Canopy CO2 enrichment permits tracing the fate of recently assimilated carbon in a mature deciduous forest

How rapidly newly assimilated carbon (C) is invested into recalcitrant structures of forests, and how closely C pools and fluxes are tied to photosynthesis, is largely unknown. A crane and a purpose-built free-air CO2 enrichment (FACE) system permitted us to label the canopy of a mature deciduous forest with 13C-depleted CO2 for 4 yr and continuously trace the flow of recent C through the forest without disturbance. Potted C4 grasses in the canopy ('isometers') served as a reference for the C-isotope input signal. After four growing seasons, leaves were completely labelled, while newly formed wood (tree rings) still contained 9% old C. Distinct labels were found in fine roots (38%) and sporocarps of mycorrhizal fungi (62%). Soil particles attached to fine roots contained 9% new C, whereas no measurable signal was detected in bulk soil. Soil-air CO2 consisted of 35% new C, indicating that considerable amounts of assimilates were rapidly returned back to the atmosphere. These data illustrate a relatively slow dilution of old mobile C pools in trees, but a pronounced allocation of very recent assimilates to C pools of short residence times.     

On the influence of tree size on the climate–growth relationship of New Zealand kauri (Agathis australis): insights from annual, monthly and daily growth patterns

Many tree-ring-based climate reconstructions are based on the assumption that the climate reaction of trees is independent of their size. Here, we test this assumption for New Zealand kauri (Agathis australis), one of the longest tree ring-based proxies for the El Niño-Southern Oscillation (ENSO). The most recent kauri chronology contains a large amount of archaeological material, e.g. timber for which the original tree size is often unknown. We analyzed the climate–growth relationship of different-sized kauri in a pristine forest using different temporal scales, i.e. annually, monthly and daily data on tree growth and climate conditions. Trees of different life stages exhibited approximately the same seasonal growth peaks during austral spring (October and November). The dormancy period overlaps with the period where weekly air temperature maxima are below ca. 17–18 °C, and where the corresponding daily minima are below ca. 8 °C. However, both correlation functions between annual growth and seasonal climate as well as Kalman filter regressions between daily growth and climate conditions suggest an influence of tree size on the climate–growth relationship for kauri. Smaller trees (DBH < 40 cm) contain weaker climate signals than larger trees. Therefore, the precautionary stripping of near-pith material (first 20 cm) from kauri chronologies may result in more uniform responses to climate forcing and thus enhance the reliability of long-term climate reconstructions.     

Tree age,site and climate controls on tree ring cellulose δ18O: A case study on oak trees from south-western France

A main concern of dendroclimatic reconstruction is to distinguish in the tree ring proxy the influence of the climate variables of interest from other controlling factors. In order to investigate age, site and climate controls on tree ring width and cellulose δ¹⁸O, measurements have been performed in nearby groups of young (145 years old) and older (310–405 years old) oak trees in south-western France, covering the period 1860–2010.Within a given site, inter-tree deviations are small, pointing to a common climatic signal. Despite a similar inter-annual variability, the average level of cellulose δ¹⁸O in the young tree group is ∼0.8‰ higher than in the old trees. Such offsets might be caused by different soil properties and differences in the fraction of the source water used by trees from different depths. The δ¹⁸O of water in the top soil layer is directly related to the current growing season precipitation, while deeper water can have a lower and more constant δ¹⁸O. Local cave drip waters at 10 m depth indeed show a constant isotopic composition, which corresponds to pluri-annual mean precipitation.A 2‰ increasing trend is observed in cellulose δ¹⁸O of young trees in the first 30 years of growth, during a period when no trend is visible in older trees. This increase can be quantitatively explained by humidity gradients under the forest canopy, and a changing microclimate around the crown as trees grow higher.While relationships between tree ring width and climate appear complex, the isotopic composition of cellulose is strongly correlated with summer maximum temperature, relative humidity and evapotranspiration (r ≈ 0.70). Weaker correlations (r ≈ 0.40) are identified with precipitation δ¹⁸O from a 15-year long local record and from the REMOiso model output. These results imply that leaf water enrichment has a stronger control on the inter-annual variability of cellulose δ¹⁸O than the δ¹⁸O of precipitation.This study demonstrates the suitability of oak tree ring cellulose δ¹⁸O for reconstructing past summer climate variability in south-western France, provided that the sampling and pooling strategy accounts for the fact that trees from different sites and of different age can introduce non-climatic signals.     

