Carbon distribution in grass (Festuca pratensis L.) during regrowth after cutting—utilization of stored and newly assimilated carbon

Distribution of net assimilated C in meadow fescue (Fectuca pratensi L.) was followed before and after cutting of the shoots. Plants were continuously labelled in a growth chamber with ¹⁴C-labelled CO₂ in the atmosphere from seedling to cutting and with ¹³C-labelled CO₂ in the atmosphere during regrowth after the cutting. Labelled C, both ¹⁴C and ¹³C, was determined at the end of the two growth periods in shoots, crowns, roots, soil and rhizosphere respiration. Distribution of net assimilated C followed almost the same pattern at the end of the two growth periods, i.e. at the end of the ¹⁴C- and the ¹³C-labelling periods. Shoots retained 71–73% of net assimilated C while 9% was detected in the roots and 11–14% was released from the roots, determined as labelled C in soil and as rhizosphere respiration. At the end of the 2nd growth period, after cutting and regrowth, 21% of the residual plant ¹⁴C at cutting (¹⁴C in crowns and roots) was found in the new shoot biomass. A minor part of the residual plant ¹⁴C, 12%, was lost from the plants. The decreases in ¹⁴C in crowns and roots during the regrowth period suggest that ¹⁴C in both crowns and roots was translocated to new shoot tissue. Approximately half of the total root C at the end of the regrowth period after cutting was ¹³C-labelled C and thus represents new root growth. Root death after cutting could not be determined in this experiment, since the decline in root ¹⁴C during the regrowth period may also be assigned to root respiration, root exudation and translocation to the shoots. ei]{gnH}{fnLambers} ei]{gnA C}{fnBorstlap}     

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 translocation to the rhizosphere of maize and wheat and influence on the turnover of native soil organic matter at different soil nitrogen levels

Wheat and maize were grown in a growth chamber with the atmospheric CO₂ continuously labelled with ¹⁴C to study the translocation of assimilated carbon to the rhizosphere. Two different N levels in soil were applied. In maize 26–34% of the net assimilated ¹⁴C was translocated below ground, while in wheat higher values (40–58%) were found. However, due to the much higher shoot production in maize the total amount of carbon translocated below ground was similar to that of wheat. At high N relatively more of the C that was translocated to the root, was released into the soil due to increased root respiration and/or root exudation and subsequent microbial utilization and respiration. The evolution rate of unlabelled CO₂ from the native soil organic matter decreased after about 25 days when wheat was grown at high N as compared to low N. This negative effect of high N in soil was not observed with maize.     

Root systems and root:mass ratio-carbon allocation under current and projected atmospheric conditions in arable crops

Roots of annual crop plants are a major sink for carbon particularly during early, vegetative growth when up to one-half of all assimilated carbon may be translocated belowground. Flowering marks a particularly important change in resource allocation, especially in determinate species, with considerably less allocation to roots and, depending on environmental conditions, there may be insufficient for maintenance. Studies with ¹⁴C indicate the rapid transfer belowground of assimilates with typically 50% translocated in young cereal plants of which 50% is respired; exudation/rhizodeposition is generally <5% of the fixed carbon. Root: total plant mass decreases through the season and is affected by soil and atmospheric conditions. Limited water availability increased the allocation of ¹³C to roots of wheat grown in columns so that at booting 0.38 of shoot C (ignoring shoot respiration) was belowground compared to 0.31 in well-watered plants. Elevated CO₂ (700 mol CO₂ mol^–1 air) increased the proportion of root:total mass by 55% compared with normal concentration, while increasing the air temperature by a mean of 3 °C decreased the proportion from 0.093 in the cool treatment to 0.055 in the warm treatment.     

