strong depletion of ¹⁵N of T. officinale. Phosphorous had no significant effect on ¹³C but depleted ¹⁵N of all species.     

Carbon isotopes and carbon turnover in cotton and wheat FACE experiments

The Maricopa cotton and wheat FACE (free-air CO₂ enrichment) experiments offer propitious opportunity to quantify carbon turnover. The commercial CO₂ (% MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGqiVu0Je9sqqrpepC0xbbL8F4rqqrFfpeea0xe9Lq-Jc9% vqaqpepm0xbba9pwe9Q8fs0-yqaqpepae9pg0FirpepeKkFr0xfr-x% fr-xb9adbaqaaeGaciGaaiaabeqaamaabaabaaGcbaGaeqiTdq2aaW% baaSqabeaacaaIXaGaaG4maaaakiaaboeacqGHijYUcqGHsislcaaI% ZaGaaG4naiaacwcaliaad+gaaaa!3FCB!\[\delta ^{13} {\text{C}} \approx - 37\% o\]) used to elevate CO₂ concentration in field plots provided a strongly ¹³C-depleted tracer. Soil CO₂ and ¹³C of soil organic carbon (SOC) in CO₂-enriched and Control plots were measured between the final cotton FACE project (October 1991) and the end of the second wheat experiment (June 1994). The initial ¹³C-depletion in SOC of cotton FACE plots (measured by the difference in ¹³C between FACE and Control plots) persisted at the same level (1.9) 1.5 years after the experiment ended. A similar depletion was observed in soil CO₂ evolved in the same plots, indicating ongoing decomposition of the new SOC. The SOC ¹³C of wheat plots before and after two growing seasons showed increasing ¹³C-depletion in FACE relative to Control. Isotopic mass balance was consistent with 5–6% new carbon input from the two wheat crops. This is lower than the 12–13% calculated for FACE cotton and perhaps a consequence of the larger root system of cotton or the 3-year duration of the cotton experiments versus 2 years for the wheat.     

Is growth of soil microorganisms in boreal forests limited by carbon or nitrogen availability?

To study whether the biomass of soil microorganisms in a boreal Pinus sylvestris-Vaccinium vitis-idaea forest was limited by the availability of carbon or nitrogen, we applied sucrose from sugar cane, a C₄ plant, to the organic mor-layer of the C₃–C dominated soil. We can distinguish between microbial mineralization of the added sucrose and respiration of endogenous carbon (root and microbial) by using the C₄-sucrose as a tracer, exploiting the difference in natural abundance of ¹³C between the added C₄-sucrose (¹³C –10.8) and the endogenous C₃–carbon (¹³C –26.6 ). In addition to sucrose, NH₄Cl (340 kg N ha^–1) was added factorially to the mor-layer. We followed the microbial activity for nine days after the treatments, by in situ sampling of CO₂ evolved from the soil and mass spectrometric analyses of ¹³C in the CO₂. We found that microbial biomass was limited by the availability of carbon, rather than nitrogen availability, since there was a 50% increase in soil respiration in situ between 1 h and 5 days after adding the sucrose. However, no further increase was observed unless nitrogen was also added. Analyses of the ¹³C ratios of the evolved CO₂ showed that increases in respiration observed between 1 h and 9 days after the additions could be accounted for by an increase in mineralization of the added C₄–C.     

Carbon fluxes to the soil in a mature temperate forest assessed by <Superscript>13</Superscript>C isotope tracing

Photosynthetic carbon uptake and respiratory C release from soil are major components of the global carbon balance. The use of ¹³C depleted CO₂ (¹³C = –30) in a free air CO₂ enrichment experiment in a mature deciduous forest permitted us to trace the carbon transfer from tree crowns to the rhizosphere of 100–120 years old trees. During the first season of CO₂ enrichment the CO₂ released from soil originated substantially from concurrent assimilation. The small contribution of recent carbon in fine roots suggests a much slower fine root turnover than is often assumed.¹³C abundance in soil air correlated best with temperature data taken from 4 to 10 days before air sampling time and is thus rapidly available for root and rhizosphere respiration. The spatial variability of ¹³C in soil air showed relationships to above ground tree types such as conifers versus broad-leaved trees. Considering the complexity and strong overlap of roots from different individuals in a forest, this finding opens an exciting new possibility of associating respiration with different species. What might be seen as signal noise does in fact contain valuable information on the spatial heterogeneity of tree-soil interaction.     

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.