Abstract. 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 isotopeCorrespondence to: N. Brüggemann (n.brueggemann@fz-juelich.de) 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 plantinternal 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 Published by Copernicus Publications on behalf of the European Geosciences Union. 3458 N. Brüggemann et al.: Plant-soil-atmosphere C fluxes 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 2 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 2 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 2 and the soil matrix, such as CO 2 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 2 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.

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. The last 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. 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 impede the interpretation of isotopic signals in CO₂ or organic compounds at the plant and ecosystem level. This is where new research approaches should be aimed at

We investigated the relationships of net ecosystem carbon exchange (NEE), soil temperature, and moisture with soil respiration rate and its components at a grassland ecosystem. Stable carbon isotopes were used to separate soil respiration into autotrophic and heterotrophic components within an eddy covariance footprint during the 2008 and 2009 growing seasons. After correction for self-correlation, rates of soil respiration and its autotrophic and heterotrophic components for both years were found to be strongly influenced by variations in daytime NEEthe amount of C retained in the ecosystem during the daytime, as derived from NEE measurements when photosynthetically active radiation was above 0 lmol m À2 s À1 . The time scale for correlation of variations in daytime NEE with fluctuations in respiration was longer for heterotrophic respiration (36-42 days) than for autotrophic respiration (4-6 days). In addition to daytime NEE, autotrophic respiration was also sensitive to soil moisture but not soil temperature. In contrast, heterotrophic respiration from soils was sensitive to changes in soil temperature, soil moisture, and daytime NEE. Our results show thatas for forestsplant activity is an important driver of both components of soil respiration in this tallgrass prairie grassland ecosystem. Heterotrophic respiration had a slower coupling with plant activity than did autotrophic respiration. Our findings suggest that the frequently observed variations in the sensitivity of soil respiration to temperature or moisture may stem from variations in the proportions of autotrophic and heterotrophic components of soil respiration. Rates of photosynthesis at seasonal time scales should also be considered as a driver of both autotrophic and heterotrophic soil respiration for ecosystem flux modeling. Eddy covariance measurementsContinuous measurements of CO 2 , water, and energy exchange were made with the eddy covariance technique, a micrometeorological method that measures the net exchange

While annual precipitation in much of the US Corn Belt is likely to remain constant, atmospheric vapor pressure deficit (VPD), the driver of crop water loss (evapotranspiration; ET), is projected to increase from~2.2 kPa today to~2.7 kPa by mid-century primarily due to the temperature increase. Without irrigation, it has been hypothesized that the increase in VPD will create a ceiling to future increases in maize yields. We calculated current and future growing season ET based on biomass, water use efficiency, and the amount of yield these levels of ET would support for maize production in the Midwest USA. We assumed that the production of more grain will necessitate a proportional increase in the production of biomass, with a corresponding increase in ET. Here we show that as VPD increases, maintaining current maize yields (2013-2016) will require a large expansion of irrigation, greater than threefold, in areas currently supported by rain. The average predicted yield for the region of 244 AE 4 bushels/acre (15,316 AE 251 kg/ha) projected for 2050, assuming yield increases observed for the past 60 yr continue, would not be possible with projected increases in VPD, creating a water ceiling to maize yields. Substantial increases in maize yields and the production of high yielding grasses for bioenergy will require developing cultivars with greater water use efficiency, a trait that has not been a priority for breeders in the past.

Soil respiration (Rsoil ) is one of the largest CO2 fluxes in the global carbon (C) cycle. Estimation of annual Rsoil requires extrapolation of survey measurements or gap filling of automated records to produce a complete time series. Although many gap filling methodologies have been employed, there is no standardized procedure for producing defensible estimates of annual Rsoil . Here, we test the reliability of nine different gap filling techniques by inserting artificial gaps into 20 automated Rsoil records and comparing gap filling Rsoil estimates of each technique to measured values. We show that although the most commonly used techniques do not, on average, produce large systematic biases, gap filling accuracy may be significantly improved through application of the most reliable methods. All methods performed best at lower gap fractions and had relatively high, systematic errors for simulated survey measurements. Overall, the most accurate technique estimated Rsoil based on the soil temperature dependence of Rsoil by assuming constant temperature sensitivity and linearly interpolating reference respiration (Rsoil at 10 °C) across gaps. The linear interpolation method was the second best-performing method. In contrast, estimating Rsoil based on a single annual Rsoil - Tsoil relationship, which is currently the most commonly used technique, was among the most poorly-performing methods. Thus, our analysis demonstrates that gap filling accuracy may be improved substantially without sacrificing computational simplicity. Improved and standardized techniques for estimation of annual Rsoil will be valuable for understanding the role of Rsoil in the global C cycle.

