Masako Dannoura

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.

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.

Fine root morphological traits determine variation in root respiration of Quercus serrata.

Fine root respiration is a significant component of carbon cycling in forest ecosystems. Although fine roots differ functionally from coarse roots, these root types have been distinguished based on arbitrary diameter cut-offs (e.g., 2 or 5 mm). Fine root morphology is directly related to physiological function, but few attempts have been made to understand the relationships between morphology and respiration of fine roots. To examine relationships between respiration rates and morphological traits of fine roots (0.15-1.4 mm in diameter) of mature Quercus serrata Murr., we measured respiration of small fine root segments in the field with a portable closed static chamber system. We found a significant power relationship between mean root diameter and respiration rate. Respiration rates of roots<0.4 mm in mean diameter were high and variable, ranging from 3.8 to 11.3 nmol CO2 g(-1) s(-1), compared with those of larger diameter roots (0.4-1.4 mm), which ranged from 1.8 to 3.0 nmol CO2 g(-1) s(-1). Fine root respiration rate was positively correlated with specific root length (SRL) as well as with root nitrogen (N) concentration. For roots<0.4 mm in diameter, SRL had a wider range (11.3-80.4 m g(-1)) and was more strongly correlated with respiration rate than diameter. Our results indicate that a more detailed classification of fine roots<2.0 mm is needed to represent the heterogeneity of root respiration and to evaluate root biomass and root morphological traits.

The carbon budget of coarse woody debris in a temperate broad-leaved secondary forest in Japan

We evaluated the carbon budget of coarse woody debris (CWD) in a temperate broad-leaved secondary forest. On the basis of a field survey conducted in 2003, the mass of CWD was estimated at 9.30 tC ha-1, with snags amounting to 60% of the total mass. Mean annual CWD input mass was estimated to be 0.61 tC ha-1 yr-1 by monitoring tree mortality in the forest from 1999 to 2004. We evaluated the CWD decomposition rate as the CO2 evolution rate from CWD by measuring CO2 emissions from 91 CWD samples (RCWD) with a closed dynamic chamber and infrared gas analysis system. The relationships between RCWD and temperature in the chamber, water content of the CWD, and other CWD characteristics were determined. By scaling the measured RCWD to the ecosystem, we estimated that the annual RCWD in the forest in 2003 was 0.50 tC ha-1 yr-1 or 10%–16% of the total heterotrophic respiration. Therefore, 0.11 tC ha-1 yr-1 or 7% of the forest net ecosystem production was sequestered by CWD. In a young forest, in which CWD input and decomposition are not balanced, the CWD carbon budget needs to be quantified for accurate evaluation of the forest carbon cycle and NEP.

Patterns of root respiration rates and morphological traits in 13 tree species in a tropical forest

The root systems of forest trees are composed of different diameters and heterogeneous physiological traits. However, the pattern of root respiration rates from finer and coarser roots across various tropical species remains unknown. To clarify how respiration is related to the morphological traits of roots, we evaluated specific root respiration and its relationships to mean root diameter (D) of various diameter and root tissue density (RTD; root mass per unit root volume; gcm(-3)) and specific root length (SRL; root length per unit root mass; mg(-1)) of the fine roots among and within 14 trees of 13 species from a primary tropical rainforest in the Pasoh Forest Reserve in Peninsular Malaysia. Coarse root (2-269mm) respiration rates increased with decreasing D, resulting in significant relationships between root respiration and diameter across species. A model based on a radial gradient of respiration rates of coarse roots simulated the exponential decrease in respiration with diameter. The respiration rate of fine roots (<2mm) was much higher and more variable than those of larger diameter roots. For fine roots, the mean respiration rates for each species increased with decreasing D. The respiration rates of fine roots declined markedly with increasing RTD and increased with increasing SRL, which explained a significant portion of the variation in the respiration among the 14 trees from 13 species examined. Our results indicate that coarse root respiration in tree species follows a basic relationship with D across species and that most of the variation in fine root respiration among species is explained by D, RTD and SRL. We found that the relationship between root respiration and morphological traits provides a quantitative basis for separating fine roots from coarse roots and that the pattern holds across different species.