不同去趋势方法对树轮气候信号识别的影响

树木生长受到气候因子、随年龄增长的内在生长趋势、环境干扰和其他扰动信号的影响。目前存在不同的去趋势方法对树木年轮进行去趋势以识别树木生长中的气候信号。以往的研究多基于单个方法识别树轮气候信号,而不同去趋势方法识别的树轮气候信号可能会有一定的差别。为了对比不同去趋势方法对树轮气候信号识别的影响,我们基于国际年轮数据库网站获取中国西部地区68个点的树轮宽度数据,采用最常用的"signal-free"方法（SsfCrn）、线性和负指数函数法（std）、67%样条函数法（spline）、firedman方法、以及基于经验模式分解去趋势方法（EEMD）5种去趋势方法分别建立树轮年表,并对比分析同一地点的不同年表对气候响应的异同。结果表明：不同去趋势方法得到的年表对温度、降水以及相对湿度等气候因素的响应具有明显差异。其中,SsfCrn去趋势方法建立的年表对温度（月平均温、月最低温、月平均最低温）响应中相关最高的样点在所有样点中占比最高;EEMD去趋势方法建立的年表对降水量、相对湿度和月最高温响应中相关最高的样点在所有样点中占比最高;firedman去趋势方法建立的年表对月平均最高温响应中相关最高的样点在所有样点中占比最高。研究结果表明SsfCrn,EEMD和firedman方法在识别树轮气候信号方法具有一定的优势。在不同研究区域中,不同去趋势方法建立的年表对不同气候条件响应有差异,因此选择不同的去趋势方法识别树木生长趋势,分析哪种方法可以更好的反应气候变化对树木生长的影响显得尤为重要。     

Genetic by environment interactions affect plant–soil linkages

The role of plant intraspecific variation in plant–soil linkages is poorly understood, especially in the context of natural environmental variation, but has important implications in evolutionary ecology. We utilized three 18‐ to 21‐year‐old common gardens across an elevational gradient, planted with replicates of five Populus angustifolia genotypes each, to address the hypothesis that tree genotype (G), environment (E), and G × E interactions would affect soil carbon and nitrogen dynamics beneath individual trees. We found that soil nitrogen and carbon varied by over 50% and 62%, respectively, across all common garden environments. We found that plant leaf litter (but not root) traits vary by genotype and environment while soil nutrient pools demonstrated genotype, environment, and sometimes G × E interactions, while process rates (net N mineralization and net nitrification) demonstrated G × E interactions. Plasticity in tree growth and litter chemistry was significantly related to the variation in soil nutrient pools and processes across environments, reflecting tight plant–soil linkages. These data overall suggest that plant genetic variation can have differential affects on carbon storage and nitrogen cycling, with implications for understanding the role of genetic variation in plant–soil feedback as well as management plans for conservation and restoration of forest habitats with a changing climate.     

