Carbon allocation and carbon transfer between t Betula papyrifera and t Pseudotsuga menziesii seedlings using a 13C pulse-labeling method

Here we describe a simple method for pulse-labeling tree seedlings with ¹³CO_2(gas), and then apply the method in two related experiments: t (i) comparison of carbon allocation patterns between t Betula papyrifera Marsh. and t Pseudotsuga menziesii (Mirb.) Franco, and t (ii) measurement of one-way belowground carbon transfer from t B. papyrifera to t P. menziesii. Intraspecific carbon allocation patterns and interspecific carbon transfer both influence resource allocation, and consequently development, in mixed communities of t B. papyrifera and t P. menziesii.In preparation for the two experiments, we first identified the appropriate ¹³CO_2(gas) pulse-chase regime for labeling seedlings: a range of pulse (100-mL and 200-mL 99 atom%¹³ CO_2(gas)) and chase (0, 3 and 6 d) treatments were applied to one year-old t B. papyrifera and t P. menziesii seedlings. The amount of ¹³CO₂ fixed immediately after 1.5 h exposure was greatest for both t B. papyrifera (40.8 mg excess ¹³C) and t P. menziesii (22.9 mg excess ¹³C) with the 200-mL pulse, but higher ¹³C loss and high sample variability resulted in little difference in excess¹³ C content between pulse treatments after 3 d for either species. The average excess ¹³C root/shoot ratio of t B. papyrifera and t P. menziesii changed from 0.00 immediately following the pulse to 0.61 and 0.87 three and six days later, which reflected translocation of 75% of fixed isotope out of foliage within 3 d following the pulse and continued enrichment in fine roots over 6 d. Based on these results, the 100-mL CO_2(gas) and 6-d chase were considered appropriate for the carbon allocation and belowground transfer experiments.In the carbon allocation experiment, we found after 6 d that t B. papyrifera allocated 49% (average 9.5 mg) and t P. menziesii 41% (average 5.8 mg) of fixed isotope to roots, of which over 55% occurred in fine roots in both species. Species differences in isotope allocation patterns paralleled differences in tissue biomass distribution. The greater pulse labeling efficiency of t B. papyrifera compared to t P. menziesii was associated with its two-fold and 13- fold greater leaf and whole seedling net photosynthetic rates, respectively, 53% greater biomass, and 35% greater root/shoot ratio.For the carbon transfer experiment, t B. papyrifera and t P. menziesii were grown together in laboratory rootboxes, with their roots intimately mingled. A pulse of 100 mL¹³ CO_2(gas) was applied to paper birch and one-way transfer to neighboring t P. menziesii was measured after 6 d. Of the excess ¹³C fixed by t B. papyrifera, 4.7% was transferred to neighboring t P. menziesii, which distributed the isotope evenly between roots and shoots. Of the isotope received by t P. menziesii, we estimated that 93% was taken up through belowground pathways, and the remaining 7% taken up by foliage as¹³ CO_2(gas) respired by t B. papyrifera shoots. These two experiments indicate that t B. papyrifera fixes more total carbon and allocates a greater proportion to its root system than does t P. menziesii, giving it a competitive edge in resource gathering; however, below-ground carbon sharing is of sufficient magnitude that it may help ensure co-existence of the two species in mixed communities.     

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.     

Partitioning of belowground C in young sugar maple forest

Background and aims

Trees allocate a high proportion of assimilated carbon belowground, but the partitioning of that C among ecosystem components is poorly understood thereby limiting our ability to predict responses of forest C dynamics to global change drivers.

Methods

We labeled sugar maple saplings in natural forest with a pulse of photosynthetic ¹³C in late summer and traced the pulse over the following 3 years. We quantified the fate of belowground carbon by measuring ¹³C enrichment of roots, rhizosphere soil, soil respiration, soil aggregates and microbial biomass.

