Increased root exudation of14C-compounds by sorghum seedlings inoculated with nitrogen-fixing bacteria

Summary Organic components leaked fromSorghum bicolor seedlings (‘root exudates’) were examined by recovering¹⁴C labelled compounds from root solutions of seedlings inoculated withAzospirillum brasilense, Azotobacter vinelandii orKlebsiella pneumoniae nif-. Up to 3.5% of the total¹⁴C recovered from shoots, roots, and nutrient solutions was found in the root solutions. Inoculation with Azospirillum and Azotobacter increased the amounts of¹⁴C and decreased the amounts of carbohydrates in the root solutions. When sucrose was added as a carbon source for the bacteria, the increase of¹⁴C in the solutions did not occur. Quantities of¹⁴C found in the root solutions were proportional to amounts of mineral nitrogen supplied to the plants. Bacterial growth also was proportional to nitrogen levels. When sorghum plants were grown in soil and labelled with¹⁴CO₂, about 15% of the total¹⁴C recovered within 48 hours exposure was found in soil leachates.     

Influence of long-term CO2 storage on δ13C in vials capped with butyl and butyl/PTFE caps and the relevance for ecological samples

Background

The carbon (C) isotope signature of solid materials such as plants and soil, or gaseous samples (atmospheric or soil air), can be used as a useful tool for investigations of the C cycle. In gaseous samples, stability of δ¹³C of carbon dioxide (CO₂) in air during storage represents a problem.

Methods

We tested the long-term storage effect of δ¹³C originated from CO₂ in vials both capped with butyl or butyl coated with polytetrafluoroethylene (PTFE) on the internal surface. Therefore, pure CO₂, depleted in ¹³C, was stored for up to 736 days. In addition, the relevance of long-term storage for ecological soil air data collected from a free-air CO₂ enrichment (FACE) experiment located in Stuttgart (Germany) during one growing period with a maximum storage period of 210 days was judged.

Results

With increasing storage time, a change in isotopic composition towards less depleted δ¹³C was observed. The changes in δ¹³C were highest at the beginning of the storage period and decreased over time, which could be described with an asymptotic model. The maximum change in δ¹³C was less than 2?‰ and lower for vials capped with butyl/PTFE septa. In the FACE experiment, the comparison between corrected and uncorrected data showed that δ¹³C originated from soil air changed within this data set by up to 1?‰. The calculation of the fraction of plant derived C resulted in an underestimation of up to 10 %.

Conclusion

The storage effect should be taken into consideration when interpreting δ¹³C values in order to avoid miscalculations.     

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.     

Photosynthesis controls of CO2 efflux from maize rhizosphere

The effects of different shading periods of maize plants on rhizosphere respiration and soil organic matter decomposition were investigated by using a ¹³C natural abundance and ¹⁴C pulse labeling simultaneously. ¹³C was a tracer for total C assimilated by maize during the whole growth period, and ¹⁴C was a tracer for recently assimilated C. CO₂ efflux from bare soil was 4 times less than the total CO₂ efflux from planted soil under normal lighting. Comparing to the normal lighting control (12/12 h day/night), eight days with reduced photosynthesis (12/36 h day/night period) and strongly reduced photosynthesis (12/84 h day/night period) resulted in 39% and 68% decrease of the total CO₂ efflux from soil, respectively. The analysis of ¹³C natural abundance showed that root-derived CO₂ efflux accounted for 82%, 68% and 56% of total CO₂ efflux from the planted soil with normal, prolonged and strongly prolonged night periods, respectively. Clear diurnal dynamics of the total CO₂ efflux from soil with normal day-night period as well as its strong reduction by prolonged night period indicated tight coupling with plant photosynthetic activity. The light-on events after prolonged dark periods led to increases of root-derived and therefore of total CO₂ efflux from soil. Any factor affecting photosynthesis, or substrate supply to roots and rhizosphere microorganisms, is an important determinant of root-derived CO₂ efflux, and thereby, total CO₂ efflux from soils. ¹⁴C labeling of plants before the first light treatment did not show any significant differences in the ¹⁴CO₂ respired in the rhizosphere between different dark periods because the assimilate level in the plants was high. Second labeling, conducted after prolonged night phases, showed higher contribution of recently assimilated C (¹⁴C) to the root-derived CO₂ efflux by shaded plants. Results from ¹³C natural abundance showed that the cultivation of maize on Chromic Luvisol decreased soil organic matter (SOM) mineralization compared to unplanted soil (negative priming effect). A more important finding is the observed tight coupling of the negative rhizosphere effect on SOM decomposition with photosynthesis.     