Spatial and age-dependent tree-ring growth responses of Larix gmelinii to climate in northeastern China

Tree-ring width chronologies from 276 Larix gmelinii cores taken in northeastern China were used to analyze spatial and age-dependent growth–climate response relationships. Tree radial growth from five localities showed similar patterns, while exhibiting different tree-ring growth responses to local climate. The rotated principal component analysis (RPCA) indicated that tree age, growing season moisture conditions, and ambient air temperature variations resulted from location differences (e.g., longitude, latitude, and altitude), which could explain the non-stationary spatial climate–growth relations observed. The study tested the fundamental assumption that the climate–growth of L. gmelinii was age independent after the removal of size trends and disturbance signals. The age-related climate–growth relationship might potentially improve the veracity of past climate reconstructions. Bootstrapped correlation function analyses suggested that the response of L. gmelinii radial growth to climate differed between trees ≥150 years old and <150 years old. Mean sensitivity and standard deviation for trees increased with age in the <150 years old tree class; whereas trees ≥150 years old had no significant relationship with age. These results showed that the assumption of age-independent climate–growth relationship is invalid at these sites. Physiological processes and/or hydraulic constraints dependent on tree age, together with detrending techniques could be the possible causal factors of clear age-dependent responses. These results suggested the importance of incorporating trees of all ages into the chronology to recover a detailed climatic signal in a reconstruction of L. gmelinii for this region.     

Linking drought legacy effects across scales: From leaves to tree rings to ecosystems

Severe drought can cause lagged effects on tree physiology that negatively impact forest functioning for years. These “drought legacy effects” have been widely documented in tree‐ring records and could have important implications for our understanding of broader scale forest carbon cycling. However, legacy effects in tree‐ring increments may be decoupled from ecosystem fluxes due to (a) postdrought alterations in carbon allocation patterns; (b) temporal asynchrony between radial growth and carbon uptake; and (c) dendrochronological sampling biases. In order to link legacy effects from tree rings to whole forests, we leveraged a rich dataset from a Midwestern US forest that was severely impacted by a drought in 2012. At this site, we compiled tree‐ring records, leaf‐level gas exchange, eddy flux measurements, dendrometer band data, and satellite remote sensing estimates of greenness and leaf area before, during, and after the 2012 drought. After accounting for the relative abundance of tree species in the stand, we estimate that legacy effects led to ~10% reductions in tree‐ring width increments in the year following the severe drought. Despite this stand‐scale reduction in radial growth, we found that leaf‐level photosynthesis, gross primary productivity (GPP), and vegetation greenness were not suppressed in the year following the 2012 drought. Neither temporal asynchrony between radial growth and carbon uptake nor sampling biases could explain our observations of legacy effects in tree rings but not in GPP. Instead, elevated leaf‐level photosynthesis co‐occurred with reduced leaf area in early 2013, indicating that resources may have been allocated away from radial growth in conjunction with postdrought upregulation of photosynthesis and repair of canopy damage. Collectively, our results indicate that tree‐ring legacy effects were not observed in other canopy processes, and that postdrought canopy allocation could be an important mechanism that decouples tree‐ring signals from GPP.     

The role of climate change in the widespread mortality of holm oak in open woodlands of Southwestern Spain

Forest decline and increasing tree mortality are of global concern and the identification of the causes is necessary to develop preventive measures. Global warming is an emerging factor responsible for the increasing tree mortality in drought-prone ecosystems. In the southwestern Iberian Peninsula, Mediterranean holm oak open woodlands currently undergo large-scale population-level tree die-off. In this region, temperature and aridity have increased during recent decades, but the possible role of climate change in the current oak mortality has not been investigated.To assess the role of climate change in oak die-off in managed open woodlands in southwestern Spain, we analyzed climate change-related signals in century-long tree ring chronologies of dead holm oaks. We examined the high/low-frequency variability in growth and the relationship between growth and climate.Similar to other Mediterranean forests, growth was favored by precipitation from autumn of the year prior to ring formation to spring of the year of ring formation, whereas high temperatures during spring limited growth. Since the 1970s, the intensity of the high-frequency response to water availability increased simultaneously with temperature and aridity. The growth trends matched those of climatic changes. Growth suppressions occurred during droughts in the 1970s, 1980s and 1990s. Widespread stand-level, age-independent mortality occurred since 2005 and affected trees that cannot be considered old for the species standards.The close relationship between growth and climate indicate that climate change strongly controlled the growth patterns. This suggests that harsher climatic conditions, especially increased aridity, affected the tree performance and could have played a significant role in the mortality process. Climate change may have exacerbated or predisposed trees to the impact of other factors (e.g. intense management and pathogens). These observations could suggest a similar future increase in oak mortality which may occur in more northern oak open woodlands if aridity further increases.     