Results

The pulse of ¹³C contributed strongly to root and rhizosphere respiration for over a year, and respiration comprised about 75 % of total belowground C allocation (TBCA) in the first year. We estimate that rhizosphere carbon flux (RCF) during the dormant season comprises at least 6 % of TBCA. After 3 years, 3.8 % of the C allocated belowground was recovered in soil organic matter, mostly in water-stable aggregates.

Conclusions

A pulse of carbon allocated belowground in temperate forest supplies root respiration, root growth and RCF throughout the following year and a small proportion becomes stabilized in soil aggregates.     

A comparison of carbon flow from pre-labelled and pulse-labelled plants

Perennial rye-grass was subjected to two different¹⁴C labelling regimes to enable a partitioning of the carbon sources contributing to rhizosphere carbon-flow. Plant/soil microcosms were designed which enabled rye-grass plants to either receive a single pulse of¹⁴C?CO₂ or to be pre-labelled using a series of¹⁴C?CO₂ pulses, allowing the fate of newly photoassimilated carbon and carbon lost by root decomposition to be followed into the soil. For young rye-grass plants grown over a short period, rhizosphere carbon flow was found to be dominated by newly photoassimilated carbon. Evidence for this came from the observed percentage of the total¹⁴C budget (i.e. total¹⁴C?CO₂ fixed by the plants) lost from the root/soil system, which was 30 times greater for the pulse labelled compared to pre-labelled plants. Root decomposition was found to be less at 10°C compared to 20–25°C, though input of¹⁴C into the soil was the same at both temperatures.     

Rate of Belowground Carbon Allocation Differs with Successional Habit of Two Afromontane Trees

Background

Anthropogenic disturbance of old-growth tropical forests increases the abundance of early successional tree species at the cost of late successional ones. Quantifying differences in terms of carbon allocation and the proportion of recently fixed carbon in soil CO₂ efflux is crucial for addressing the carbon footprint of creeping degradation.

Methodology

We compared the carbon allocation pattern of the late successional gymnosperm Podocarpus falcatus (Thunb.) Mirb. and the early successional (gap filling) angiosperm Croton macrostachyus Hochst. es Del. in an Ethiopian Afromontane forest by whole tree ¹³CO₂ pulse labeling. Over a one-year period we monitored the temporal resolution of the label in the foliage, the phloem sap, the arbuscular mycorrhiza, and in soil-derived CO₂. Further, we quantified the overall losses of assimilated ¹³C with soil CO₂ efflux.

Principal Findings

¹³C in leaves of C. macrostachyus declined more rapidly with a larger size of a fast pool (64% vs. 50% of the assimilated carbon), having a shorter mean residence time (14 h vs. 55 h) as in leaves of P. falcatus. Phloem sap velocity was about 4 times higher for C. macrostachyus. Likewise, the label appeared earlier in the arbuscular mycorrhiza of C. macrostachyus and in the soil CO₂ efflux as in case of P. falcatus (24 h vs. 72 h). Within one year soil CO₂ efflux amounted to a loss of 32% of assimilated carbon for the gap filling tree and to 15% for the late successional one.

Conclusions

Our results showed clear differences in carbon allocation patterns between tree species, although we caution that this experiment was unreplicated. A shift in tree species composition of tropical montane forests (e.g., by degradation) accelerates carbon allocation belowground and increases respiratory carbon losses by the autotrophic community. If ongoing disturbance keeps early successional species in dominance, the larger allocation to fast cycling compartments may deplete soil organic carbon in the long run.     

Carbon input by roots into the soil: Quantification of rhizodeposition from root to ecosystem scale