Herbivore-induced changes in plant carbon allocation: assessment of below-ground C fluxes using carbon-14

Effects of above-ground herbivory on short-term plant carbon allocation were studied using maize (Zea mays) and a generalist lubber grasshopper (Romalea guttata). We hypothesized that above-ground herbivory stimulates current net carbon assimilate allocation to below-ground components, such as roots, root exudation and root and soil respiration. Maize plants 24 days old were grazed (c. 25–50% leaf area removed) by caging grasshoppers around individual plants and 18 h later pulse-labelled with¹⁴CO₂. During the next 8 h,¹⁴C assimilates were traced to shoots, roots, root plus soil respiration, root exudates, rhizosphere soil, and bulk soil using carbon-14 techniques. Significant positive relationships were observed between herbivory and carbon allocated to roots, root exudates, and root and soil respiration, and a significant negative relationship between herbivory and carbon allocated to shoots. No relationship was observed between herbivory and¹⁴C recovered from soil. While herbivory increased root and soil respiration, the peak time for¹⁴CO₂ evolved as respiration was not altered, thereby suggesting that herbivory only increases the magnitude of respiration, not patterns of translocation through time. Although there was a trend for lower photosynthetic rates of grazed plants than photosynthetic rates of ungrazed plants, no significant differences were observed among grazed and ungrazed plants. We conclude that above-ground herbivory can increase plant carbon fluxes below ground (roots, root exudates, and rhizosphere respiration), thus increasing resources (e.g., root exudates) available to soil organisms, especially microbial populations.     

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.     

Spatial distribution and turnover of root-derived carbon in alfalfa rhizosphere depending on top- and subsoil properties and mycorrhization

Aims

This study analyzed the extent to which root exudates diffuse from the root surface towards the soil depending on topsoil and subsoil properties and the effect of arbuscular mycorrhizal fungal hyphae on root-derived C distribution in the rhizosphere.

Methods

Alfalfa was grown in three-compartment pots. Nylon gauze prevented either roots alone or roots and arbuscular mycorrhizal fungal hyphae from penetrating into the rhizosphere compartments. ¹⁴CO₂ pulse labeling enabled the measurement of ¹⁴C-labeled exudates in dissolved (DOC) and total organic carbon (TOC) in the rhizosphere, distributed either by diffusion alone or by diffusion, root hair and hyphal transport.

Results

Root exudation and microbial decomposition of exudates was higher in the rhizosphere with topsoil compared to subsoil properties. Exudates extended over 28 mm (DOC) and 20 mm (TOC). Different soil properties and mycorrhization, likely caused by the low arbuscular mycorrhizal colonization of roots (13?±?4 % (topsoil properties) and 18?±?5 % (subsoil properties)), had no effect.

Conclusions

Higher microbial decomposition compensated for higher root exudation into the rhizosphere with topsoil properties, which resulted in equal exudate extent when compared to the rhizosphere with subsoil properties. Higher ¹⁴C activity used for labeling compared with previous studies enabled the detection of low exudate concentrations at longer distances from the root surface.     