Assimilate partitioning affects 13C fractionation of recently assimilated carbon in maize

Coupling ¹³C natural abundance and ¹⁴C pulse labelling enabled us to investigate the dependence of ¹³C fractionation on assimilate partitioning between shoots, roots, exudates, and CO₂ respired by maize roots. The amount of recently assimilated C in these four pools was controlled by three levels of nutrient supply: full nutrient supply (NS), 10 times diluted nutrient supply (DNS), and deionised water (DW). After pulse labelling of maize shoots in a ¹⁴CO₂ atmosphere, ¹⁴C was traced to determine the amounts of recently assimilated C in the four pools and the δ¹³C values of the four pools were measured. Increasing amounts of recently assimilated C in the roots (from 8% to 10% of recovered ¹⁴C in NS and DNS treatments) led to a 0.3‰ ¹³C enrichment from NS to DNS treatments. A further increase of C allocation in the roots (from 10% to 13% of recovered ¹⁴C in DNS and DW treatments) resulted in an additional enrichment of the roots from DNS to DW treatments by 0.3‰. These findings support the hypothesis that ¹³C enrichment in a pool increases with an increasing amount of C transferred into that pool. δ¹³C of CO₂ evolved by root respiration was similar to that of the roots in DNS and DW treatments. However, if the amount of recently assimilated C in root respiration was reduced (NS treatment), the respired CO₂ became 0.7‰ ¹³C depleted compared to roots. Increasing amounts of recently assimilated C in the CO₂ from NS via DNS to DW treatments resulted in a 1.6‰ δ¹³C increase of root respired CO₂ from NS to DW treatments. Thus, for both pools, i.e. roots and root respiration, increasing amounts of recently assimilated C in the pool led to a δ¹³C increase. In DW and DNS plants there was no ¹³C fractionation between roots and exudates. However, high nutrient supply decreased the amount of recently assimilated C in exudates compared to the other two treatments and led to a 5.3‰ ¹³C enrichment in exudates compared to roots. We conclude that ¹³C discrimination between plant pools and within processes such as exudation and root respiration is not constant but strongly depends on the amount of C in the respective pool and on partitioning of recently assimilated C between plant pools. Section Editor: H. Lambers     

Differences in rhizosphere carbon-partitioning among plant species of different families

Interspecific variations in carbon (C) allocation and partitioning in the rhizosphere were investigated on 12 Mediterranean species belonging to different family groups (grasses, legumes, non-legume forbs) and having different life cycles. Plants grown individually in artificial soil, in a greenhouse and inoculated with rhizosphere microflora were labelled with ¹⁴CO₂ for 3 h at the vegetative stage. Rhizosphere respiration was measured during 6 days after which labelled C partitioning between shoots, roots, soil, root washing solution and respiration was estimated. The percentage of assimilated ¹⁴C allocated below ground differed significantly between species (41 – 76%) but no significant difference was found between grasses, legumes and non-legume forbs. When expressed as percentage of below-ground ¹⁴C, rhizosphere respiration was significantly smaller for non-legume forbs (42%) than for grasses (46%) and legumes (51%). Consequently more ¹⁴C was incorporated into root biomass in the former. Half-life of ¹⁴CO₂ evolution through respiration ranged from 23 h in legumes to 27 h for non-legume forbs and 37 h for grasses. This suggested differences in microbial activities due to quantities and quality of root exuded C. Rhizosphere respiration was positively correlated with the amount of ¹⁴C in the solution used to wash the roots on one hand, and root N concentration on the other hand. This led to a functional hierarchy between plant family groups of the overall rhizosphere activity. It went from non-legume forbs being the less active (except Crepis sancta)in terms of respiration and exudation, to grasses and then legumes, the most active but also the richest in nitrogen.     

Forest soil respiration rate and δ13C is regulated by recent above ground weather conditions

Soil respiration, a key component of the global carbon cycle, is a major source of uncertainty when estimating terrestrial carbon budgets at ecosystem and higher levels. Rates of soil and root respiration are assumed to be dependent on soil temperature and soil moisture yet these factors often barely explain half the seasonal variation in soil respiration. We here found that soil moisture (range 16.5–27.6% of dry weight) and soil temperature (range 8–17.5°C) together explained 55% of the variance (cross-validated explained variance; Q²) in soil respiration rate (range 1.0–3.4 mol C m^–2 s^–1) in a Norway spruce (Picea abies) forest. We hypothesised that this was due to that the two components of soil respiration, root respiration and decomposition, are governed by different factors. We therefore applied PLS (partial least squares regression) multivariate modelling in which we, together with below ground temperature and soil moisture, used the recent above ground air temperature and air humidity (vapour pressure deficit, VPD) conditions as x-variables. We found that air temperature and VPD data collected 1–4 days before respiration measurements explained 86% of the seasonal variation in the rate of soil respiration. The addition of soil moisture and soil temperature to the PLS-models increased the Q² to 93%. ¹³C analysis of soil respiration supported the hypotheses that there was a fast flux of photosynthates to root respiration and a dependence on recent above ground weather conditions. Taken together, our results suggest that shoot activities the preceding 1–6 days influence, to a large degree, the rate of root and soil respiration. We propose this above ground influence on soil respiration to be proportionally largest in the middle of the growing season and in situations when there is large day-to-day shifts in the above ground weather conditions. During such conditions soil temperature may not exert the major control on root respiration.     