Despite its fundamental role for carbon (C) and nutrient cycling, rhizodeposition remains ‘the hidden half of the hidden half’: it is highly dynamic and rhizodeposits are rapidly incorporated into microorganisms, soil organic matter, and decomposed to CO₂. Therefore, rhizodeposition is rarely quantified and remains the most uncertain part of the soil C cycle and of C fluxes in terrestrial ecosystems. This review synthesizes and generalizes the literature on C inputs by rhizodeposition under crops and grasslands (281 data sets). The allocation dynamics of assimilated C (after ¹³C‐CO₂ or ¹⁴C‐CO₂ labeling of plants) were quantified within shoots, shoot respiration, roots, net rhizodeposition (i.e., C remaining in soil for longer periods), root‐derived CO₂, and microorganisms. Partitioning of C pools and fluxes were used to extrapolate belowground C inputs via rhizodeposition to ecosystem level. Allocation from shoots to roots reaches a maximum within the first day after C assimilation. Annual crops retained more C (45% of assimilated ¹³C or ¹⁴C) in shoots than grasses (34%), mainly perennials, and allocated 1.5 times less C belowground. For crops, belowground C allocation was maximal during the first 1–2 months of growth and decreased very fast thereafter. For grasses, it peaked after 2–4 months and remained very high within the second year causing much longer allocation periods. Despite higher belowground C allocation by grasses (33%) than crops (21%), its distribution between various belowground pools remains very similar. Hence, the total C allocated belowground depends on the plant species, but its further fate is species independent. This review demonstrates that C partitioning can be used in various approaches, e.g., root sampling, CO₂ flux measurements, to assess rhizodeposits’ pools and fluxes at pot, plot, field and ecosystem scale and so, to close the most uncertain gap of the terrestrial C cycle.     

Peatland carbon efflux partitioning reveals that Sphagnum photosynthate contributes to the DOC pool

Over half of the world's peat originated from Sphagnum, representing 10–15% of the terrestrial carbon stock. However, information regarding the release and exudation of organic carbon by living Sphagnum plants into the surface peat is scarce. In this study, we examined the contribution of recent Sphagnum subnitens (Russ. and Warnst.) photosynthate carbon to the peatland dissolved organic carbon (DOC) pool. This was done using a ¹³CO₂ pulse-chase experimental approach during the growing season. Despite the importance of Sphagnum in long-term carbon accumulation, results showed that the Sphagnum community rapidly contributes recently synthesized carbon to the peatland DOC pool. We estimate that by 4 h up to 4% of the total DOC in peat leachate was derived from ¹³CO₂ pulse labelling at ambient CO₂ concentrations. Nonetheless, a huge 64% of the ¹³C initially assimilated by photosynthesis was retained in Sphagnum subnitens for 23 days after labelling, consistent with the role of Sphagnum in peatland carbon accumulation. The majority of ¹³C loss as respired CO₂ came within the few days post ¹³CO₂ labelling, suggesting that it was derived from plant respiration of photosynthates.     

Soil CO2 efflux and soil carbon balance of a tropical rubber plantation

Natural rubber is a valuable source of income in many tropical countries and rubber trees are increasingly planted in tropical areas, where they contribute to land-use changes that impact the global carbon cycle. However, little is known about the carbon balance of these plantations. We studied the soil carbon balance of a 15-year-old rubber plantation in Thailand and we specifically explored the seasonal dynamic of soil CO₂ efflux (F _S) in relation to seasonal changes in soil water content (W _S) and soil temperature (T _S), assessed the partitioning of F _S between autotrophic (R _A) and heterotrophic (R _H) sources in a root trenching experiment and estimated the contribution of aboveground and belowground carbon inputs to the soil carbon budget. A multiplicative model combining both T _S and W _S explained 58 % of the seasonal variation of F _S. Annual soil CO₂ efflux averaged 1.88 kg C m^?2 year^?1 between May 2009 and April 2011 and R _A and R _H accounted for respectively 63 and 37 % of F _S, after corrections of F _S measured on trenched plots for root decomposition and for difference in soil water content. The 4-year average annual aboveground litterfall was 0.53 kg C m^?2 year^?1 while a conservative estimate of belowground carbon input into the soil was much lower (0.17 kg C m^?2 year^?1). Our results highlighted that belowground processes (root and rhizomicrobial respiration and the heterotrophic respiration related to belowground carbon input into the soil) have a larger contribution to soil CO₂ efflux (72 %) than aboveground litter decomposition.     