Bacterial inoculation of maize affects carbon allocation to roots and carbon turnover in the rhizosphere

To assess the influence of bacteria inoculation on carbon flow through maize plant and rhizosphere,¹⁴C allocation after¹⁴CO₂ application to shoots over a 5-day period was determined. Plants were grown on C- and N-free quartz sand in two-compartment pots, separating root and shoot space. While one treatment remained uninoculated, treatments two and three were inoculated withPantoea agglomerans (D5/23) andPseudomonas fluorescens (Ps I A12), respectively, five days after planting. Bacterial inoculation had profound impacts on carbon distribution within the system. Root/rhizosphere respiration was increased and more carbon was allocated to roots of plants being inoculated. After five days of¹⁴CO₂ application, more ethanol-soluble substances were found in roots of inoculated treatments and lower rhizodeposition indicated intensive C turnover in the rhizosphere. In both inoculated treatments the intensity of photosynthesis measured as net-CO₂-assimilation rates were increased when compared to the uninoculated plants. However, high C turnover in the rhizosphere reduced shoot growth of D5/23 inoculated plants, with no effect on shoot growth of Ps I A12 inoculated plants. A separation of labeled compounds in roots and rhizodeposition revealed that neutral substances (sugars) constituted the largest fraction. The relative fractions of sugars, amino acids and organic acids in roots and rhizodeposition suggest that amino acid exudation was particularly stimulated by bacterial inoculation and that turnover of this substance group is high in the rhizosphere.     

Dynamics of photoassimilated carbon in douglas fir seedlings

The relations between CO₂ uptake, translocation, and carbon accumulation in several vegetative components of Douglas fir seedlings (Pseudotsuga menziesii [Mirb.] Franco) have been quantified using ¹⁴CO₂. Seedlings were exposed to a constant specific radioactivity of ¹⁴CO₂ and a repeating daily pattern of temperature and light for 4 consecutive days. Results of ¹⁴C analysis, which indicated a transitory pattern of photoassimilated carbon movement, were extrapolated to a “steady rate” using a compartment analysis. Accumulation rates of photoassimilated carbon, relative to tissue carbon, were new needles, 0.94%/day, old needles, 1.14%/day, new shoots 0.38%/day, stem, 0.16%/day, and roots, 0.50%/day. Therefore, the source of carbon, the needles, is also the strongest sink.     

Rice biomass production and carbon cycling in 13CO2 pulse-labeled microcosms with different soils under submerged conditions

Aims

Rice fields are an important source for the greenhouse gas methane. Plants play an essential role in carbon supply for soil microbiota, but the influence of the microbial community on carbon cycling is not well understood.

Methods

Microcosms were prepared using sand-vermiculite amended with different soils and sediments, and planted with rice. The microcosms at different growth stages were pulse-labeled with ¹³CO₂ followed by tracing ¹³C in plant, soil and atmospheric carbon pools and quantifying the abundance of methanogenic archaea in rhizosphere soil.

Results

Overall,?>85 % of the freshly assimilated carbon was allocated in aboveground plant biomass, approximately 10 % was translocated into the roots and?4, but emission of ¹³C-labeled CH₄ started immediately and ¹³C enrichment revealed that plant-derived carbon was an important source for methanogenesis. The results further demonstrated that carbon assimilation and translocation processes, microbial abundance and gas emission were not only affected by the plant growth stage, but also by the content and type of soil in which the rice plants grew.

Conclusions

The study illustrates the close ties between plant physiology, soil properties and microbial communities for carbon turnover and ecosystem functioning.     

Allocation of carbon to shoots,roots, soil and rhizosphere respiration by barrel medic (Medicago truncatula) before and after defoliation

The allocation of carbon to shoots, roots, soil and rhizosphere respiration in barrel medic (Medicago truncatulaGaertn.) before and after defoliation was determined by growing plants in pots in a labelled atmosphere in a growth cabinet. Plants were grown in a ¹⁴CO₂-labelled atmosphere for 30 days, defoliated and then grown in a ¹³CO₂-labelled atmosphere for 19 days. Allocation of ¹⁴C-labelled C to shoots, roots, soil and rhizosphere respiration was determined before defoliation and the allocation of ¹⁴C and ¹³C was determined for the period after defoliation. Before defoliation, 38.4% of assimilated C was allocated below ground, whereas after defoliation it was 19.9%. Over the entire length of the experiment, the proportion of net assimilated carbon allocated below ground was 30.3%. Of this, 46% was found in the roots, 22% in the soil and 32% was recovered as rhizosphere respiration. There was no net translocation of assimilate from roots to new shoot tissue after defoliation, indicating that all new shoot growth arose from above-ground stores and newly assimilated carbon. The rate of rhizosphere respiration decreased immediately after defoliation, but after 8 days, was at comparable levels to those before defoliation. It was not until 14 days after defoliation that the amount of respiration from newly assimilated C (¹³C) exceeded that of C assimilated before defoliation (¹⁴C). This revised version was published online in June 2006 with corrections to the Cover Date.     

