Tree-girdling to separate root and heterotrophic respiration in two Eucalyptus stands in Brazil

The release of carbon as CO₂ from belowground processes accounts for about 70% of total ecosystem respiration. Insights about factors controlling soil CO₂ efflux are constrained by the challenge of apportioning sources of CO₂ between autotrophic tree roots (and mycorrhizal fungi) and heterotrophic microorganisms. In some temperate conifer forests, the reduction in soil CO₂ efflux after girdling (phloem removal) has been used to separate these sources. Girdling stops the flow of carbohydrates to the belowground portion of the ecosystem, which should slow respiration by roots and mycorrhizae while heterotrophic respiration should remain constant or be enhanced by the decomposition of newly dead roots. Therefore, the reduction in CO₂ efflux after girdling should be a conservative estimate of the belowground flux of C from trees. We tested this approach in two tropical Eucalyptus plantations. Tree canopies remained intact for more than 3 months after girdling, showing no reduction in light interception. The reduction in soil CO₂ efflux averaged 16–24% for the 3-month period after girdling. The reduction in CO₂ efflux was similar for plots with one half of the trees girdled and those with all of the trees girdled. Girdling did not reduce live fine root biomass for at least 5 months after treatment, indicating that large reserves of carbohydrates in the root systems of Eucalyptus trees maintained the roots and root respiration. Our results suggest that the girdling approach is unlikely to provide useful insights into the contribution of tree roots and heterotrophs to soil CO₂ efflux in this type of forest ecosystem.     

Allocation of carbon in a mature eucalypt forest and some effects of soil phosphorus availability

Pools and annual fluxes of carbon (C) were estimated for a mature Eucalyptus pauciflora (snowgum) forest with and without phosphorus (P) fertilizer addition to determine the effect of soil P availability on allocation of C in the stand. Aboveground biomass was estimated from allometric equations relating stem and branch diameters of individual trees to their biomass. Biomass production was calculated from annual increments in tree diameters and measurements of litterfall. Maintenance and construction respiration were calculated for each component using equations given by Ryan (1991a). Total belowground C flux was estimated from measurements of annual soil CO₂ efflux less the C content of annual litterfall (assuming forest floor and soil C were at approximate steady state for the year that soil CO₂ efflux was measured). The total C content of the standing biomass of the unfertilized stand was 138 t ha^-1, with approximately 80% aboveground and 20% belowground. Forest floor C was 8.5 t ha^-1. Soil C content (0–1 m) was 369 t ha^-1 representing 70% of the total C pool in the ecosystem. Total gross annual C flux aboveground (biomass increment plus litterfall plus respiration) was 11.9 t ha^-1 and gross flux belowground (coarse root increment plus fine root production plus root respiration) was 5.1 t ha^-1. Total annual soil efflux was 7.1 t ha^-1, of which 2.5 t ha^-1 (35%) was contributed by litter decomposition.The short-term effect of changing the availability of P compared with C on allocation to aboveground versus belowground processes was estimated by comparing fertilized and unfertilized stands during the year after treatment. In the P-fertilized stand annual wood biomass increment increased by 30%, there was no evidence of change in canopy biomass, and belowground C allocation decreased by 19% relative to the unfertilized stand. Total annual C flux was 16.97 and 16.75 t ha^-1 yr^-1 and the ratio of below- to aboveground C allocation was 0.43 and 0.35 in the unfertilized and P-fertilized stands, respectively. Therefore, the major response of the forest stand to increased soil P availability appeared to be a shift in C allocation; with little change in total productivity. These results emphasise that both growth rate and allocation need to be estimated to predict changes in fluxes and storage of C in forests that may occur in response to disturbance or climate change.     