Induced carbon reallocation and compensatory growth as root herbivore tolerance mechanisms

Upon attack by leaf herbivores, many plants reallocate photoassimilates below ground. However, little is known about how plants respond when the roots themselves come under attack. We investigated induced resource allocation in maize plants that are infested by the larvae Western corn rootworm Diabrotica virgifera virgifera. Using radioactive ¹¹CO₂, we demonstrate that root‐attacked maize plants allocate more new ¹¹C carbon from source leaves to stems, but not to roots. Reduced meristematic activity and reduced invertase activity in attacked maize root systems are identified as possible drivers of this shoot reallocation response. The increased allocation of photoassimilates to stems is shown to be associated with a marked thickening of these tissues and increased growth of stem‐borne crown roots. A strong quantitative correlation between stem thickness and root regrowth across different watering levels suggests that retaining photoassimilates in the shoots may help root‐attacked plants to compensate for the loss of belowground tissues. Taken together, our results indicate that induced tolerance may be an important strategy of plants to withstand belowground attack. Furthermore, root herbivore‐induced carbon reallocation needs to be taken into account when studying plant‐mediated interactions between herbivores.     

Fast assimilate turnover revealed by in situ 13CO2 pulse-labelling in Subarctic tundra

Climatic changes in Arctic regions are likely to have significant impacts on vegetation composition and physiological responses of different plant types, with implications for the regional carbon (C) cycle. Here, we explore differences in allocation and turnover of assimilated C in two Subarctic tundra communities. We used an in situ ¹³C pulse at mid-summer in Swedish Lapland to investigate C allocation and turnover in four contrasting tundra plant communities. We found a high rate of turnover of assimilated C in leaf tissues of Betula nana and graminoid vegetation at the height of the growing season, with a mean residence time of pulse-derived ¹³C of 1.1 and 0.7?days, respectively. One week after the pulse, c. 20 and 15%, respectively, of assimilated label-C remained in leaf biomass, representing most likely allocation to structural biomass. For the perennial leaf tissue of the graminoid communities, a remainder of approximately 5% of the pulse-derived C was still traceable after 1?year, whereas none was detectable in Betula foliage. The results indicate a relatively fast C turnover and small belowground allocation during the active growing season of recent assimilates in graminoid communities, with comparatively slower turnover and greater investment in belowground allocation by B. nana vegetation.     

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.     

Allocation and Turnover of Photosynthetically Assimilated CO(2) in Leaves of Glycine max L. Clark

The allocation and turnover of photosynthetically assimilated ¹⁴CO₂ in lipid and protein fractions of soybean (Glycine max L. Clark) leaves and stem materials was measured. In whole plant labeling experiments, allocation of photosynthate from a pulse of ¹⁴CO₂ into polymeric compounds was: 25% to proteins in 4 days, 20% to metabolically inert cell wall products in 1 to 2 days, 10% to lipids in 4 days, and 4% to starch in 1 day. The amount of ¹⁴C labeled photosynthate that an actively growing leaf (leaf 4) used for its own lipid synthesis immediately following pulse labeling was about 25%. The ¹⁴C of labeled proteins turned over with half-lives of 3.8, 3.3, and 4.1 days in leaves 1, 2, and 3, respectively; and turnover of ¹⁴C in total shoot protein proceeded with a half-life of 5.2 days. Three kinetic ¹⁴C turnover patterns were observed in lipids: a rapid turnover fraction (within a day), an intermediate fraction (half-life about 5 days), and a slow turnover fraction. These results are discussed in terms of previously published accounts of translocation, carbon budgets, carbon use, and turnover in starch, lipid, protein, and cell wall materials of various plants including soybeans.     