Global assessment of relationships between climate and tree growth

Tree‐ring records provide global high‐resolution information on tree‐species responses to global change, forest carbon and water dynamics, and past climate variability and extremes. The underlying assumption is a stationary (time‐stable), quasi‐linear relationship between tree growth and environment, which however conflicts with basic ecological and evolutionary theory. Indeed, our global assessment of the relevant tree‐ring literature demonstrates non‐stationarity in the majority of tested cases, not limited to specific proxies, environmental parameters, regions or species. Non‐stationarity likely represents the general nature of the relationship between tree‐growth proxies and environment. Studies assuming stationarity however score two times more citations influencing other fields of science and the science–policy interface. To reconcile ecological reality with the application of tree‐ring proxies for climate or environmental estimates, we provide a clarification of the stationarity concept, propose a simple confidence framework for the re‐evaluation of existing studies and recommend the use of a new statistical tool to detect non‐stationarity in tree‐ring proxies. Our contribution is meant to stimulate and facilitate discussion in light of our results to help increase confidence in tree‐ring‐based climate and environmental estimates for science, the public and policymakers.     

Biodiversity correlates with regional patterns of forest litter pools

We compared litter pools of more than 1,000 forests differing in tree species diversity over a large scale in Catalonia (NE Spain). Monospecific forests always had smaller litter pools than mixed (from 2 to 5 tree species) forests. Whether there was a positive effect beyond two species mixtures depended on the species and functional identity of the dominant tree species. In sclerophyllous forests the positive effect of diversity was a step-function from one to more species. However, in conifers, litter pools increased constantly with tree diversity. The identity of the dominant tree species and functional type had also a significant effect on litter pools. For instance, forests dominated by sclerophyllous tree species had larger litter pools than forests dominated by deciduous and conifer tree species. When other forest structure parameters (i.e. tree basal area, wood production, successional stage, shrub cover and leaf area index) and environmental factors (i.e. mean annual temperature, mean annual precipitation, annual evapotranspiration and hillside position) where included in the analysis only leaf area index, basal area, wood production and mean temperature influenced litter pools positively. Our analysis emphasizes that at the regional scale, the litter compartment can be as influenced by biodiversity components as by other forest structure and climate components. In mixed forests, species and functional identity of the trees determine whether litter pools increase with tree diversity.     

Latitudinal Patterns and Climatic Drivers of Leaf Litter Multiple Nutrients in Chinese Broad-Leaved Tree Species: Does Leaf Habit Matter?

Leaf litter nutrients play a key role in nutrient cycling in forest ecosystems, yet our current knowledge of the ways in which climate controls leaf litter nutrients remains uncertain, especially for broad-leaved tree species in China. We performed a meta-analysis of geographic patterns of leaf litter nutrients of Chinese broad-leaved tree species in relation to climatic variables and leaf habit (as a discrete classification of tree species). We found that mean leaf litter carbon (C), nitrogen (N), phosphorus (P), potassium (K), calcium (Ca) and magnesium (Mg) were 458.36, 10.11, 0.72, 6.37, 14.22 and 2.59 mg^?1 g, respectively. Leaf litter nutrients did not diverge between leaf habits where they coexisted. These leaf litter nutrients displayed significant latitudinal trends, partly driven by climatic factors and a shift in leaf habit. Mean annual precipitation explained the largest amount of total variation in leaf litter C, N, P and K, and mean annual temperature was the most important predictor for leaf litter Mg, whereas leaf habit was the largest contributor to total variation in leaf litter Ca. We further found that the relationships between climate and leaf litter nutrients were distinguishable for evergreen and deciduous broad-leaved tree species. Collectively, our study differed from previous studies that evaluated leaf litter nutrients and only focused on N and P, and substantiated that leaf litter nutrients in forest ecosystems were affected by climate and leaf habit, but the strengths of the influences of these factors were strongly contingent on leaf litter nutrient identity. Therefore, alteration of climate would directly and indirectly (via a shift in species composition) affect latitudinal patterns of leaf litter nutrients and thus the associated nutrient flux and ecosystem functioning. Our study also underlined the need to include multiple nutrients to explore the influence of climate on leaf litter nutrient stoichiometry.     