Contribution of Lolium perenne rhizodeposition to carbon turnover of pasture soil

Carbon rhizodeposition and root respiration during eight development stages of Lolium perenne were studied on a loamy Gleyic Cambisol by ¹⁴CO₂ pulse labelling of shoots in a two compartment chamber under controlled laboratory conditions. Total ¹⁴CO₂ efflux from the soil (root respiration, microbial respiration of exudates and dead roots) in the first 8 days after ¹⁴C pulse labelling decreased during plant development from 14 to 6.5% of the total ¹⁴C input. Root respiration accounted for was between 1.5 and 6.5% while microbial respiration of easily available rhizodeposits and dead root remains were between 2 and 8% of the ¹⁴C input. Both respiration processes were found to decline during plant development, but only the decrease in root respiration was significant. The average contribution of root respiration to total ¹⁴CO₂ efflux from the soil was approximately 41%. Close correlation was found between cumulative ¹⁴CO₂ efflux from the soil and the time when maximum ¹⁴CO₂ efflux occurred (r=0.97). The average total of CO₂ Defflux from the soil with Lolium perenne was approximately 21 μg C-CO₂ d⁻¹ g⁻¹. It increased slightly during plant development. The contribution of plant roots to total CO₂ efflux from the soil, calculated as the remainder from respiration of bare soil, was about 51%. The total ¹⁴C content after 8 days in the soil with roots ranged from 8.2 to 27.7% of assimilated carbon. This corresponds to an underground carbon transfer by Lolium perenne of 6–10 g C m⁻² at the beginning of the growth period and 50–65 g C m⁻² towards the end of the growth period. The conventional root washing procedure was found to be inadequate for the determination of total carbon input in the soil because 90% of the young fine roots can be lost. This revised version was published online in June 2006 with corrections to the Cover Date. This revised version was published online in June 2006 with corrections to the Cover Date. This revised version was published online in June 2006 with corrections to the Cover Date.     

Distribution of assimilated carbon in plants and rhizosphere soil of basket willow (Salix viminalis L.)

Willow is often used in bio-energy plantations for its potential to function as a renewable energy source, but knowledge about its effect on soil carbon dynamics is limited. Therefore, we investigated the temporal variation in carbon dynamics in willow, focusing on below-ground allocation and sequestration to soil carbon pools. Basket willow plants (Salix viminalis L.) in their second year of growth were grown in pots in a greenhouse. At five times during the plants growth, namely 0, 1, 2, 3 and 4 months after breaking winter dormancy, a subset of the plants were continuously labelled with ¹⁴CO₂ in an ESPAS growth chamber for 28 days. After the labelling, the plants were harvested and separated into leaves, first and second year stems and roots. The soil was analysed for total C and ¹⁴C content as well as soil microbial biomass. Immediately after breaking dormancy, carbon stored in the first year stems was relocated to developing roots and leaves. Almost half the newly assimilated C was used for leaf development the first month of growth, dropping to below 15% in the older plants. Within the second month of growth, secondary growth of the stem became the largest carbon sink in the system, and remained so for the older age classes. Between 31 and 41% of the recovered ¹⁴C was allocated to below-ground pools. While the fraction of assimilated ¹⁴C in roots and root+soil respiration did not vary with plant age, the amount allocated to soil and soil microbial biomass increased in the older plants, indicating an increasing rhizodeposition. The total amount of soil microbial biomass was 30% larger in the oldest age class than in an unplanted control soil. The results demonstrate a close linkage between photosynthesis and below-ground carbon dynamics. Up to 13% of the microbial biomass consisted of carbon assimilated by the willows within the past 4 weeks, up to 11% of the recovered ¹⁴C was found as soil organic matter.     

