Influence of Nutrient Availability, Stand Age, and Canopy Structure on Isoprene Flux in a Eucalyptus saligna Experimental Forest

Eucalyptus plantations occupy approximately 10 million ha of land in the tropics and, increasingly, afforestation and reforestation projects are relying on this genus to provide rapid occupation of degraded sites, large quantities of high-quality wood products, and high rates of carbon sequestration. Members of the genus Eucalyptus are also very high emitters of isoprene, the dominant volatile organic compound emitted by trees in tropical ecosystems, which significantly influences the oxidative capacity of the atmosphere. While fertilization growth response of these trees has been intensively studied, little is known about how fertilization and tree age alter isoprene production from plantations of these trees. Here we examined the effects of fertilization and tree age on leaf-level isoprene flux from 2- and 6-year-old trees in a Eucalyptus saligna experimental forest in Hawaii. Leaf-level emission at a given canopy height did not differ between fertilized and unfertilized 6-year-old trees likely because leaf nitrogen content did not vary with fertilization. Across treatments, however, the standardized emission rate of isoprene (emission at a standard light and temperature) followed patterns of leaf N content and declined with canopy depth. Although leaf nitrogen content was similar between 2-year and 6-year fertilized trees, leaf-level emission rates declined with stand age. Surprisingly, despite differences in stand leaf area and leaf area distribution, modeled canopy-level isoprene flux was similar across stands varying in fertilization and tree age. Model results suggest that leaf area index was high enough in all treatments to absorb most of the light penetrating the canopy, leading to similar canopy flux rates despite the very different sized canopies

Carbon allocation and carbon isotope fluxes in the plant-soil-atmosphere continuum: a review

The terrestrial carbon (C) cycle has received increasing interest over the past few decades, however, there is still a lack of understanding of the fate of newly assimilated C allocated within plants and to the soil, stored within ecosystems and lost to the atmosphere. Stable carbon isotope studies can give novel insights into these issues. In this review we provide an overview of an emerging picture of plant-soil-atmosphere C fluxes, as based on C isotope studies, and identify processes determining related C isotope signatures. The first part of the review focuses on isotopic fractionation processes within plants during and after photosynthesis. The second major part elaborates on plant-internal and plant-rhizosphere C allocation patterns at different time scales (diel, seasonal, interannual), including the speed of C transfer and time lags in the coupling of assimilation and respiration, as well as the magnitude and controls of plant-soil C allocation and respiratory fluxes. Plant responses to changing environmental conditions, the functional relationship between the physiological and phenological status of plants and C transfer, and interactions between C, water and nutrient dynamics are discussed. The role of the C counterflow from the rhizosphere to the aboveground parts of the plants, e.g. via CO<sub>2</sub> dissolved in the xylem water or as xylem-transported sugars, is highlighted. The third part is centered around belowground C turnover, focusing especially on above- and belowground litter inputs, soil organic matter formation and turnover, production and loss of dissolved organic C, soil respiration and CO<sub>2</sub> fixation by soil microbes. Furthermore, plant controls on microbial communities and activity via exudates and litter production as well as microbial community effects on C mineralization are reviewed. A further part of the paper is dedicated to physical interactions between soil CO<sub>2</sub> and the soil matrix, such as CO<sub>2</sub> diffusion and dissolution processes within the soil profile. Finally, we highlight state-of-the-art stable isotope methodologies and their latest developments. From the presented evidence we conclude that there exists a tight coupling of physical, chemical and biological processes involved in C cycling and C isotope fluxes in the plant-soil-atmosphere system. Generally, research using information from C isotopes allows an integrated view of the different processes involved. However, complex interactions among the range of processes complicate or currently impede the interpretation of isotopic signals in CO<sub>2</sub> or organic compounds at the plant and ecosystem level. This review tries to identify present knowledge gaps in correctly interpreting carbon stable isotope signals in the plant-soil-atmosphere system and how future research approaches could contribute to closing these gaps

Response of CO₂ and H₂O fluxes in a mountainous tropical rainforest in equatorial Indonesia to El Niño events