Carbon loss from the roots of tomato and pea seedlings grown in soil

Tomato and pea seedlings were grown for 14d and 28d with shoots in constant specific activity¹⁴CO₂ and the amounts and distribution of carbon within the plants and of that released into the soil from the roots were measured. The estimates of carbon loss were derived from measurements of¹⁴CO₂ respired from both the root and the accompanying microbial population and from the root derived¹⁴C-labelled organic carbon compounds in the soil. The relationship between plant growth and the loss of carbon and distribution of carbon within the plants are discussed.     

Losses of14C from roots of pulse-labelled wheat and barley during washing from soil

In crop carbon budget studies losses of root material during storage and washing of samples may cause considerable errors. To correct data from field experiments where rhizosphere C fluxes in wheat and barley were determined by¹⁴C pulse-labelling at different development stages, experiments were performed to quantify losses of¹⁴C from roots during washing. Losses of¹⁴C from wheat roots grown on nutrient solution and stored in different ways, decreased from on average 45% of total¹⁴C content 8 days after labelling to 27% after 21 days. This decrease was probably related to the incorporation of¹⁴C into structural compounds. During washing of oven-dried soil cores of held-grown wheat and barley 3 weeks after labelling, different size classes of losses of¹⁴C from the roots increased substantially with the development stage of the crop at labelling. The 0.3–0.6 mm size class increased from 5% of the¹⁴C in roots > 0.3 mm in young plants to 25% at ripening, and the < 0.3 mm size class increased from 8 to 41% of total¹⁴C content. The latter size class was, however, determined by washing handpicked roots and may therefore partly consist of adhering exudates, mucilages and microorganisms. The effect of development stage on root washing losses was attributed to root senescence which increases the fragility of roots. Thus, especially at the rate development stages root washing losses caused a severe underestimation of the root¹⁴C content. However, with these results the¹⁴C distribution patterns of the field experiments could be adequately corrected.Communication No. 77 of the Dutch Programme on Soil Ecology of Arable Farming Systems.     

Effect of blast infection on the photosynthetic activity of rice seedlings

Photosynthesis was affected considerably when the cultivars Zenith, Lacrosse, Usen and Co. 13 belonging to indica and japonica groups of rice were inoculated with physiological races IC₁₃ and ID₁ ofPyricularia oryzae Cav. The 21-day seedlings after inoculation withP. oryzae and uninoculated control were exposed to¹⁴CO₂ under natural sunlight (190 Wm^-2). The extent of reduction in photosynthetic rate in the cultivars varied on inoculation with different races ofP. oryzae used, the reduction being more apparent with race ID₁.     

Estimation of rhizodeposition at field scale: upscaling of a 14C labeling study

Background and aims

Rhizodeposition of plants is the most uncertain component of the carbon (C) cycle. By existing approaches the amount of rhizodeposition can only roughly be estimated since its persistence in soil is very short compared to other organic C pools. We suggest an approach to quantify rhizodeposition at the field scale by assuming a constant ratio between rhizodeposited-C to root-C.

Methods

Maize plants were pulse-labeled with ¹⁴CO₂ under controlled conditions and the soil ¹⁴CO₂ efflux was separated into root and rhizomicrobial respiration. The latter and the ¹⁴C activity remaining in the soil corresponded to total rhizodeposition. By relating rhizodeposited-¹⁴C to root-¹⁴C a rhizodeposition-to-root ratio of 0.56 was calculated. This ratio was applied to the root biomass C measured in the field to estimate rhizodeposition under field conditions.