How does nitrogen availability alter rhizodeposition in Lolium multiflorum Lam. during vegetative growth?

The objective of this work was to determine if the impact of nitrogen (N) on the release of organic carbon (C) into the soil by roots (rhizodeposition) correlated with the effect of this nutrient on some variables of plant growth. Lolium multiflorum Lam. was grown at two levels of N supply, either in sterile sand percolated with nutrient solution or in non-sterile soil. The axenic sand systems allowed continuous quantification of rhizodeposition and accurate analysis of root morphology whilst the soil microcosms allowed the study of ¹⁴C labelled C flows in physico-chemical and biological conditions relevant to natural soils. In the axenic sand cultures, enhanced N supply strongly increased the plant biomass, the plant N content and the shoot to root ratio. N supply altered the root morphology by increasing the root surface area and the density of apices, both being significantly positively correlated with the rate of organic C release by plant roots before sampling. This observation is consistent with the production of mucilage by root tips and with mechanisms of root exudation reported previously in the literature, i.e. the passive diffusion of roots solutes along the root with increased rate behind the root apex. We proposed a model of root net exudation, based on the number of root apices and on root soluble C that explained 60% of the variability in the rate of C release from roots at harvest. The effects of N on plant growth were less marked in soil, probably related to the relatively high supply of N from non-fertiliser soil-sources. N fertilization increased the shoot N concentration of the plants and the shoot to root ratio. Increased N supply decreased the partitioning of ¹⁴C to roots. In parallel, N fertilisation increased the root soluble ¹⁴C and the ¹⁴C recovered in the soil per unit of root biomass, suggesting a stimulation of root exudation by N supply. However, due to the high concentration of N in our unfertilised plants, this stimulation was assumed to be very weak because no significant effect of N was observed on the microbial C and on the bacterial abundance in the rhizosphere. Considering the difficulties in evaluating rhizodeposition in non sterile soil, it is suggested that the root soluble C, the root surface area and the root apex density are additional relevant variables that should be useful to measure along with the variables that are commonly determined when investigating how plant functioning impacts on the release of C by roots (i.e soil C, C of the microbial biomass, rhizosphere respiration).     

Influence of light intensity on the distribution of carbon and consequent effects on mineralization of soil nitrogen in a barley (Hordeum vulgare L.)-soil system

A pot experiment was conducted in a ¹⁴C-labelled atmosphere to study the influence of living plants on organic-N mineralization. The soil organic matter had been labelled, by means of a 200-days incubation, with ¹⁵N. The influence of the carbon input from the roots on the formation of microbial biomass was evaluated by using two different light intensities (I). Mineralization of ¹⁵N-labelled soil N was examined by following its fate in both the soil biomass and the plants. Less dry matter accumulated in shoots and roots at the lower light intensity. Furthermore, in all the plant-soil compartments examined, with the exception of rhizosphere respiration, the proportion of net assimilated ¹⁴C was lower in the low-I treatment than in the high-I treatment. The lower rates of ¹⁴C and ¹⁵N incorporation into the soil biomass were associated with less root-derived ¹⁴C. During the chamber period (¹⁴CO₂-atmosphere), mineralized amounts of ¹⁵N (measured as plant uptake of ¹⁵N) were small and represented about 6.8 to 7.8% of the initial amount of organic ¹⁵N in the soil. Amounts of unlabelled N found in the plants, as a percentage of total soil N, were 2.5 to 3.3%. The low availability of labelled N to microorganisms was the result of its stabilization during the 210 days of soil incubation. Differences in carbon supply resulted in different rates of N mineralization which is consistent with the hypothesis that roots induce N mineralization. N mineralization was higher in the high-I treatment. On the other hand, the rate of mineralization of unlabelled stable soil N was lower than labelled soil ¹⁵N which was stabilized. The amounts of ¹⁵N mineralized in planted soil during the chamber period (43 days) which were comparable with those mineralized in unplanted soil incubated for 210 days, also suggested that living plants increased the turnover rate of soil organic matter.     