BAAD: a biomass and allometry database for woody plants

Understanding how plants are constructed—i.e., how key size dimensions and the amount of mass invested in different tissues varies among individuals—is essential for modeling plant growth, carbon stocks, and energy fluxes in the terrestrial biosphere. Allocation patterns can differ through ontogeny, but also among coexisting species and among species adapted to different environments. While a variety of models dealing with biomass allocation exist, we lack a synthetic understanding of the underlying processes. This is partly due to the lack of suitable data sets for validating and parameterizing models. To that end, we present the Biomass And Allometry Database (BAAD) for woody plants. The BAAD contains 259 634 measurements collected in 176 different studies, from 21 084 individuals across 678 species. Most of these data come from existing publications. However, raw data were rarely made public at the time of publication. Thus, the BAAD contains data from different studies, transformed into standard units and variable names. The transformations were achieved using a common workflow for all raw data files. Other features that distinguish the BAAD are: (i) measurements were for individual plants rather than stand averages; (ii) individuals spanning a range of sizes were measured; (iii) plants from 0.01–100 m in height were included; and (iv) biomass was estimated directly, i.e., not indirectly via allometric equations (except in very large trees where biomass was estimated from detailed sub-sampling). We included both wild and artificially grown plants. The data set contains the following size metrics: total leaf area; area of stem cross-section including sapwood, heartwood, and bark; height of plant and crown base, crown area, and surface area; and the dry mass of leaf, stem, branches, sapwood, heartwood, bark, coarse roots, and fine root tissues. We also report other properties of individuals (age, leaf size, leaf mass per area, wood density, nitrogen content of leaves and wood), as well as information about the growing environment (location, light, experimental treatment, vegetation type) where available. It is our hope that making these data available will improve our ability to understand plant growth, ecosystem dynamics, and carbon cycling in the worlds vegetation.

Detection of Cryptomeria japonica roots with ground penetrating radar

Abstract Coarse tree roots, which are responsible for most root carbon storage, are usually measured by destructive methods such as excavation and coring. Ground penetrating radar (GPR) is a non-destructive tool that could be used to detect coarse roots in forest soils. In this study, we examined whether the roots of Cryptomeria japonica, a major plantation species in Japan, can be detected with GPR. We also looked for factors that impact the analysis and detection of roots. Roots and wooden dowels of C. japonica were buried 30 cm deep in sandy granite soil. From GPR measurements with a 900 MHz antenna, the distribution and diameter of samples in several transects were recorded. The buried roots were detected clearly and could be distinguished at diameters of 1.1–5.2 cm. There were significant positive relationships between root diameter and parameters extracted from the resultant GPR waveform. The difference in water content between roots and soil is a crucial factor impacting the ability to detect roots with GPR. We conclude that GPR can be used as a non-destructive tool, but further investigation is needed to determine optimal conditions (e.g. water content) and analytical methods for using GPR to examine roots in forest sites.

Development of an automatic chamber system for long-term measurements of CO2 flux from roots

To separate CO2 efflux from roots (Rr) and soil (Rs), we developed a system to measure Rr continuously. Using this system, seasonal variation in Rr was obtained in a temperate forest in Japan. We measured Rs, CO2 efflux from mineral soil (Rm) and environmental factors simultaneously, and the characteristic and seasonality of Rr were analysed in comparison with Rs. Rr and Rs showed different responses to soil water content: Rs decreased with decreasing soil water content, whereas Rr peaked at relatively low soil water content. Rr/Rs decreased from 64.8% to 27.3% as soil water content increased from 0.075 to 0.225 cm cm-3. The relationship between respiration and temperature appears to change seasonally in response to phenological and biological factors. Rr showed clear seasonal variation as a function of soil temperature. During the growing period, Rr exhibited a higher rate at the same soil temperature than during other periods, which may be due to phenological influences such as fine root dynamics. Rs decreased during the summer despite high soil temperatures. The seasonal peak for Rs occurred earlier than that for soil temperature. Rr/Rs ranged between 25% and 60% over the course of the year.

Components and seasonal variation of night‐time total ecosystem respiration in a Japanese broad‐leaved secondary forest

The Yamashiro Experimental Forest is a broad-leaved secondary forest in Kyoto, Japan. On its complex terrain, low wind speed, high air stability, and local advection are common at night. To reduce the uncertainty in measuring woodytissue respiration at night, we used automated stem chambers to measure stem respiration continuously (for 5 min at 30 min intervals) on stems of Quercus serrata Murr. (deciduous) and Ilex pedunculosa Miq. (evergreen) throughout 2003. Using these data, we estimated night-time respiration for the total ecosystem and its various components, and we report foliar and soil respiration rates for 2003. Annual average night-time respiration of soil, evergreen leaf, deciduous leaf, evergreen woody tissue and deciduous woody tissue were estimated as 0.0794 (63.2%), 0.0101 (8.0%), 0.0160 (12.7%), 0.0064 (5.1%) and 0.0137 (10.9%) mg CO2 m-2 s-1, respectively. The contribution of soil respiration to the total ecosystem respiration rate reached its minimum (49.1%) on 12 June (DOY 163) and its maximum (82.4%) on 29 November (DOY 333). Seasonal change of growth respiration was marked, indicating that the seasonal variation of growth respiration must be evaluated carefully to estimate total ecosystem respiration. Therefore, long-term continuous measurement using automated chambers and averaging provides an effective means of evaluating the annual night-time ecosystem respiration.