Total Belowground Carbon Allocation in a Fast-growing Eucalyptus Plantation Estimated Using a Carbon Balance Approach

Trees allocate a large portion of gross primary production belowground for the production and maintenance of roots and mycorrhizae. The difficulty of directly measuring total belowground carbon allocation (TBCA) has limited our understanding of belowground carbon (C) cycling and the factors that control this important flux. We measured TBCA over 4 years using a conservation of mass, C balance approach in replicate stands of fast growing Eucalyptus saligna Smith with different nutrition management and tree density treatments. We measured TBCA as surface carbon dioxide (CO₂) efflux (“soil” respiration) minus C inputs from aboveground litter plus the change in C stored in roots, litter, and soil. We evaluated this C balance approach to measuring TBCA by examining (a) the variance in TBCA across replicate plots; (b) cumulative error associated with summing components to arrive at our estimates of TBCA; (c) potential sources of error in the techniques and assumptions; (d) the magnitude of changes in C stored in soil, litter, and roots compared to TBCA; and (e) the sensitivity of our measures of TBCA to differences in nutrient availability, tree density, and forest age. The C balance method gave precise estimates of TBCA and reflected differences in belowground allocation expected with manipulations of fertility and tree density. Across treatments, TBCA averaged 1.88 kg C m⁻² y⁻¹ and was 18% higher in plots planted with 10⁴ trees/ha compared to plots planted with 1111 trees/ha. TBCA was 12% lower (but not significantly so) in fertilized plots. For all treatments, TBCA declined linearly with stand age. The coefficient of variation (CV) for TBCA for replicate plots averaged 17%. Averaged across treatments and years, annual changes in C stored in soil, the litter layer, and coarse roots (−0.01, 0.06, and 0.21 kg C m⁻² y⁻¹, respectively) were small compared with surface CO₂ efflux (2.03 kg C m⁻² y⁻¹), aboveground litterfall (0.42 kg C m⁻² y⁻¹), and our estimated TBCA (1.88 kg C m⁻² y⁻¹). Based on studies from similar sites, estimates of losses of C through leaching, erosion, or storage of C in deep soil were less than 1% of annual TBCA. Received 6 March 2001; accepted 7 January 2002.     

Variation in below-ground carbon fluxes along a Populus hybridization gradient

Here, soil CO(2) efflux, minirhizotron fine root production (FRP), and estimated total below-ground carbon allocation (TBCA) were examined along an elevation and hybridization gradient between two cottonwood species. FRP was 72% greater under high-elevation Populus angustifolia, but soil CO(2) efflux and TBCA were 62% and 94% greater, respectively, under low-elevation stands dominated by Populus fremontii, with a hybrid stand showing intermediate values. Differences between the responses of FRP, soil CO(2) efflux and TBCA may potentially be explained in terms of genetic controls; while plant species and hybridization explained variance in carbon flux, we found only weak correlations of FRP and TBCA with soil moisture, and no correlations with soil temperature or nitrogen availability. Soil CO(2) efflux and TBCA were uncorrelated with FRP, suggesting that, although below-ground carbon fluxes may change along environmental and genetic gradients, major components of below-ground carbon flux may be decoupled.     

Belowground carbon cycling in a humid tropical forest decreases with fertilization

Only a small fraction of the carbon (C) allocated belowground by trees is retained by soils in long-lived, decay-resistant forms, yet because of the large magnitude of terrestrial primary productivity, even small changes in C allocation or retention can alter terrestrial C storage. The humid tropics exert a disproportionately large influence over terrestrial C storage, but C allocation and belowground retention in these ecosystems remain poorly quantified. Using mass balance and ¹³C isotope methods, we examined the effects of afforestation and fertilization, two land-use changes of large-scale importance, on belowground C cycling at a humid tropical site in Hawaii. Here we report that in unfertilized plots, 80% of the C allocated belowground by trees to roots and mycorrhizae was returned to the atmosphere within 1 year; 9% of the belowground C flux was retained in coarse roots and 11% was retained as new soil C. The gains in new soil C were offset entirely by losses of old soil C. Further, while fertilization early in stand development increased C storage in the litter layer and in coarse roots, it reduced by 22% the flux of C moving through roots and mycorrhizae into mineral soils. Because soil C formation rates related strongly to rhizosphere C flux, fertilization may reduce an already limited capacity of these forests to sequester decay-resistant soil C.     