The possible impact of El Niño–Southern Oscillation (ENSO) events on the main components of CO₂ and H₂O fluxes between the tropical rainforest and the atmosphere is investigated. The fluxes were continuously measured in an old-growth mountainous tropical rainforest in Central Sulawesi in Indonesia using the eddy covariance method for the period from January 2004 to June 2008. During this period, two episodes of El Niño and one episode of La Niña were observed. All these ENSO episodes had moderate intensity and were of the central Pacific type. The temporal variability analysis of the main meteorological parameters and components of CO₂ and H₂O exchange showed a high sensitivity of evapotranspiration (ET) and gross primary production (GPP) of the tropical rainforest to meteorological variations caused by both El Niño and La Niña episodes. Incoming solar radiation is the main governing factor that is responsible for ET and GPP variability. Ecosystem respiration (RE) dynamics depend mainly on the air temperature changes and are almost insensitive to ENSO. Changes in precipitation due to moderate ENSO events did not have any notable effect on ET and GPP, mainly because of sufficient soil moisture conditions even in periods of an anomalous reduction in precipitation in the region

The impact of extreme summer drought on the short-term carbon coupling of photosynthesis to soil CO<sub>2</sub> efflux in a temperate grassland

Along with predicted climate change, increased risks for summer drought are projected for Central Europe. However, large knowledge gaps exist in terms of how drought events influence the short-term ecosystem carbon cycle. Here, we present results from 13CO2 pulse labeling experiments at an intensively managed lowland grassland in Switzerland. We investigated the effect of extreme summer drought on the short-term coupling of freshly assimilated photosynthates in shoots to roots as well as to soil CO2 efflux. Summer drought was simulated using rainout shelters during two field seasons (2010 and 2011). Soil CO2 efflux and its isotopic composition were measured with custom-built chambers coupled to a quantum cascade laser spectrometer (QCLAS-ISO, Aerodyne Research Inc., MA, USA). During the 90 min pulse labeling experiments, we added 99.9 atom % 13CO2 to the grass sward. In addition to the isotopic analysis of soil CO2 efflux, this label was traced over 31 days into bulk shoots, roots and soil. Drought reduced the incorporation of recently fixed carbon into the shoots, but increased the relative allocation of fresh assimilates below ground compared to the control grasslands. Contrary to our hypothesis, we did not find a change of allocation speed in response to drought. Although drought clearly reduced soil CO2 efflux rates, about 75% of total tracer uptake in control plots was lost via soil CO2 efflux during 19 days after pulse labeling, compared to only about 60% under drought conditions. Thus, the short-term coupling of above- and below-ground processes was reduced in response to summer drought. The occurrence of a natural spring drought in 2011 lead to comparable albeit weaker drought responses increasing the confidence in the generalizability of our findings

The impact of extreme summer drought on the short-term carbon coupling of photosynthesis to soil CO2 efflux in a temperate grassland

Abstract. Along with predicted climate change, increased risks for summer drought are projected for Central Europe. However, large knowledge gaps exist in terms of how drought events influence the short-term ecosystem carbon cycle. Here, we present results from 13 CO 2 pulse labeling experiments at an intensively managed lowland grassland in Switzerland. We investigated the effect of extreme summer drought on the short-term coupling of freshly assimilated photosynthates in shoots to roots as well as to soil CO 2 efflux. Summer drought was simulated using rainout shelters during two field seasons (2010 and 2011). Soil CO 2 efflux and its isotopic composition were measured with custombuilt chambers coupled to a quantum cascade laser spectrometer (QCLAS-ISO, Aerodyne Research Inc., MA, USA). During the 90 min pulse labeling experiments, we added 99.9 atom % 13 CO 2 to the grass sward. In addition to the isotopic analysis of soil CO 2 efflux, this label was traced over 31 days into bulk shoots, roots and soil. Drought reduced the incorporation of recently fixed carbon into the shoots, but increased the relative allocation of fresh assimilates below ground compared to the control grasslands. Contrary to our hypothesis, we did not find a change of allocation speed in response to drought. Although drought clearly reduced soil CO 2 efflux rates, about 75 % of total tracer uptake in control plots was lost via soil CO 2 efflux during 19 days after pulse labeling, compared to only about 60 % under drought conditions. Thus, the short-term coupling of above-and below-ground processes was reduced in response to summer drought. The occurrence of a natural spring drought in 2011 lead to comparable albeit weaker drought responses increasing the confidence in the generalizability of our findings