Leaf Decay Processes during and after a Supra‐Seasonal Hydrological Drought in a Temperate Lowland Stream

Climate change models for Central Europe predict hydrological drought with fragmentation into pools during periods of high litter input in numerous lowland streams, presumably affecting in‐stream leaf decay processes. To investigate this assumption, we measured physicochemical parameters, macro‐invertebrate colonization, microbial activity, and decay rates of exposed leaves during and after a supra‐seasonal drought in a German lowland stream. Microbial activity, shredder colonization and leaf decay rates during fragmentation were low, presumably caused by drought‐related environmental conditions. Microbial activity and temperature‐corrected decay rates increased after the flow resumption but not leaf mass loss and shredder colonization. During both periods, exposed leaves appeared physically unaffected suggesting strongly reduced shredder‐mediated leaf decay despite shredder presence. Our results indicate that hydrological drought can affect organisms and processes in temperate lowland streams even after flow resumption, and should be considered in climate change scenarios. (© 2011 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Drivers of radial growth and carbon isotope discrimination of bur oak (Quercus macrocarpa Michx.) across continental gradients in precipitation,vapour pressure deficit and irradiance

Tree‐ring characteristics are commonly used to reconstruct climate variables, but divergence from the assumption of a single biophysical control may reduce the accuracy of these reconstructions. Here, we present data from bur oaks (Quercus macrocarpa Michx.) sampled within and beyond the current species bioclimatic envelope to identify the primary environmental controls on ring‐width indices (RWIs) and carbon stable isotope discrimination (Δ¹³C) in tree‐ring cellulose. Variation in Δ¹³C and RWI was more strongly related to leaf‐to‐air vapour pressure deficit (VPD) at the centre and western edge of the range compared with the northern and wettest regions. Among regions, Δ¹³C of tree‐ring cellulose was closely predicted by VPD and light responses of canopy‐level Δ¹³C estimated using a model driven by eddy flux and meteorological measurements (R² = 0.96, P = 0.003). RWI and Δ¹³C were positively correlated in the drier regions, while they were negatively correlated in the wettest region. The strength and direction of the correlations scaled with regional VPD or the ratio of precipitation to evapotranspiration. Therefore, the correlation strength between RWI and Δ¹³C may be used to infer past wetness or aridity from paleo wood by determining the degree to which carbon gain and growth have been more limited by moisture or light.     

How to Replicate the Functions and Biodiversity of a Threatened Tree Species? The Case of <Emphasis Type="Italic">Fraxinus excelsior</Emphasis> in Britain