Results

Maize allocated 298 kg C ha^?1 as root-C and 166 kg C ha^?1 as rhizodeposited-C belowground, 50 % of which were recovered in the upper 10 cm. The fate of rhizodeposits was estimated based on the ¹⁴C data, which showed that 62 % of total rhizodeposition was mineralized within 16 days, 7 % and 0.3 % was incorporated into microbial biomass and DOC, respectively, and 31 % was recovered in the soil.

Conclusions

We conclude that the present approach allows for an improved estimation of total rhizodeposition, since it accounts not only for the fraction of rhizodeposits remaining in soil, but also for that decomposed by microorganisms and released from the soil as CO₂.     

Carbon fluxes in the rhizosphere of sweet chestnut seedlings (Castanea sativa) grown under two atmospheric CO2 concentrations: 14C partitioning after pulse labelling

Partitioning of ¹⁴C was assessed in sweet chestnut seedlings (Castanea sativa Mill.) grown in ambient and elevated atmospheric [CO₂] environments during two vegetative cycles. The seedlings were exposed to ¹⁴CO₂ atmosphere in both high and low [CO₂] environments for a 6-day pulse period under controlled laboratory conditions. Six days after exposure to ¹⁴CO₂, the plants were harvested, their dry mass and the radioactivity were evaluated. ¹⁴C concentration in plant tissues, root-soil system respiratory outputs and soil residues (rhizodeposition) were measured. Root production and rhizodeposition were increased in plants growing in elevated atmospheric [CO₂]. When measuring total respiration, i.e. CO₂ released from the root/soil system, it is difficult to separate CO₂ originating from roots and that coming from the rhizospheric microflora. For this reason a model accounting for kinetics of exudate mineralization was used to estimate respiration of rhizospheric microflora and roots separately. Root activity (respiration and exudation) was increased at the higher atmospheric CO₂ concentration. The proportion attributed to root respiration accounted for 70 to 90% of the total respiration. Microbial respiration was related to the amount of organic carbon available in the rhizosphere and showed a seasonal variation dependent upon the balance of root exudation and respiration. The increased carbon assimilated by plants grown under elevated atmospheric [CO₂] stayed equally distributed between these increased root activities. ei]H Lambers     

Regulation of biosynthesis of secondary metabolites

AcetylCoA carboxylase (E.C.6.4.1.2) enzymatic activity was studied in high- and low-producing strains ofStreptomyces aureofaciens. Study of the rate of H¹⁴CO₃ ^? incorporation into acetylCoA and of CH₃ ¹⁴COO^? incorporation into the fatty acids showed that specific activity attained maximum values after about 24 hours' cultivation. AcetylCoA carboxylase activity in the producing strain was approximately double the value in the low-producing strain. The subcellular particles present in the supernatant fractions from cells disintegrated by spinning one hour at 600g and 10,000g did not influence the activity of the enzyme. Avidin inhibited it. The relationship between specific activity of the enzyme and the rate of oligoketide synthesis is discussed.     

Modifications of the carbon and nitrogen allocations in the plant (Triticum aestivum L.) soil system in response to increased atmospheric CO2 concentration