The fate and path of assimilation products in the stem of 8-year-old Scots pine (Pinus sylvestris L.) trees

Summary ¹⁴CO₂ at ambient concentration was administered to a section of an upper branch of 8-year-old Scots pines and the import of radiocarbon into the stem and roots was determined after various chase periods. ¹⁴CO₂ fixation was performed in October when export of carbon into the stems and roots was maximal. In the short-term experiments the trees were harvested 1 h, 2 days and 5 days after a 3-h ¹⁴C pulse, while chase periods of 5 or 8 months were used in the long-term experiments. Loss of ¹⁴C was initially substantial, and even after a 5-day chase had not come down to a rate which indicated decrease only by respiration. After 5 days, more than 10% of the recovered radiocarbon (53% of the ¹⁴C translocated into the stem) had entered the roots and approximately the same amount was found in the stem. Extension of the chase period beyond 5 months did not result in a further significant loss of ¹⁴C by respiration, and the bulk of the label could be localized in the cell-wall fraction. No substantial redistribution of radiocarbon prior and subsequent to the formation of the new shoots could be observed, thus indicating that the stored material was utilized for thickening the stem and roots. Radioautography of stem cross-sections revealed a narrow helical strip of ¹⁴C from the feeding branch to the root in the phloem region. In the tree harvested after bud break the utilization of the ¹⁴C-labelled material stored in the stem for the production of the first layers of earlywood and the corresponding phloem was apparent.     

Carbon distribution and nitrogen partitioning in a soil-plant system with barley (Hordeum vulgare L.), ryegrass (Lolium perenne) and rape (Brassica napus L.) grown in a14CO2-atmosphere

To examine the influence of plant-microorganism interactions on soil-N transformations (e.g. net mineralization, net immobilization) a pot experiment was conducted in a¹⁴C-labelled atmosphere by using different (two annuals, one perennial) plants species. It was assumed that variation in below-ground, microorganism-available C would influence N transformations in soil. Plant species were fertilized (low rate) with¹⁵N-labelled nitrogen and grown, during days 13 and 62 after germination, in a growth chamber with a¹⁴C-labelled atmosphere. Nitrification was inhibited by using nitrapyrin (N-Serve). During the chamber period, shoots were harvested, and associated roots and soil were collected on two sampling occasionm, e.g. after 4 and 7 weeks in the growth chamber.The distribution of net (%) assimilated¹⁴C was significantly affected by both plant and time factors, and there was a significant plant × time interaction. There were significant differences between plants in all plant-soil compartments examined as well as in the degree of the plant × time interaction.Differences in the¹⁴C distribution between plants were due to both interspecific and developmental variation. In general, when comparing¹⁵N and¹⁴C quantities between species, many of the differences found between plants can be explained by the differences determined in the weight of shoot or root parts. Despite the fact that amounts of C released were greater in ryegrass than in the other plant-treatments no unequivocal evidence was found to show that the effects of plant-microorganism interactions on soil-N mineralization were greater under ryegrass. Possible mechanisms accounting for the partitioning of N found among plant biomass, soil biomass and soil residues are discussed.     

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.     

The use of C14O2 canopy techniques for measuring carbon transfer through the plant-soil system

Summary Methods for labelling growing plants by exposing them to C¹⁴O₂ under a cellulose acetate-butyrate canopy have been developed for laboratory and field use. The length of labelling ranged from 2 to 33 days and the C¹⁴O₂ content of the atmosphere was automatically controlled. This made it possible to measure carbon assimilation by the plants, transfer of photosynthates beneath ground and respiration of the roots.In the laboratory, root respiration of wheat plants was measured by separating the above and beneath ground plant parts using a RTV rubber partition. Half to two thirds of the assimilated carbon was found above ground, 15 to 25 per cent in the roots and shoot bases below the partition and 17 to 25 per cent was lost by underground respiration. The variability of these proportions was related to the stage of maturity of the plants.On native grassland, the relative above and beneath ground productivity was 50 per cent. The time required for the photosynthates to reach the roots at various depths ranged from 1 to 5 days and the amount of material deposited in the roots changed with time and soil moisture content. The use of tubes inserted at various depths beneath the canopy permitted sampling of soil air for C¹⁴ and CO₂ measurements. The soil C¹⁴O₂ flux indicated that root respiration during 8 days accounted for 24 per cent of the labelled carbon translocated to the roots after a two days labelling period.     

