Daniel Epron

The production and turnover of extramatrical mycelium of ectomycorrhizal fungi in forest soils: role in carbon cycling

There is growing evidence of the importance of extramatrical mycelium (EMM) of mycorrhizal fungi in carbon (C) cycling in ecosystems. However, our understanding has until recently been mainly based on laboratory experiments, and knowledge of such basic parameters as variations in mycelial production, standing biomass and turnover as well as the regulatory mechanisms behind such variations in forest soils is limited. Presently, the production of EMM by ectomycorrhizal (EM) fungi has been estimated at ~140 different forest sites to be up to several hundreds of kg per ha per year, but the published data are biased towards Picea abies in Scandinavia. Little is known about the standing biomass and turnover of EMM in other systems, and its influence on the C stored or lost from soils. Here, focussing on ectomycorrhizas, we discuss the factors that regulate the production and turnover of EMM and its role in soil C dynamics, identifying important gaps in this knowledge. C availability seems to be the key factor determining EMM production and possibly its standing biomass in forests but direct effects of mineral nutrient availability on the EMM can be important. There is great uncertainty about the rate of turnover of EMM. There is increasing evidence that residues of EM fungi play a major role in the formation of stable N and C in SOM, which highlights the need to include mycorrhizal effects in models of global soil C stores.

Pulse-labelling trees to study carbon allocation dynamics: a review of methods, current knowledge and future prospects

Pulse-labelling of trees with stable or radioactive carbon (C) isotopes offers the unique opportunity to trace the fate of labelled CO(2) into the tree and its release to the soil and the atmosphere. Thus, pulse-labelling enables the quantification of C partitioning in forests and the assessment of the role of partitioning in tree growth, resource acquisition and C sequestration. However, this is associated with challenges as regards the choice of a tracer, the methods of tracing labelled C in tree and soil compartments and the quantitative analysis of C dynamics. Based on data from 47 studies, the rate of transfer differs between broadleaved and coniferous species and decreases as temperature and soil water content decrease. Labelled C is rapidly transferred belowground-within a few days or less-and this transfer is slowed down by drought. Half-lives of labelled C in phloem sap (transfer pool) and in mature leaves (source organs) are short, while those of sink organs (growing tissues, seasonal storage) are longer. (13)C measurements in respiratory efflux at high temporal resolution provide the best estimate of the mean residence times of C in respiratory substrate pools, and the best basis for compartmental modelling. Seasonal C dynamics and allocation patterns indicate that sink strength variations are important drivers for C fluxes. We propose a conceptual model for temperate and boreal trees, which considers the use of recently assimilated C versus stored C. We recommend best practices for designing and analysing pulse-labelling experiments, and identify several topics which we consider of prime importance for future research on C allocation in trees: (i) whole-tree C source-sink relations, (ii) C allocation to secondary metabolism, (iii) responses to environmental change, (iv) effects of seasonality versus phenology in and across biomes, and (v) carbon-nitrogen interactions. Substantial progress is expected from emerging technologies, but the largest challenge remains to carry out in situ whole-tree labelling experiments on mature trees to improve our understanding of the environmental and physiological controls on C allocation.

Effects of copper on growth and on photosynthesis of mature and expanding leaves in cucumber plants

Abstract The aim of this study was to study the relationship between growth inhibition in Cu-treated Cucumis sativus L. seedlings and photosynthesis. Twenty-two days after sowing, copper was added to the nutrient solution for 5 days, leading to final Cu supplementary concentration in sand of, respectively 0 (control) and 10 μg g −1 (Cu stress). The responses of cucumber leaves to copper addition depend on their growth stage. Young expanding leaves showed a reduction in leaf area, while mature leaves exhibited a significant decline in photosynthesis. Sucrose and starch content rose in both types of leaves. For mature leaf, net CO 2 assimilation declined at a nearly constant intercellular CO 2 mole fraction, indicating that stomatal closure did not account for the inhibition of photosynthesis. Maximal photochemical yield of PSII in dark-acclimated leaves was unaffected, indicating that the dark phase of photosynthesis rather than the light phase was affected. Relative growth rate and leaf area ratio of stressed plants were significantly below that of control plants while no significant decrease of net assimilation rate was observed for Cu-stressed plants. Thus despite a reduction in mature leaf photosynthesis, growth reduction is more likely due to a reduction in whole plant leaf area. This decline in photosynthesis is probably a consequence of an altered source–sink relationship, rather than due to a toxic effect of copper on photosynthesis.

Exploitation of northern peatlands and biodiversity maintenance: a conflict between economy and ecology.