Labile carbon retention compensates for CO2 released by priming in forest soils

Increase of belowground C allocation by plants under global warming or elevated CO₂ may promote decomposition of soil organic carbon (SOC) by priming and strongly affects SOC dynamics. The specific effects by priming of SOC depend on the amount and frequency of C inputs. Most previous priming studies have investigated single C additions, but they are not very representative for litterfall and root exudation in many terrestrial ecosystems. We evaluated effects of ¹³C‐labeled glucose added to soil in three temporal patterns: single, repeated, and continuous on dynamics of CO₂ and priming of SOC decomposition over 6 months. Total and ¹³C labeled CO₂ were monitored to analyze priming dynamics and net C balance between SOC loss caused by priming and the retention of added glucose‐C. Cumulative priming ranged from 1.3 to 5.5 mg C g^?1 SOC in the subtropical, and from ?0.6 to 5.5 mg C g^?1 SOC in the tropical soils. Single addition induced more priming than repeated and continuous inputs. Therefore, single additions of high substrate amounts may overestimate priming effects over the short term. The amount of added glucose C remaining in soil after 6 months (subtropical: 8.1–11.2 mg C g^?1 SOC or 41‐56% of added glucose; tropical: 8.7–15.0 mg C g^?1 SOC or 43–75% of glucose) was substantially higher than the net C loss due to SOC decomposition including priming effect. This overcompensation of C losses was highest with continuous inputs and lowest with single inputs. Therefore, raised labile organic C input to soils by higher plant productivity will increase SOC content even though priming accelerates decomposition of native SOC. Consequently, higher continuous input of C belowground by plants under warming or elevated CO₂ can increase C stocks in soil despite accelerated C cycling by priming in soils.     

Drought and recovery effects on belowground respiration dynamics and the partitioning of recent carbon in managed and abandoned grassland

The supply of soil respiration with recent photoassimilates is an important and fast pathway for respiratory loss of carbon (C). To date it is unknown how drought and land‐use change interactively influence the dynamics of recent C in soil‐respired CO₂. In an in situ common‐garden experiment, we exposed soil‐vegetation monoliths from a managed and a nearby abandoned mountain grassland to an experimental drought. Based on two ¹³CO₂ pulse‐labelling campaigns, we traced recently assimilated C in soil respiration during drought, rewetting and early recovery. Independent of grassland management, drought reduced the absolute allocation of recent C to soil respiration. Rewetting triggered a respiration pulse, which was strongly fuelled by C assimilated during drought. In comparison to the managed grassland, the abandoned grassland partitioned more recent C to belowground respiration than to root C storage under ample water supply. Interestingly, this pattern was reversed under drought. We suggest that these different response patterns reflect strategies of the managed and the abandoned grassland to enhance their respective resilience to drought, by fostering their resistance and recovery respectively. We conclude that while severe drought can override the effects of abandonment of grassland management on the respiratory dynamics of recent C, abandonment alters strategies of belowground assimilate investment, with consequences for soil‐CO₂ fluxes during drought and drought‐recovery.     

Effects of temperature and nitrate on phosphomonoesterase activities between carbon source and sink tissues in Zostera marina L.

Inorganic phosphorus (P_i) is important in the regulation of many carbon and nitrogen metabolic processes of plants. In this study, we examined alterations of phosphomonoesterase activity (PA; both alkaline and acid) in a submersed marine angiosperm, Zostera marina, grown in P_i non-limiting conditions under elevated temperature and/or nitrate enrichment. Control plants (ambient water-column NO₃⁻ < 2.5 μM, with weekly mean water temperatures between 26.5-27.0 °C based on a 20-yr data set in a local embayment) were compared to treated plants that were exposed to increased water-column nitrate (8 μM NO₃⁻ above ambient, pulsed daily at 0900 h), and/or increased temperature (ca. 3 °C above weekly means) over eight weeks in late summer-fall. Under both nitrate regimes, increased temperature resulted in periodic increased leaf and root-rhizome tissue carbon content, and increased acid and alkaline PA activities (AcPAs and AlPAs, respectively). There was a positive correlation between AlPA and AcPA activities and sucrose synthase activities in belowground structures, and a negative correlation between AlPA activities and sucrose concentrations. There were also periodic changes in PA partitioning between carbon source and sink tissues. In high-temperature and high-nitrate treatments, AcPAs significantly increased in leaves relative to activities in root-rhizome tissues (up to 12-fold higher in aboveground than belowground tissues in as little as 3 weeks after initiation of treatments). These responses were not observed in control plants, which maintained comparable AcPA activities in above- and belowground tissues. In addition, AlPA activity was significantly higher in leaf than in root-rhizome tissues of plants in high-temperature (weeks 3 and 6) and high temperature combined with high nitrate treatments (week 8), relative to AlPA activities in control plants. The observed changes in PAs were not related to P_i growth limitation, and may allow Z. marina to alter its carbon metabolism during periods of increased carbon demand/mobilization. This response would make it possible for Z. marina to meet short-term P requirements to maximize carbon production/allocation. Such a mechanism could help to explain the variability in PA activities that has been observed for many plant species during periods when environmental P_i exceeds requirements for optimal growth.     