Estimating seasonal and annual carbon inputs,and root decomposition rates in a temperate pasture following field 14C pulse-labelling

Using a ¹⁴C pulse-labelling technique, we studied the seasonal changes in assimilation and partitioning of photoassimilated C in the plant–root–soil components of a temperate pasture. Pasture and soil samples were taken after 4-h, and 35-day chase periods, to examine these seasonal ¹⁴C fluxes. Total C and ¹⁴C were determined in the shoot, root and soil system. The amounts of C translocated annually to roots and soil were also estimated from the seasonal ¹⁴C distribution and pasture growth. The in situ field decomposition of newly formed roots during different seasons, also using ¹⁴C-labelling, was studied for one year in undisturbed rhizosphere soil. The ¹⁴C-labelled roots were sampled five times and decomposition rates were calculated assuming first-order decomposition.Annual pasture production at the site was 16 020 kg DM ha^–1, and pasture growth varied with season being highest (75–79 kg ha^–1 d^–1) in spring and lowest (18–20 kg ha^–1 d^–1) in winter. The above- and below-ground partitioning of ¹⁴C also varied with the season. The respiratory ¹⁴C–CO₂ losses, calculated as the difference between the total amounts of ¹⁴C recovered in the soil-plant system at 4 h and 35 days, were high (66–70%) during the summer, autumn and winter season, and low (37–39%) during the spring and late-spring season. Pasture plants partitioned more C below-ground during spring compared with summer, autumn and winter seasons. Overall, at this high fertility dairy pasture site, 18 220 kg C/ha was respired, 6490 kg remained above-ground in the shoot, and 6820 kg was translocated to roots and 1320 kg to soil. Root decomposition rate constant (k) differed widely with the season and were the highest for the autumn roots. The half-life was highest (111 days) for autumn roots and lowest (64 days) for spring roots. About one-third of the root label measured in the spring season disappeared in the first 5 weeks after the initial 35 Day of allocation period. The late spring, summer, late summer and winter roots had intermediate half-lives (88–94 days). These results indicate that seasonal changes in root growth and decomposition should be accounted for to give a better quantification of root turnover.     

Model for rhizodeposition and CO2 efflux from planted soil and its validation by 14C pulse labelling of ryegrass

A model for rhizodeposition and root respiration was developed and parameterised based on ¹⁴C pulse labelling of Lolium perenne. The plants were grown in a two-compartment chamber on a loamy Haplic Luvisol under controlled laboratory conditions. The dynamics of ¹⁴CO₂ efflux from the soil and ¹⁴C content in shoots, roots, micro-organisms, dissolved organic carbon (DOC) and soil were measured during the first 11 days after labelling. Modelled parameters were estimated by fitting on measured ¹⁴C dynamics in the different pools. The model and the measured ¹⁴C dynamics in all pools corresponded well (r ²=0.977). The model describes well ¹⁴CO₂ efflux from the soil and ¹⁴C dynamics in shoots, roots and soil, but predicts unsatisfactorily the ¹⁴C content in micro-organisms and DOC. The model also allows for division of the total ¹⁴CO₂ efflux from the soil in ¹⁴CO₂ derived from root respiration and ¹⁴CO₂ derived from rhizomicrobial respiration by use of exudates and root residues. Root respiration and rhizomicrobial respiration amounted for 7.6% and 6.0% of total assimilated C, respectively, which accounts for 56% and 44% of root-derived ¹⁴CO₂ efflux from the soil planted with 43-day-old Lolium perenne, respectively. The sensitivity analysis has shown that root respiration rate affected the curve of ¹⁴CO₂ efflux from the soil mainly during the first day after labelling. The changes in the exudation rate influenced the ¹⁴CO₂ efflux later than first 24 h after labelling.     