Peatlands are ecosystems of exceptional conservation value because of their beauty, biodiversity, importance in global geochemical cycles, and the paleoenvironmental records they preserve. Commercial extraction and drainage for forestry or agriculture have caused the destruction of many peatlands, especially in or close to urban areas of the northern temperate zone. Are these commercial and environmental interests irreconcilable? A close analysis suggests that limited peat extraction may actually increase biodiversity in some cases, and may be sustainable over the long term. As we learn more about how peatlands spontaneously regenerate following disturbance, and what conditions govern the re-establishment of a diverse community and the ability to sequester carbon, we increase our chances of being able to restore damaged peatlands. Preserving the chronological records hidden in the peat profile, the natural heritage value of peatlands, and the bulk of sequestered carbon, however, will remain incompatible with any form of exploitation.

Soil CO2 efflux in a beech forest: comparison of two closed dynamic systems

The aim of this study was to understand why two closed dynamic systems with a very similar design gave large differences in soil CO2 efflux measurements (PP systems and LI-COR). Both in the field (forest beech stand) and in the laboratory, the PPsystems gave higher estimations of soil CO2 efflux than the LI-COR system (ranging from 30% to 50%). The difference in wind speed occurring within the soil respiration chambers (0.9 m s−1 within the SRC-1 and 0.4 m s−1 within the LI-6000-09 chambers) may account for the discrepancy between the two systems. An excessive air movement inside the respiration chamber is thought to disrupt the high laminar boundary layer over the forest floor. This would promote an exhaust of the CO2 accumulated into the upper soil layers into the chamber and a lateral diffusion of CO2 in the soil towards the respiration chamber. The discrepancy between the two systems was reduced (i) by decreasing fan speed within the SRC-1, (ii) by increasing wind speed over the soil surface outside the respiration chamber, or (iii) by using an artificial soil design without high CO2 concentration in soil pores. We show that wind speed is an important component of soil CO2 diffusion which must be taken into account when measuring soil CO2 efflux, even on very fine textured soil like silt-loam soil. Proper measurement can be achieved by maintaining wind speed inside the chamber below 0.4 m s−1 since low wind speed conditions predominate under forest canopies. However, more accurate measurements will be obtained by regulating wind speeds within the chamber at a velocity representative of the wind speed recorded simultaneously at the floor surface.

Climatic Influences on Seasonal and Spatial Differences in Soil CO2 Efflux

The efflux of C02 from the soil is characterized by large seasonal fluctuations due to seasonal changes in root and microbial respiration. Although several biotic and abiotic factors influence root and microbial activity (see Chap. 3), the control exerted by temperature, and in some cases moisture, is usually dominant. In the absence of water stress, variation in soil temperature accounts for most of the seasonal and diurnal variation in soil C02 efflux. Where water stress frequently occurs, soil C02 efflux may not be correlated with soil temperature, but with its moisture content (Rout and Gupta 1989). Thus, C02 release from the soil appears to respond to temperature or moisture, whichever is most limiting at the time of measurement (Schlentner and van Cleve 1985).

Tracing of recently assimilated carbon in respiration at high temporal resolution in the field with a tuneable diode laser absorption spectrometer after in situ 13CO2 pulse labelling of 20-year-old beech trees

The study of the fate of assimilated carbon in respiratory fluxes in the field is needed to resolve the residence and transfer times of carbon in the atmosphere-plant-soil system in forest ecosystems, but it requires high frequency measurements of the isotopic composition of evolved CO2. We developed a closed transparent chamber to label the whole crown of a tree and a labelling system capable of delivering a 3-h pulse of 99% 13CO2 in the field. The isotopic compositions of trunk and soil CO2 effluxes were recorded continuously on two labelled and one control trees by a tuneable diode laser absorption spectrometer during a 2-month chase period following the late summer labelling. The lag times for trunk CO2 effluxes are consistent with a phloem sap velocity of about 1 m h(-1). The isotopic composition (delta13C) of CO2 efflux from the trunk was maximal 2-3 days after labelling and declined thereafter following two exponential decays with a half-life of 2-8 days for the first and a half-life of 15-16 days for the second. The isotopic composition of the soil CO2 efflux was maximal 3-4 days after labelling and the decline was also well fitted with a sum of two exponential functions with a half-life of 3-5 days for the first exponential and a half-life of 16-18 days for the second. The amount of label recovered in CO2 efflux was around 10-15% of the assimilated 13CO2 for soil and 5-13% for trunks. As labelling occurred late in the growing season, substantial allocation to storage is expected.