Increases in the flux of carbon belowground stimulate nitrogen uptake and sustain the long-term enhancement of forest productivity under elevated CO₂

The earth's future climate state is highly dependent upon changes in terrestrial C storage in response to rising concentrations of atmospheric CO?. Here we show that consistently enhanced rates of net primary production (NPP) are sustained by a C-cascade through the root-microbe-soil system; increases in the flux of C belowground under elevated CO? stimulated microbial activity, accelerated the rate of soil organic matter decomposition and stimulated tree uptake of N bound to this SOM. This process set into motion a positive feedback maintaining greater C gain under elevated CO? as a result of increases in canopy N content and higher photosynthetic N-use efficiency. The ecosystem-level consequence of the enhanced requirement for N and the exchange of plant C for N belowground is the dominance of C storage in tree biomass but the preclusion of a large C sink in the soil.     

Physiological girdling of pine trees via phloem chilling: proof of concept

Quantifying below-ground carbon (C) allocation is particularly difficult as methods usually disturb the root-mycorrhizal-soil continuum. We reduced C allocation below ground of loblolly pine trees by: (1) physically girdling trees and (2) physiologically girdling pine trees by chilling the phloem. Chilling reduced cambium temperatures by approximately 18 degrees C. Both methods rapidly reduced soil CO2 efflux, and after approximately 10 days decreased net photosynthesis (P(n)), the latter indicating feedback inhibition. Chilling decreased soil-soluble C, indicating that decreased soil CO2 efflux may have been mediated by a decrease in root C exudation that was rapidly respired by microbes. These effects were only observed in late summer/early autumn when above-ground growth was minimal, and not in the spring when above-ground growth was rapid. All of the effects were rapidly reversed when chilling was ceased. In fertilized plots, both chilling and physical girdling methods reduced soil CO2 efflux by approximately 8%. Physical girdling reduced soil CO2 efflux by 26% in non-fertilized plots. This work demonstrates that phloem chilling provides a non-destructive alternative to reducing the movement of recent photosynthate below the point of chilling to estimate C allocation below ground on large trees.     

Effects of soil phosphorus availability,temperature and moisture on soil respiration in Eucalyptus pauciflora forest

Rates of soil respiration (CO₂ efflux) were measured for a year in a mature Eucalyptus pauciflora forest in unfertilized and phosphorus-fertilized plots. Soil CO₂ efflux showed a distinct seasonal trend, and average daily rates ranged from 124 to 574 mg CO₂ m^–2 hr^–1. Temperature and moisture are the main variables that cause variation in soil CO₂ efflux; hence their effects were investigated over a year so as to then differentiate the treatment effect of phosphorus (P) nutrition.Soil temperature had the greatest effect on CO₂ efflux and exhibited a highly significant logarithmic relationship (r² = 0.81). Periods of low soil and litter moisture occurred during summer when temperatures were greater than 10 °C, and this resulted in depression of soil CO₂ efflux. During winter, when temperatures were less than 10 °C, soil and litter moisture were consistently high and thus their variation had little effect on soil CO₂ efflux. A multiple regression model including soil temperature, and soil and litter moisture accounted for 97% of the variance in rates of CO₂ efflux, and thus can be used to predict soil CO₂ efflux at this site with high accuracy. Total annual efflux of carbon from soil was estimated to be 7.11 t C ha^–1 yr^–1. The model was used to predict changes in this annual flux if temperature and moisture conditions were altered. The extent to which coefficients of the model differ among sites and forest types requires testing.Increased soil P availability resulted in a large increase in stem growth of trees but a reduction in the rate of soil CO₂ efflux by approximately 8%. This reduction is suggested to be due to lower root activity resulting from reduced allocation of assimilate belowground. Root activity changed when P was added to microsites within plots, and via the whole tree root system at the plot level. These relationships of belowground carbon fluxes with temperature, moisture and nutrient availability provide essential information for understanding and predicting potential changes in forest ecosystems in response to land use management or climate change.     