Turnover and distribution of root exudates of Zea mays

Decomposition and distribution of root exudates of Zea mays L. were studied by means of ¹⁴CO₂ pulse labeling of shoots on a loamy Haplic Luvisol. Plants were grown in two-compartment pots, where the lower part was separated from the roots by monofilament gauze. Root hairs, but not roots, penetrated through the gauze into the lower part of the soil. The root-free soil in the lower compartment was either sterilized with cycloheximide and streptomycin or remained non-sterile. In order to investigate exudate distribution, 3 days after the ¹⁴C labeling, the lower soil part was frozen and sliced into 15, one-mm thick layers using a microtome. Cumulative ¹⁴CO₂ efflux from the soil during the first 3 days after ¹⁴C pulse labeling did not change during plant growth and amounted to about 13–20% of the total recovered ¹⁴C (41–55% of the carbon translocated below ground). Nighttime rate of total CO₂ efflux was 1.5 times lower than during daytime because of tight coupling of exudation with photosynthesis intensity. The average CO₂ efflux from the soil with Zea mays was about 74 g C g^–1 day^–1 (22 g C m^–2 day^–1), although, the contribution of plant roots to the total CO₂ efflux from the soil was about 78%, and only 22% was respired from the soil organic matter. Zea mays transferred about 4 g m^–2 of carbon under ground during 26 days of growth. Three zones of exudate concentrations were identified from the distribution of the ¹⁴C-activity in rhizosphere profiles after two labeling periods: (1) 1–2 (3) mm (maximal concentration of exudates) 2) 3–5 mm (presence of exudates is caused by their diffusion from the zone 1); (3) 6–10 mm (very insignificant amounts of exudates diffused from the previous zones). At the distance further than 10 mm no exudates were found. The calculated coefficient of exudate diffusion in the soil was 1.9 × 10^–7 cm² s^–1.     

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 and nitrogen in the root-zone of barley (Hordeum vulgare L.) supplied with nitrogen fertilizer at two rates

Below-ground carbon (C) production and nitrogen (N) flows in the root-zone of barley supplied with high or low amounts of N-fertilizer were investigated. Interest was focused on the effect of the level of N-fertilizer on the production of root-derived C and on gross immobilization (i) and gross mineralization (m) rates. The plants were grown for 46 days in a sandy loam soil. Principles of pool dilution and changes in ¹⁵N pool abundances were used in conjunction with mathematical modelling to calculate the flows of N. N was applied at a high or a low rate, as (¹⁵NH₄)₂SO₄ solution (17.11 atom% ¹⁵N excess), before sowing. Nitrification was inhibited by using nitrapyrin (N-Serve). Pots were sampled four or five times during the experimental period, i.e. 0, 22, 30, 38 and 46 days after germination. On the three last sampling occasions, samples were also collected from pots in a growth chamber with ¹⁴C-labelled atmosphere.The release of ¹⁴C, measured as the proportion of the total ¹⁴C translocated below ground, was higher in the high-N treatment, but the differences between treatments were small. Our results were not conclusive in demonstrating that high-N levels stimulate the decomposition and microbial utilization of root-released materials. However, the internal circulation of soil-N, calculated N fluxes (m), which were in accordance with C mineralization rates and amounts of unlabelled N found in the plants (PU), suggested that the decomposition of native soil organic matter was hampered in the high-N treatment. Apparently, towards the end of the experimental period, microorganisms in the low-N treatment used C from soil organic matter to a greater extent than C they used from root released material, presumably because lower amounts of mineral N were available to microorganisms in the low-N treatment. Immobilization of N appeared to be soil driven (organisms decomposing soil organic matter account for the N demand) at low-N and root-driven (organisms decomposing roots and root-derived C account for the N demand) at high-N.Abbreviations AU Ammonium N-unlabelled - AL Ammonium N-labelled - AT Ammonium N-labelled and unlabelled (total) - NU Nitrate N-unlabelled - OU Organic N-unlabelled - OL Organic N-labelled - OT Organic N-total - PU Plant N-unlabelled (shoots and roots) - PL Plant N-labelled (shoots and roots) - PT Plant N-total (shoots and roots) - SL Sink or source of N-labelled - S Source or sink of N, mainly to and from the outer part of the cylinder - SU Sink or source of N-unlabelled - m Mineralization rate - i Immobilization rate - ua Uptake of ammonium - un Uptake of nitrate - la Loss of ammonium.