Studies on long-term effects of plants grown at elevated CO 2 are scarce and mechanisms of such responses are largely unknown. To gain mechanistic understanding on respiratory acclimation to elevated CO 2 , the Crassulacean acid metabolism Mediterranean invasive Opuntia ficus-indica Miller was grown at various CO 2 concentrations. Respiration rates, maximum activity of cytochrome c oxidase, and active mitochondrial number consistently decreased in plants grown at elevated CO 2 during the 9 months of the study when compared to ambient plants. Plant growth at elevated CO 2 also reduced cytochrome pathway activity, but increased the activity of the alternative pathway. Despite all these effects seen in plants grown at high CO 2 , the specific oxygen uptake rate per unit of active mitochondria was the same for plants grown at ambient and elevated CO 2 . Although decreases in photorespiration activity have been pointed out as a factor contributing to the long-term acclimation of plant respiration to growth at elevated CO 2 , the homeostatic maintenance of specific respiratory rate per unit of mitochondria in response to high CO 2 suggests that photorespiratory activity may play a small role on the long-term acclimation of respiration to elevated CO 2 . However, despite growth enhancement and as a result of the inhibition in cytochrome pathway activity by elevated CO 2 , total mitochondrial ATP production was decreased by plant growth at elevated CO 2 when compared to ambient-grown plants. Because plant growth at elevated CO 2 increased biomass but reduced respiratory machinery, activity, and ATP yields while maintaining O 2 consumption rates per unit of mitochondria, we suggest that acclimation to elevated CO 2 results from physiological adjustment of respiration to tissue ATP demand, which may not be entirely driven by nitrogen metabolism as previously suggested.

Perennial grasses are promising feedstocks for bioenergy production in the Midwestern USA. Few experiments have addressed how drought influences their carbon fluxes and storage. This study provides a direct comparison of ecosystem-scale measurements of carbon fluxes associated with miscanthus (Miscanthus × giganteus), switchgrass (Panicum virgatum), restored native prairie and maize (Zea mays)/soybean (Glycine max) ecosystems. The main objective of this study was to assess the influence of a naturally occurring drought during 2012 on key components of the carbon cycle and plant development relative to non-extreme years. The perennials reached full maturity 3-5 years after establishment. Miscanthus had the highest gross primary production (GPP) and lowest net ecosystem exchange (NEE) in 2012 followed by similar values for switchgrass and prairie, and the row crops had the lowest GPP and highest NEE. A post-drought effect was observed for miscanthus. Over the duration of the experiment, perennial ecosystems were carbon sinks, as indicated by negative net ecosystem carbon balance (NECB), while maize/soybean was a net carbon source. Our observations suggest that perennial ecosystems, and in particular miscanthus, can provide a high yield and a large potential for CO2 fixation even during drought, although drought may negatively influence carbon uptake in the following year, questioning the long-term consequence of its maintained productivity.

The production of perennial cellulosic feedstocks for bioenergy presents the potential to diversify regional economies and the national energy supply, while also serving as climate 'regulators' due to a number of biogeochemical and biogeophysical differences relative to row crops. Numerous observational and model-based approaches have investigated biogeochemical trade-offs, such as increased carbon sequestration and increased water use, associated with growing cellulosic feedstocks. A less understood aspect is the biogeophysical changes associated with the difference in albedo (a), which could alter the local energy balance and cause local to regional cooling several times larger than that associated with offsetting carbon. Here, we established paired fields of Miscanthus 9 giganteus (miscanthus) and Panicum virgatum (switchgrass), two of the leading perennial cellulosic feedstock candidates, and traditional annual row crops in the highly productive 'Corn-belt'. Our results show that miscanthus did and switchgrass did not have an overall higher a than current row crops, but a strong seasonal pattern existed. Both perennials had consistently higher growing season a than row crops and winter a did not differ. The lack of observed differences in winter a, however, masked an interaction between snow cover and species differences, with the perennial species, compared with the row crops, having a higher a when snow was absent and a much lower a when snow was present. Overall, these changes resulted in an average net reduction in annual absorbed energy of about 5 W m À2 for switchgrass and about 8 W m À2 for miscanthus relative to annual crops. Therefore, the conversion from annual row to perennial crops alters the radiative balance of the surface via changes in a and could lead to regional cooling.