The autotrophic contribution to soil respiration in a northern temperate deciduous forest and its response to stand disturbance

The goal of this study was to evaluate the contribution of oak trees (Quercus spp.) and their associated mycorrhizal fungi to total community soil respiration in a deciduous forest (Black Rock Forest) and to explore the partitioning of autotrophic and heterotrophic respiration. Trees on twelve 75 × 75-m plots were girdled according to four treatments: girdling all the oaks on the plot (OG), girdling half of the oak trees on a plot (O50), girdling all non-oaks on a plot (NO), and a control (C). In addition, one circular plot (diameter 50 m) was created where all trees were girdled (ALL). Soil respiration was measured before and after tree girdling. A conservative estimate of the total autotrophic contribution is approximately 50%, as indicated by results on the ALL and OG plots. Rapid declines in carbon dioxide (CO₂) flux from both the ALL and OG plots, 37 and 33%, respectively, were observed within 2 weeks following the treatment, demonstrating a fast turnover of recently fixed carbon. Responses from the NO and O50 treatments were statistically similar to the control. A non-proportional decline in respiration rates along the gradient of change in live aboveground biomass complicated partitioning of the overall rate of soil respiration and indicates that belowground carbon flux is not linearly related to aboveground disturbance. Our findings suggest that in this system there is a threshold disturbance level between 35 and 74% of live aboveground biomass loss, beyond which belowground dynamics change dramatically.     

Transpiration alters the contribution of autotrophic and heterotrophic components of soil CO2 efflux

? An unbiased partitioning of autotrophic and heterotrophic components of soil CO(2) efflux is important to estimate forest carbon budgets and soil carbon sequestration. The contribution of autotrophic sources to soil CO(2) efflux (F(A)) may be underestimated during the daytime as a result of internal transport of CO(2) produced by root respiration through the transpiration stream. ? Here, we tested the hypothesis that carbon isotope composition of soil CO(2) efflux (δ(FS)) in a Eucalyptus plantation grown on a C(4) soil is enriched during the daytime, which will indicate a decrease in F(A) during the periods of high transpiration. ? Mean δ(FS) of soil CO(2) efflux decreased to -25.7‰ during the night and increased to -24.7‰ between 11:00 and 15:00 h when the xylem sap flux density was at its maximum. ? Our results indicate a decrease in the contribution of root respiration to soil CO(2) efflux during the day that may be interpreted as a departure of root-produced CO(2) in the transpiration stream, leading to a 17% underestimation of autotrophic contribution to soil CO(2) efflux on a daily timescale.     

Genetic Identity of Populus tremuloides Litter Influences Decomposition and Nutrient Release in a Mixed Forest Stand

Recent research has shown that genetic variation can directly impact community and ecosystem level processes. Populus tremuloides (trembling aspen) is an extremely widespread and genetically diverse tree species important to many North American forest ecosystems. Using leaf litter from five genotypes grown in a common garden under two nutrient treatments, we tracked litter decomposition in a natural aspen stand for 1 year. Here we show that aspen leaf litter decomposes and releases carbon, nitrogen, and sulfur in relation to its genetic identity. In a secondary experiment, we show that the genetic diversity of aspen litter mixtures can influence decomposition, however weakly so. Overall, nutrient treatments influenced leaf litter decomposition the most, followed by genetic identity, and then by genetic diversity (if at all in some cases). In this widespread, genetically diverse, and dominant species, genetic variation within a single species is important to ecosystem functioning. The relatively weak effect of genetic diversity on the processes measured here does not preclude its importance to ecosystem functioning, but does suggest that genetic identity and composition are more important than genetic diversity per se.     

Is Tree Species Diversity or Species Identity the More Important Driver of Soil Carbon Stocks,C/N Ratio,and pH?