The suitability of alternative tree species to replace species that are either threatened by pests/disease or at risk from climate change is commonly assessed by their ability to grow in a predicted future climate, their resistance to disease and their production potential. The ecological implications of a change in tree species are seldom considered. Here, we develop and test 3 methods to assess the ecological suitability of alternative trees. We use as our case study the systematic search for an alternative tree species to Fraxinus excelsior (currently declining throughout Europe due to Hymenoscyphus fraxineus). Those trees assessed as most similar to F. excelsior in selected ecosystem functions (decomposition, leaf litter and soil chemistry) (Method A) were least similar when assessed by the number of ash-associated species that also use them (Method B) and vice versa. Method C simultaneously assessed ecosystem functions and species use, allowing trade-offs between supporting ecosystem function and species use to be identified. Using Method C to develop hypothetical scenarios of different tree species mixtures showed that prioritising ecosystem function and then increasing the mixture of tree species to support the greatest number of ash-associated species possible, results in a mixture of trees more ecologically similar to F. excelsior than by simply mixing tree species together to support the greatest number of ash-associated species. We conclude that establishing alternative tree species results in changes in both ecosystem function and species supported and have developed a general method to assess suitability that simultaneously integrates both ecosystem function and the ‘number of species supported’.     

Carbon Accumulation Patterns During Post-Fire Succession in Cajander Larch (Larix cajanderi) Forests of Siberia

Increased fire activity within boreal forests could affect global terrestrial carbon (C) stocks by decreasing stand age or altering tree recruitment, leading to patterns of forest regrowth that differ from those of pre-fire stands. To improve our understanding of post-fire C accumulation patterns within boreal forests, we evaluated above- and belowground C pools within 17 Cajander larch (Larix cajanderi) stands of northeastern Siberia that varied in both years since fire and stand density. Early-successional stands (<20-year old) exhibited low larch recruitment, and consequently, low density, aboveground larch biomass, and aboveground net primary productivity (ANPP_tree). Mid-successional stands (21- to 70-year old) were even-aged with considerable variability in stand density. High-density mid-successional stands had 21 times faster rates of ANPP_tree than low-density stands (252 vs. 12?g?C?m^?2?y^?1) and 26 times more C in aboveground larch biomass (2,186 vs. 85?g?C?m^?2). Density had little effect on total soil C pools. During late-succession (>70-year old), aboveground larch biomass, ANPP_tree, and soil organic layer C pools increased with stand age. These stands were low density and multi-aged, containing both mature trees and new recruits. The rapid accumulation of aboveground larch biomass in high-density, mid-successional stands allowed them to obtain C stocks similar to those in much older low-density stands (~8,000?g?C?m^?2). If fire frequency increases without altering stand density, landscape-level C storage could decline, but if larch density also increases, large aboveground C pools within high-density stands could compensate for a shorter successional cycle.     

西大别山小林海黄山松树轮宽度的气候意义

基于树轮年代学研究方法,在鄂、豫、皖交界的西大别山北坡进行黄山松研究,建立了1915—2011年的树轮宽度标准年表（STD）.结果表明： 年表中较高的平均敏感度表明树轮中含有较多的气候变化的高频信息;较高的一阶自相关系数表明树轮生长存在显著的前期生长滞后效应;高信噪比和样本解释总量暗示树轮中含有较多的环境信息.标准年表序列指数与1959—2011年间气象因子的相关分析表明,黄山松树轮宽度生长受生长季末（9—10月）温度、降水量和相对湿度的影响较大;与9—10月帕尔默干旱指数呈显著正相关. 9、10月的水热组合是影响小林海黄山松树轮生长的主要因子.
     

Leaf litter carbon,nitrogen, and phosphorus stoichiometric patterns as related to climatic factors and leaf habits across Chinese broad-leaved tree species