Carbon allocation and carbon isotope fluxes in the plant-soil-atmosphere continuum: A review

The terrestrial carbon (C) cycle has received increasing interest over the past few decades, however, there is still a lack of understanding of the fate of newly assimilated C allocated within plants and to the soil, stored within ecosystems and lost to the atmosphere. Stable carbon isotope studies can give novel insights into these issues. In this review we provide an overview of an emerging picture of plant-soil-atmosphere C fluxes, as based on C isotope studies, and identify processes determining related C isotope signatures. The first part of the review focuses on isotopic fractionation processes within plants during and after photosynthesis. The second major part elaborates on plant-internal and plant-rhizosphere C allocation patterns at different time scales (diel, seasonal, interannual), including the speed of C transfer and time lags in the coupling of assimilation and respiration, as well as the magnitude and controls of plant-soil C allocation and respiratory fluxes. Plant responses to changing environmental conditions, the functional relationship between the physiological and phenological status of plants and C transfer, and interactions between C, water and nutrient dynamics are discussed. The role of the C counterflow from the rhizosphere to the aboveground parts of the plants, e.g. via CO2 dissolved in the xylem water or as xylem-transported sugars, is highlighted. The third part is centered around belowground C turnover, focusing especially on above- and belowground litter inputs, soil organic matter formation and turnover, production and loss of dissolved organic C, soil respiration and CO2 fixation by soil microbes. Furthermore, plant controls on microbial communities and activity via exudates and litter production as well as microbial community effects on C mineralization are reviewed. A further part of the paper is dedicated to physical interactions between soil CO2 and the soil matrix, such as CO2 diffusion and dissolution processes within the soil profile. Finally, we highlight state-of-the-art stable isotope methodologies and their latest developments. From the presented evidence we conclude that there exists a tight coupling of physical, chemical and biological processes involved in C cycling and C isotope fluxes in the plant-soil-atmosphere system. Generally, research using information from C isotopes allows an integrated view of the different processes involved. However, complex interactions among the range of processes complicate or currently impede the interpretation of isotopic signals in CO2 or organic compounds at the plant and ecosystem level. This review tries to identify present knowledge gaps in correctly interpreting carbon stable isotope signals in the plant-soil-atmosphere system and how future research approaches could contribute to closing these gaps

Atmospheric deposition, CO2, and change in the land carbon sink

Concentrations of atmospheric carbon dioxide (CO2) have continued to increase whereas atmospheric deposition of sulphur and nitrogen has declined in Europe and the USA during recent decades. Using time series of flux observations from 23 forests distributed throughout Europe and the USA, and generalised mixed models, we found that forest-level net ecosystem production and gross primary production have increased by 1% annually from 1995 to 2011. Statistical models indicated that increasing atmospheric CO2 was the most important factor driving the increasing strength of carbon sinks in these forests. We also found that the reduction of sulphur deposition in Europe and the USA lead to higher recovery in ecosystem respiration than in gross primary production, thus limiting the increase of carbon sequestration. By contrast, trends in climate and nitrogen deposition did not significantly contribute to changing carbon fluxes during the studied period. Our findings support the hypothesis of a general CO2-fertilization effect on vegetation growth and suggest that, so far unknown, sulphur deposition plays a significant role in the carbon balance of forests in industrialized regions. Our results show the need to include the effects of changing atmospheric composition, beyond CO2, to assess future dynamics of carbon-climate feedbacks not currently considered in earth system/climate modelling.Peer reviewe

Biogenic Volatile Organic Compound and Respiratory CO2 Emissions after 13C-Labeling: Online Tracing of C Translocation Dynamics in Poplar Plants