We explored tree species diversity effects on soil C stock, C/N ratio, and pH as compared with effects of tree species identity. We sampled forest floors and mineral soil (0–40 cm) in a diversity gradient of 1–5 tree species composed of conifers and broadleaves in Bia?owie?a Forest, Poland. Diversity was a weaker driver than identity of soil C stocks, C/N ratio, and pH in the soil profile. However, there were significant non-additive effects of diversity and significant effects of identity on C stock and C/N ratio within different parts of the soil profile. More diverse forests had higher C stocks and C/N ratios in the 20–40 cm layer, whereas identity in terms of conifer proportion increased C stocks and C/N ratios only in forest floors. A positive relationship between C stocks and root biomass in the 30–40 cm layer suggested that belowground niche complementarity could be a driving mechanism for higher root carbon input and in turn a deeper distribution of C in diverse forests. Diversity and identity affected soil pH in topsoil with positive and negative impacts, respectively. More diverse forests would lead to higher soil nutrient status as reflected by higher topsoil pH, but there was a slight negative effect on N status as indicated by higher C/N ratios in the deeper layers. We conclude that tree species diversity increases soil C stocks and nutrient status to some extent, but tree species identity is a stronger driver of the studied soil properties, particularly in the topsoil.     

Evidence of a strong coupling between root exudation, C and N availability, and stimulated SOM decomposition caused by rhizosphere priming effects

Increased temperatures and concomitant changes in vegetation patterns are expected to dramatically alter the functioning of northern ecosystems over the next few decades. Predicting the ecosystem response to such a shift in climate and vegetation is complicated by the lack of knowledge about the links between aboveground biota and belowground process rates. Current models suggest that increasing temperatures and rising concentrations of atmospheric CO(2) will be partly mitigated by elevated C sequestration in plant biomass and soil. However, empirical evidence does not always support this assumption, as elevated temperature and CO(2) concentrations also accelerate the belowground C flux, in many cases extending to increased decomposition of soil organic matter (SOM) and ultimately resulting in decreased soil C stocks. The mechanism behind the increase has remained largely unknown, but it has been suggested that priming might be the causative agent. Here, we provide quantitative evidence of a strong coupling between root exudation, SOM decomposition, and release of plant available N caused by rhizosphere priming effects. As plants tend to increase belowground C allocation with increased temperatures and CO(2) concentrations, priming effects need to be considered in our long-term analysis of soil C budgets in a changing environment. The extent of priming seems to be intimately linked to resource availability, as shifts in the stoichiometric nutrient demands of plants and microorganisms will lead to either cooperation (resulting in priming) or competition (no priming will occur). The findings lead us on the way to resolve the varying response of primary production, SOM decomposition, and release of plant available N to elevated temperatures, CO(2) concentrations, and N availability.     

Share the wealth: Trees with greater ectomycorrhizal species overlap share more carbon

The mutualistic symbiosis between forest trees and ectomycorrhizal fungi (EMF) is among the most ubiquitous and successful interactions in terrestrial ecosystems. Specific species of EMF are known to colonize specific tree species, benefitting from their carbon source, and in turn, improving their access to soil water and nutrients. EMF also form extensive mycelial networks that can link multiple root‐tips of different trees. Yet the number of tree species connected by such mycelial networks, and the traffic of material across them, are just now under study. Recently we reported substantial belowground carbon transfer between Picea, Pinus, Larix and Fagus trees in a mature forest. Here, we analyze the EMF community of these same individual trees and identify the most likely taxa responsible for the observed carbon transfer. Among the nearly 1,200 EMF root‐tips examined, 50%–70% belong to operational taxonomic units (OTUs) that were associated with three or four tree host species, and 90% of all OTUs were associated with at least two tree species. Sporocarp ¹³C signals indicated that carbon originating from labelled Picea trees was transferred among trees through EMF networks. Interestingly, phylogenetically more closely related tree species exhibited more similar EMF communities and exchanged more carbon. Our results show that belowground carbon transfer is well orchestrated by the evolution of EMFs and tree symbiosis.     