The aim of this work was to examine the response of wheat plants to a doubling of the atmospheric CO₂ concentration on: (1) carbon and nitrogen partitioning in the plant; (2) carbon release by the roots; and (3) the subsequent N uptake by the plants. The experiment was performed in controlled laboratory conditions by exposing fast-growing spring wheat plants, during 28 days, to a ¹⁴CO₂ concentration of 350 or 700 L L^–1 at two levels of soil nitrogen fertilization. Doubling CO₂ availability increased total plant production by 34% for both N treatment. In the N-fertilized soil, the CO₂ enrichment resulted in an increase in dry mass production of 41% in the shoots and 23% in the roots; without N fertilization this figure was 33% and 37%, respectively. In the N-fertilized soil, the CO₂ increase enhanced the total N uptake by 14% and lowered the N concentration in the shoots by 23%. The N concentration in the roots was unchanged. In the N-fertilized soil, doubling CO₂ availability increased N uptake by 32% but did not change the N concentrations, in either shoots or roots. The CO₂ enrichment increased total root-derived carbon by 12% with N fertilization, and by 24% without N fertilization. Between 85 and 90% of the total root derived-¹⁴C came from respiration, leaving only 10 to 15% in the soil as organic ¹⁴C. However, when total root-derived ¹⁴C was expressed as a function of root dry weight, these differences were only slightly significant. Thus, it appears that the enhanced carbon release from the living roots in response to increased atmospheric CO₂, is not due to a modification of the activity of the roots, but is a result of the increased size of the root system. The increase of root dry mass also resulted in a stimulation of the soil N mineralization related to the doubling atmospheric CO₂ concentration. The discussion is focused on the interactions between the carbon and nitrogen allocation, especially to the root system, and the implications for the acquisition of nutrients by plants in response to CO₂ increase.Abbreviations N soil fertilization without nitrogen - N soil fertilization with nitrogen     

Elevated CO2 temporally enhances phosphorus immobilization in the rhizosphere of wheat and chickpea

Aims

The efficient management of phosphorus (P) in cropping systems remains a challenge due to climate change. We tested how plant species access P pools in soils of varying P status (Olsen-P 3.2–17.6 mg?kg^?1), under elevated atmosphere CO₂ (eCO₂).

Methods

Chickpea (Cicer arietinum L.) and wheat (Triticum aestivum L.) plants were grown in rhizo-boxes containing Vertosol or Calcarosol soil, with two contrasting P fertilizer histories for each soil, and exposed to ambient (380 ppm) or eCO₂ (700 ppm) for 6 weeks.

Results

The NaHCO₃-extractable inorganic P (Pi) in the rhizosphere was depleted by both wheat and chickpea in all soils, but was not significantly affected by CO₂ treatment. However, NaHCO₃-extractable organic P (Po) accumulated, especially under eCO₂ in soils with high P status. The NaOH-extractable Po under eCO₂ accumulated only in the Vertosol with high P status. Crop species did not exhibit different eCO₂-triggered capabilities to access any P pool in either soil, though wheat depleted NaHCO₃-Pi and NaOH-Pi in the rhizosphere more than chickpea. Elevated CO₂ increased microbial biomass C in the rhizosphere by an average of 21 %. Moreover, the size in Po fractions correlated with microbial C but not with rhizosphere pH or phosphatase activity.

Conclusion

Elevated CO₂ increased microbial biomass in the rhizosphere which in turn temporally immobilized P. This P immobilization was greater in soils with high than low P availability.     

Effects of Elevated Atmospheric Carbon Dioxide and Temperature on Soil Respiration in a Boreal Forest Using δ13C as a Labeling Tool

This study examines the effect of elevated atmospheric carbon dioxide [CO₂] (+340 ppm, ¹³C-depleted) and/or elevated air temperature (2.8–3.5°C) on the rate and δ¹³C of soil respiration. The study was conducted in a boreal Norway spruce forest using temperature-controlled whole-tree chambers and ¹³C as a marker for root respiration. The δ¹³C of needle carbohydrates was followed after the onset of the CO₂ treatment in August 2001 and during a 2.5-week period in the summer of 2002. Averaged over the growing seasons of 2002 and 2003, we observed a 48% and 62% increase, respectively, in soil respiration in response to elevated [CO₂], but no response to elevated air temperature. The percentage increase in response to elevated [CO₂] varied seasonally (between 10% and 190% relative to the control), but the absolute increase varied less (39 ± 11 mg C m⁻² h⁻¹; mean ± SD). Data on δ¹³C of soil respiration indicate that this increase in soil respiration rate resulted from increased root/rhizosphere respiration of recently fixed carbon. Our results support the hypothesis that root/rhizosphere respiration is sensitive to variation in substrate availability.