Globally plants are the primary sink of atmospheric CO(2), but are also the major contributor of a large spectrum of atmospheric reactive hydrocarbons such as terpenes (e.g. isoprene) and other biogenic volatile organic compounds (BVOC). The prediction of plant carbon (C) uptake and atmospheric oxidation capacity are crucial to define the trajectory and consequences of global environmental changes. To achieve this, the biosynthesis of BVOC and the dynamics of C allocation and translocation in both plants and ecosystems are important.We combined tunable diode laser absorption spectrometry (TDLAS) and proton transfer reaction mass spectrometry (PTR-MS) for studying isoprene biosynthesis and following C fluxes within grey poplar (Populus x canescens) saplings. This was achieved by feeding either (13)CO(2) to leaves or (13)C-glucose to shoots via xylem uptake. The translocation of (13)CO(2) from the source to other plant parts could be traced by (13)C-labeled isoprene and respiratory (13)CO(2) emission.In intact plants, assimilated (13)CO(2) was rapidly translocated via the phloem to the roots within 1 hour, with an average phloem transport velocity of 20.3±2.5 cm h(-1). (13)C label was stored in the roots and partially reallocated to the plants' apical part one day after labeling, particularly in the absence of photosynthesis. The daily C loss as BVOC ranged between 1.6% in mature leaves and 7.0% in young leaves. Non-isoprene BVOC accounted under light conditions for half of the BVOC C loss in young leaves and one-third in mature leaves. The C loss as isoprene originated mainly (76-78%) from recently fixed CO(2), to a minor extent from xylem-transported sugars (7-11%) and from photosynthetic intermediates with slower turnover rates (8-11%).We quantified the plants' C loss as respiratory CO(2) and BVOC emissions, allowing in tandem with metabolic analysis to deepen our understanding of ecosystem C flux

Quality control of CarboEurope flux data – Part I: Footprint analyses to evaluate sites in forest ecosystems

International audienceWe applied a site evaluation approach combining Lagrangian Stochastic footprint modelling with a quality assessment approach for eddy-covariance data to 25 forested sites of the CarboEurope-IP network. The analysis addresses the spatial representativeness of the flux measurements, instrumental effects on data quality, spatial patterns in the data quality, and the performance of the coordinate rotation method. Our findings demonstrate that application of a footprint filter could strengthen the CarboEurope-IP flux database, since only one third of the sites is situated in truly homogeneous terrain. Almost half of the sites experience a significant reduction in eddy-covariance data quality under certain conditions, though these effects are mostly constricted to a small portion of the dataset. Reductions in data quality of the sensible heat flux are mostly induced by characteristics of the surrounding terrain, while the latent heat flux is subject to instrumentation-related problems. The Planar-Fit coordinate rotation proved to be a reliable tool for the majority of the sites using only a single set of rotation angles. Overall, we found a high average data quality for the CarboEurope-IP network, with good representativeness of the measurement data for the specified target land cover types

Quality Control of CarboEurope Flux Data - Part I: Coupling Footprint Analyses with Flux Data Quality Assessment to Evaluate Sites in Forest Ecosystems

We applied a site evaluation approach combining Lagrangian Stochastic footprint modelling with a quality assessment approach for eddy-covariance data to 25 forested sites of the CarboEurope-IP network. The analysis addresses the spatial representativeness 5 of the flux measurements, instrumental effects on data quality, spatial patterns in the data quality, and the performance of the coordinate rotation method. Our findings demonstrate that application of a footprint filter could strengthen the CarboEurope-IP flux database, since only one third of the sites is situated in truly homogeneous terrain. Almost half of the sites experience a significant reduction in eddy-covariance data 10 quality under certain conditions, though these effects are mostly constricted to a small portion of the dataset. Reductions in data quality of the sensible heat flux are mostly induced by characteristics of the surrounding terrain, while the latent heat flux is subject to instrumentation-related problems. The Planar-Fit coordinate rotation proved to be a reliable tool for the majority of the sites using only a single set of rotation angles. 15 Overall, we found a high average data quality for the CarboEurope-IP network, with good representativeness of the measurement data for the specified target land cover types.JRC.H.2-Air and Climat