Urban Tree Effects on Soil Organic Carbon

Urban trees sequester carbon into biomass and provide many ecosystem service benefits aboveground leading to worldwide tree planting schemes. Since soils hold ∼75% of ecosystem organic carbon, understanding the effect of urban trees on soil organic carbon (SOC) and soil properties that underpin belowground ecosystem services is vital. We use an observational study to investigate effects of three important tree genera and mixed-species woodlands on soil properties (to 1 m depth) compared to adjacent urban grasslands. Aboveground biomass and belowground ecosystem service provision by urban trees are found not to be directly coupled. Indeed, SOC enhancement relative to urban grasslands is genus-specific being highest under Fraxinus excelsior and Acer spp., but similar to grasslands under Quercus robur and mixed woodland. Tree cover type does not influence soil bulk density or C∶N ratio, properties which indicate the ability of soils to provide regulating ecosystem services such as nutrient cycling and flood mitigation. The trends observed in this study suggest that genus selection is important to maximise long-term SOC storage under urban trees, but emerging threats from genus-specific pathogens must also be considered.     

Increased litterfall in tropical forests boosts the transfer of soil CO2 to the atmosphere

Aboveground litter production in forests is likely to increase as a consequence of elevated atmospheric carbon dioxide (CO(2)) concentrations, rising temperatures, and shifting rainfall patterns. As litterfall represents a major flux of carbon from vegetation to soil, changes in litter inputs are likely to have wide-reaching consequences for soil carbon dynamics. Such disturbances to the carbon balance may be particularly important in the tropics because tropical forests store almost 30% of the global soil carbon, making them a critical component of the global carbon cycle; nevertheless, the effects of increasing aboveground litter production on belowground carbon dynamics are poorly understood. We used long-term, large-scale monthly litter removal and addition treatments in a lowland tropical forest to assess the consequences of increased litterfall on belowground CO(2) production. Over the second to the fifth year of treatments, litter addition increased soil respiration more than litter removal decreased it; soil respiration was on average 20% lower in the litter removal and 43% higher in the litter addition treatment compared to the controls but litter addition did not change microbial biomass. We predicted a 9% increase in soil respiration in the litter addition plots, based on the 20% decrease in the litter removal plots and an 11% reduction due to lower fine root biomass in the litter addition plots. The 43% measured increase in soil respiration was therefore 34% higher than predicted and it is possible that this 'extra' CO(2) was a result of priming effects, i.e. stimulation of the decomposition of older soil organic matter by the addition of fresh organic matter. Our results show that increases in aboveground litter production as a result of global change have the potential to cause considerable losses of soil carbon to the atmosphere in tropical forests.     

Carbon and nitrogen dynamics in a Pinus densiflora forest with low and high stand densities

Aims Understanding carbon (C) and nitrogen (N) dynamics and their dependence on the stand density of an even-aged, mature forest provides knowledge that is important for forest management. This study investigated the differences in ecosystem total C and N storage and flux between a low-density stand (LD) and a high-density stand (HD) and examined the effects of stand density on aboveground net primary productivity (ANPP), total belowground C allocation (TBCA) and net ecosystem production (NEP) in a naturally regenerated, 65- to 75-year-old Pinus densiflora S. et Z. forest.Methods LD (450 trees ha^-1) and HD (842 trees ha^-1) were established in an even-aged, mature P. densiflora forest in September 2006. The forest had been naturally regenerated following harvesting, and the stand density was naturally maintained without any artificial management such as thinning. The diameter at breast height (DBH ≥ 5.0cm) of all live stems within the stands was measured yearly from 2007 to 2011. To compare C and N storage and fluxes in LD and HD, C and N pools in aboveground and belowground biomass, the forest floor, coarse woody debris (CWD) and soil; soil CO₂ efflux (R S); autotrophic respiration (R A); litter production; and soil N availability were measured. Further, ANPP, TBCA and NEP were estimated from plot-based measurement data.Important findings Ecosystem C (Mg C ha^-1) and N (Mg N ha^-1) storage was, respectively, 173.0±7.3 (mean ± SE) and 4.69±0.30 for LD and 162±11.8 and 4.08±0.18 for HD. There were no significant differences in C and N storage in the ecosystem components, except for soils, between the two stands. In contrast, there were significant differences in aboveground ANPP and TBCA between the two stands (P < 0.05). Litterfall, biomass increment and R S were major C flux components with values of, respectively, 3.89, 3.74 and 9.07 Mg C ha^-1 year^-1 in LD and 3.15, 2.94 and 7.06 Mg C ha^-1 year^-1 in HD. Biometric-based NEP (Mg C ha^-1 year^-1) was 4.18 in LD and 5.50 in HD. Although the even-aged, mature P. densiflora forest had similar C and N allocation patterns, it showed different C and N dynamics depending on stand density. The results of the current study will be useful for elucidating the effects of stand density on C and N storage and fluxes, which are important issues in managing natural mature forest ecosystems.     