In situ assessment of the velocity of carbon transfer by tracing 13C in trunk CO2 efflux after pulse labelling: variations among tree species and seasons

Phloem is the main pathway for transferring photosynthates belowground. In situ(13) C pulse labelling of trees 8-10 m tall was conducted in the field on 10 beech (Fagus sylvatica) trees, six sessile oak (Quercus petraea) trees and 10 maritime pine (Pinus pinaster) trees throughout the growing season. Respired (13) CO2 from trunks was tracked at different heights using tunable diode laser absorption spectrometry to determine time lags and the velocity of carbon transfer (V). The isotope composition of phloem extracts was measured on several occasions after labelling and used to estimate the rate constant of phloem sap outflux (kP ). Pulse labelling together with high-frequency measurement of the isotope composition of trunk CO2 efflux is a promising tool for studying phloem transport in the field. Seasonal variability in V was predicted in pine and oak by bivariate linear regressions with air temperature and soil water content. V differed among the three species consistently with known differences in phloem anatomy between broadleaf and coniferous trees. V increased with tree diameter in oak and beech, reflecting a nonlinear increase in volumetric flow with increasing bark cross-sectional area, which suggests changes in allocation pattern with tree diameter in broadleaf species. Discrepancies between V and kP indicate vertical changes in functional phloem properties.

Seasonal and daily time course of the 13C composition in soil CO2 efflux recorded with a tunable diode laser spectrophotometer (TDLS).

Temporal variations of carbon isotope composition of soil CO2 efflux (FS and δ13CFS) at different time scales should reflect both temporal variations of the climate conditions that affect canopy functioning and temporal changes in the relative contribution of autotrophic respiration to total FS. A tunable diode laser spectrophotometer (TDLS) was installed in the Hesse forest (northeast of France) early during the 2007 growing season to determine the seasonal and daily variability in δ13CFS. This method, based on the measurement of the absorption of an infrared laser emission at specific wave lengths of the 13CO2 and 12CO2, allows the continuous monitoring of the two isotopologues. The concentrations of the two isotopologues in FS were continuously monitored from June to November 2007 using chamber method and Keeling plots drawn from nocturnal accumulation of CO2 below the canopy. These TDLS measurements and isotope ratio mass spectrometer based Keeling plots gave very similar values of δ13CFS, showing the reliability of the TDLS system in this context. Results were analysed with regard to seasonal and daily changes in climatic and edaphic variables and compared with the δ13C of CO2 respired by roots, litter and soil incubated under controlled conditions. Pronounced daily as well as seasonal variations in δ13CFS were recorded (up to 1.5‰). The range of variation of δ13CFS was of the same order of magnitude at both diurnal and seasonal scales. δ13CFS observed in the field fluctuated between values of litter and of root respiration recorded during incubation, suggesting that temporal (and probably spatial) variations were associated with changes in the relative contribution of the two compartments during the day and during the season.

Production and carbon allocation in monocultures and mixed-species plantations of Eucalyptus grandis and Acacia mangium in Brazil

Introducing nitrogen-fixing tree species in fast-growing eucalypt plantations has the potential to improve soil nitrogen availability compared with eucalypt monocultures. Whether or not the changes in soil nutrient status and stand structure will lead to mixtures that out-yield monocultures depends on the balance between positive interactions and the negative effects of interspecific competition, and on their effect on carbon (C) uptake and partitioning. We used a C budget approach to quantify growth, C uptake and C partitioning in monocultures of Eucalyptus grandis (W. Hill ex Maiden) and Acacia mangium (Willd.) (treatments E100 and A100, respectively), and in a mixture at the same stocking density with the two species at a proportion of 1 : 1 (treatment MS). Allometric relationships established over the whole rotation, and measurements of soil CO(2) efflux and aboveground litterfall for ages 4-6 years after planting were used to estimate aboveground net primary production (ANPP), total belowground carbon flux (TBCF) and gross primary production (GPP). We tested the hypotheses that (i) species differences for wood production between E. grandis and A. mangium monocultures were partly explained by different C partitioning strategies, and (ii) the observed lower wood production in the mixture compared with eucalypt monoculture was mostly explained by a lower partitioning aboveground. At the end of the rotation, total aboveground biomass was lowest in A100 (10.5 kg DM m(-2)), intermediate in MS (12.2 kg DM m(-2)) and highest in E100 (13.9 kg DM m(-2)). The results did not support our first hypothesis of contrasting C partitioning strategies between E. grandis and A. mangium monocultures: the 21% lower growth (ΔB(w)) in A100 compared with E100 was almost entirely explained by a 23% lower GPP, with little or no species difference in ratios such as TBCF/GPP, ANPP/TBCF, ΔB(w)/ANPP and ΔB(w)/GPP. In contrast, the 28% lower ΔB(w) in MS than in E100 was explained both by a 15% lower GPP and by a 15% lower fraction of GPP allocated to wood growth, thus partially supporting our second hypothesis: mixing the two species led to shifts in C allocations from above- to belowground, and from growth to litter production, for both species.