Partitioning pattern of carbon flux in a Kobresia grassland on the Qinghai‐Tibetan Plateau revealed by field 13C pulse‐labeling

Characterizing the carbon turnover in terrestrial ecosystems is critical for understanding and predicting carbon dynamics in ecosystems. We used in situ¹³C pulse labeling to track photosynthetic carbon fluxes from shoot to roots and to soil in a Kobresia humilis meadow on the Qinghai‐Tibet Plateau. We found that about 36.7% of labeled carbon was translocated out from the shoots within the first 24 h after photosynthetic uptake. This is equivalent to 66.1% of total ¹³C moving out from the shoot during the 32‐day chase period, indicating a rapid and large translocation of newly fixed carbon to belowground parts in these alpine plants. 58.7% of the assimilated ¹³C was transferred belowground. At the end of the chase phase, 30.9% was retained in living roots, 3.4% in dead roots, 17.2% lost as belowground respiration and 7.3% remained in the soil. In the four carbon pools (i.e., shoots, living roots, dead roots, and soil pools), living roots consistently had the highest proportion of ¹³C in the plant–soil system during the 32 days. Based on the ¹³C partitioning pattern and biomass production, we estimate a total of 4930 kg C ha^?1 was allocated belowground during the vegetation growth season in this alpine meadow. Of this, roots accumulated 2868 kg C ha^?1 and soils accumulated 613 kg C ha^?1. This study suggests that carbon storage in belowground carbon pools plays the most important role in carbon cycles in the alpine meadow.     

Use of decreasing foliar carbon isotope discrimination during water limitation as a carbon tracer to study whole plant carbon allocation

Foliar carbon isotope discrimination (Δ) of C₃ plants decreases in water‐deficit situations as discrimination by the photosynthetic primary carboxylation reaction decreases. This diminished Δ in leaves under water deficit can be used as a tracer to study whole plant carbon allocation patterns. Carbon isotope composition (δ¹³C value) of leaf hot water extracts or leaf tissue sap represents a short‐term integral of leaf carbon isotope discrimination and thus represents the δ¹³C value of source carbon that may be distributed within a plant in water‐deficit situations. By plotting the δ¹³C values of source carbon against the δ¹³C values of sink tissues, such as roots or stems, it is possible to assess carbon allocation to and incorporation into sink organs in relation to already present biomass. This natural abundance labelling method has been tested in three independent experiments, a one‐year field study with the fruit tree species Ziziphus mauritiana and peach (Prunus persica), a medium‐term drought stress experiment with Ziziphus rotundifolia trees in the glasshouse, and a short‐term drought stress experiment with soybean (Glycine max). The data show that the natural abundance labelling method can be applied to qualitatively assess carbon allocation in drought‐stressed plants. Although it is not possible to estimate exact fluxes of assimilated carbon during water deficit the method represents an easy to use tool to study integrated plant adaptations to drought stress. In addition, it is a less laborious method that can be applied in field studies as well as in controlled experiments, with plants from any developmental stage.     