Roots and fungi accelerate carbon and nitrogen cycling in forests exposed to elevated CO2

A common finding in multiple CO(2) enrichment experiments in forests is the lack of soil carbon (C) accumulation owing to microbial priming of 'old' soil organic matter (SOM). However, soil C losses may also result from the accelerated turnover of 'young' microbial tissues that are rich in nitrogen (N) relative to bulk SOM. We measured root-induced changes in soil C dynamics in a pine forest exposed to elevated CO(2) and N enrichment by combining stable isotope analyses, molecular characterisations of SOM and microbial assays. We find strong evidence that the accelerated turnover of root-derived C under elevated CO(2) is sufficient in magnitude to offset increased belowground inputs. In addition, the C losses were associated with accelerated N cycling, suggesting that trees exposed to elevated CO(2) not only enhance N availability by stimulating microbial decomposition of SOM via priming but also increase the rate at which N cycles through microbial pools.     

Environmental change and carbon limitation in trees: a biochemical, ecophysiological and ecosystem appraisal

As C(3) photosynthesis is not yet CO(2)-saturated, forests offer the possibility of enhanced growth and carbon (C) sequestration with rising atmospheric CO(2). However, at an ecosystem scale, increased photosynthetic rates are not always translated into faster tree growth, and in free air carbon enrichment (FACE) experiments with trees, the stimulation in above-ground growth often declines with time. So is tree growth C-limited? The evidence is reviewed here at three different scales. First, at the biochemical scale, the role of Rubisco is discussed by considering its evolution and role as a nitrogen (N) storage protein. Second, at the ecophysiological scale, C allocation to gain nutrients from the soil is considered and it is argued that any C limitation is only through a limitation to soil nutrient cycling. Finally, the response of forest ecosystems to rising atmospheric CO(2) concentrations is considered and evidence from FACE experiments is discussed. From the three lines of evidence we conclude that the growth of trees is not C-limited, with the key to understanding future responses to climate change being turnover of soil organic matter and nutrient cycling.     

Long‐term changes in forest carbon under temperature and nitrogen amendments in a temperate northern hardwood forest

Currently, forests in the northeastern United States are net sinks of atmospheric carbon. Under future climate change scenarios, the combined effects of climate change and nitrogen deposition on soil decomposition, aboveground processes, and the forest carbon balance remain unclear. We applied carbon stock, flux, and isotope data from field studies at the Harvard forest, Massachusetts, to the ForCent model, which integrates above‐ and belowground processes. The model was able to represent decadal‐scale measurements in soil C stocks, mean residence times, fluxes, and responses to a warming and N addition experiment. The calibrated model then simulated the longer term impacts of warming and N deposition on the distribution of forest carbon stocks. For simulation to 2030, soil warming resulted in a loss of soil organic matter (SOM), decreased allocation to belowground biomass, and gain of aboveground carbon, primarily in large wood, with an overall small gain in total system carbon. Simulated nitrogen addition resulted in a small increase in belowground carbon pools, but a large increase in aboveground large wood pools, resulting in a substantial increase in total system carbon. Combined warming and nitrogen addition simulations showed a net gain in total system carbon, predominately in the aboveground carbon pools, but offset somewhat by losses in SOM. Hence, the impact of continuation of anthropogenic N deposition on the hardwood forests of the northeastern United States may exceed the impact of warming in terms of total ecosystem carbon stocks. However, it should be cautioned that these simulations do not include some climate‐related processes, different responses from changing tree species composition. Despite uncertainties, this effort is among the first to use decadal‐scale observations of soil carbon dynamics and results of multifactor manipulations to calibrate a model that can project integrated aboveground and belowground responses to nitrogen and climate changes for subsequent decades.