Our understanding of how climate and leaf habit (evergreen vs. deciduous) drive leaf litter carbon (C), nitrogen (N), and phosphorus (P) stoichiometric patterns is largely limited but is particularly important for broad-leaved forests, since the forest is sensitive to climate change. Here, we investigated leaf litter C, N, and P stoichiometric ratios of broad-leaved tree species in relation to climate and leaf habit using previous publications and our additional samplings across China. We found that mean leaf litter C:N across Chinese broad-leaved tree species was within the range of the global flora, whereas C:P was lower and N:P higher. Evergreen species displayed higher leaf litter C:N, C:P, and N:P than their deciduous counterparts. Both leaf litter C:P and N:P for all species pooled were negatively correlated with latitude, driven by mean annual precipitation (MAP) and mean annual temperature, respectively, while leaf litter C:N displayed no clear latitudinal trend. The direction and magnitude of leaf litter C, N, and P stoichiometric ratios in response to climate diverged between leaf habits. For example, evergreen leaf litter C:N was negatively correlated with MAP, while deciduous counterparts did not respond significantly to MAP. We conclude that leaf litter C, N, and P stoichiometric ratios shifted along the climatic gradient, and the strength of such shifts differed between leaf habits. Therefore, leaf litter stoichiometric patterns across leaf habits suggest that any climate change-driven shift in species distribution may potentially alter the ecosystem’s nutrient cycling processes of evergreen- and deciduous-dominated broad-leaved forests differentially.     

Disturbance legacies and climate jointly drive tree growth and mortality in an intensively studied boreal forest

Most North American forests are at some stage of post‐disturbance regrowth, subject to a changing climate, and exhibit growth and mortality patterns that may not be closely coupled to annual environmental conditions. Distinguishing the possibly interacting effects of these processes is necessary to put short‐term studies in a longer term context, and particularly important for the carbon‐dense, fire‐prone boreal forest. The goals of this study were to combine dendrochronological sampling, inventory records, and machine‐learning algorithms to understand how tree growth and death have changed at one highly studied site (Northern Old Black Spruce, NOBS) in the central Canadian boreal forest. Over the 1999–2012 inventory period, mean tree diameter increased even as stand density and basal area declined significantly. Tree mortality averaged 1.4 ± 0.6% yr^?1, with most mortality occurring in medium‐sized trees; new recruitment was minimal. There have been at least two, and probably three, significant influxes of new trees since stand initiation, but none in recent decades. A combined tree ring chronology constructed from sampling in 2001, 2004, and 2012 showed several periods of extreme growth depression, with increased mortality lagging depressed growth by ~5 years. Higher minimum and maximum air temperatures exerted a negative influence on tree growth, while precipitation and climate moisture index had a positive effect; both current‐ and previous‐year data exerted significant effects. Models based on these variables explained 23–44% of the ring‐width variability. We suggest that past climate extremes led to significant mortality still visible in the current forest structure, with decadal dynamics superimposed on slower patterns of fire and succession. These results have significant implications for our understanding of previous work at NOBS, the carbon sequestration capability of old‐growth stands in a disturbance‐prone landscape, and the sustainable management of regional forests in a changing climate.     

Carbon isotopes in terrestrial ecosystem pools and CO2 fluxes

Stable carbon isotopes are used extensively to examine physiological, ecological, and biogeochemical processes related to ecosystem, regional, and global carbon cycles and provide information at a variety of temporal and spatial scales. Much is known about the processes that regulate the carbon isotopic composition (delta(13)C) of leaf, plant, and ecosystem carbon pools and of photosynthetic and respiratory carbon dioxide (CO(2)) fluxes. In this review, systematic patterns and mechanisms underlying variation in delta(13)C of plant and ecosystem carbon pools and fluxes are described. We examine the hypothesis that the delta(13)C of leaf biomass can be used as a reference point for other carbon pools and fluxes, which differ from the leaf in delta(13)C in a systematic fashion. Plant organs are typically enriched in (13)C relative to leaves, and most ecosystem pools and respiratory fluxes are enriched relative to sun leaves of dominant plants, with the notable exception of root respiration. Analysis of the chemical and isotopic composition of leaves and leaf respiration suggests that growth respiration has the potential to contribute substantially to the observed offset between the delta(13)C values of ecosystem respiration and the bulk leaf. We discuss the implications of systematic variations in delta(13)C of ecosystem pools and CO(2) fluxes for studies of carbon cycling within ecosystems, as well as for studies that use the delta(13)C of atmospheric CO(2) to diagnose changes in the terrestrial biosphere over annual to millennial time scales.