Carbon flow in an upland grassland: effect of liming on the flux of recently photosynthesized carbon to rhizosphere soil

The effect of liming on the flow of recently photosynthesized carbon to rhizosphere soil was studied using ¹³CO₂ pulse labelling, in an upland grassland ecosystem in Scotland. The use of ¹³C enabled detection, in the field, of the effect of a 4‐year liming period of selected soil plots on C allocation from plant biomass to soil, in comparison with unlimed plots. Photosynthetic rates and carbon turnover were higher in plants grown in limed soils than in those from unlimed plots. Higher δ¹³C‰ values were detected in shoots from limed plants than in those from unlimed plants in samples clipped within 15 days of the end of pulse labelling. Analysis of the aboveground plant production corresponding to the 4‐year period of liming indicated that the standing biomass was higher in plots that received lime. Lower δ¹³C‰ values in limed roots compared with unlimed roots were found, whereas no significant difference was detected between soil samples. Extrapolation of our results indicated that more C has been lost through the soil than has been gained via photosynthetic assimilation because of pasture liming in Scotland during the period 1990–1998. However, the uncertainty associated with such extrapolation based on this single study is high and these estimates are provided only to set our findings in the broader context of national soil carbon emissions.     

Photoassimilated carbon allocation in a wheat plant-soil system as affected by soil fertility and land-use history

Background and aims

Carbon (C) cycling in terrestrial ecosystems is influenced by the distribution of photo-assimilated C in the plant-soil system. Photo-assimilated C allocation in a wheat cropping system was examined to identify the links between soil fertility, C partitioning and soil C sequestration.

Methods

A pulse labelling experiment was conducted where ¹⁴CO₂ was introduced to wheat plants grown in two groups of soils of varying fertility: arable soils spiked with nutrients, and soils with differing land-use histories. Wheat shoot, root and soil samples were taken 1, 14 and 28 days after pulse labelling to examine the fluxes of ¹⁴C in the plant-root-soil system.

Results

The partitioning of ¹⁴C in wheat plant-root-soil system was found to vary with time, nutrient spiked soil fertility and land-use history. At the end of the experiment using spiked soils, a positive correlation was observed between the allocation of ¹⁴C in the shoots and soil fertility, whereas in the roots, this relationship was negative. The overall allocation of ¹⁴C in the plant-root system differed significantly between the land-use histories; while in the spiked arable soils ¹⁴C allocation in the shoots and roots systematically followed their fertility status.

Conclusions

There was a weak relationship between C allocation and soil fertility in the soils of different land-use history compared to the strong relationship in the spiked arable soils. This suggests that other factors in the soils under different land uses were more important than nutrient status alone in driving photo-assimilated C allocation. This study demonstrated that soil fertility and land-use history have a crucial role in the allocation of photo-assimilated C in the plant-soil system and are important factors by which C sequestration in soil may be impacted.     

Soil temperature affects carbon allocation within arbuscular mycorrhizal networks and carbon transport from plant to fungus

How soil carbon balance will be affected by plant–mycorrhizal interactions under future climate scenarios remains a significant unknown in our ability to forecast ecosystem carbon storage and fluxes. We examined the effects of soil temperature (14, 20, 26 °C) on the structure and extent of a multispecies community of arbuscular mycorrhizal (AM) fungi associated with Plantago lanceolata. To isolate fungi from roots, we used a mesh‐divided pot system with separate hyphal compartments near and away from the plant. A ¹³C pulse label was then used to trace the flow of recently fixed photosynthate from plants into belowground pools and respiration. Temperature significantly altered the structure and allocation of the AM hyphal network, with a switch from more vesicles (storage) in cooled soils to more extensive extraradical hyphal networks (growth) in warmed soils. As soil temperature increased, we also observed an increase in the speed at which plant photosynthate was transferred to and respired by roots and AM fungi coupled with an increase in the amount of carbon respired per unit hyphal length. These differences were largely independent of plant size and rates of photosynthesis. In a warmer world, we would therefore expect more carbon losses to the atmosphere from AM fungal respiration, which are unlikely to be balanced by increased growth of AM fungal hyphae.