Regenerating temperate forest mesocosms in elevated CO2: belowground growth and nitrogen cycling

The response of temperate forest ecosystems to elevated atmospheric CO₂ concentrations is important because these ecosystems represent a significant component of the global carbon cycle. Two important but not well understood processes which elevated CO₂ may substantially alter in these systems are regeneration and nitrogen cycling. If elevated CO₂ leads to changes in species composition in regenerating forest communities then the structure and function of these ecosystems may be affected. In most temperate forests, nitrogen appears to be a limiting nutrient. If elevated CO₂ leads to reductions in nitrogen cycling through increased sequestration of nitrogen in plant biomass or reductions in mineralization rates, long-term forest productivity may be constrained. To study these processes, we established mesocosms of regenerating forest communities in controlled environments maintained at either ambient (375 ppm) or elevated (700 ppm) CO₂ concentrations. Mesocosms were constructed from intact monoliths of organic forest soil. We maintained these mesocosms for 2 years without any external inputs of nitrogen and allowed the plants naturally present as seeds and rhizomes to regenerate. We used ¹⁵N pool dilution techniques to quantify nitrogen fluxes within the mesocosms at the end of the 2 years. Elevated atmospheric CO₂ concentration significantly affected a number of plant and soil processes in the experimental regenerating forest mesocosms. These changes included increases in total plant biomass production, plant C/N ratios, ectomycorrhizal colonization of tree fine roots, changes in tree fine root architecture, and decreases in plant NH₄ ⁺ uptake rates, gross NH₄ ⁺ mineralization rates, and gross NH₄ ⁺ consumption rates. In addition, there was a shift in the relative biomass contribution of the two dominant regenerating tree species; the proportion of total biomass contributed by white birch (Betula papyrifera) decreased and the proportion of total biomass contributed by yellow birch (B. alleghaniensis) increased. However, elevated CO₂ had no significant effect on the total amount of nitrogen in plant and soil microbial biomass. In this study we observed a suite of effects due to elevated CO₂, some of which could lead to increases in potential long term growth responses to elevated CO₂, other to decreases. The reduced plant NH₄ ⁺ uptake rates we observed are consistent with reduced NH₄ ⁺ availability due to reduced gross mineralization rates. Reduced NH₄ ⁺ mineralization rates are consistent with the increases in C/N ratios we observed for leaf and fine root material. Together, these data suggest the positive increases in plant root architectural parameters and mycorrhizal colonization may not be as important as the potential negative effects of reduced nitrogen availability through decreased decomposition rates in a future atmosphere with elevated CO₂. Received: 10 January 1997 / Accepted: 25 July 1997     

Forest fine-root production and nitrogen use under elevated CO2: contrasting responses in evergreen and deciduous trees explained by a common principle

Despite the importance of nitrogen (N) limitation of forest carbon (C) sequestration at rising atmospheric CO₂ concentration, the mechanisms responsible are not well understood. To elucidate the interactive effects of elevated CO₂ (eCO₂) and soil N availability on forest productivity and C allocation, we hypothesized that (1) trees maximize fitness by allocating N and C to maximize their net growth and (2) that N uptake is controlled by soil N availability and root exploration for soil N. We tested this model using data collected in Free‐Air CO₂ Enrichment sites dominated by evergreen (Pinus taeda; Duke Forest) and deciduous [Liquidambar styraciflua; Oak Ridge National Laboratory (ORNL)] trees. The model explained 80–95% of variation in productivity and N‐uptake data among eCO₂, N fertilization and control treatments over 6 years. The model explains why fine‐root production increased, and why N uptake increased despite reduced soil N availability under eCO₂ at ORNL and Duke. In agreement with observations at other sites, the model predicts that soil N availability reduced below a critical level diminishes all eCO₂ responses. At Duke, a negative feedback between reduced soil N availability and N uptake prevented progressive reduction in soil N availability at eCO₂. At ORNL, soil N availability progressively decreased because it did not trigger reductions in N uptake; N uptake was maintained at ORNL through a large increase in the production of fast turnover fine roots. This implies that species with fast root turnover could be more prone to progressive N limitation of carbon sequestration in woody biomass than species with slow root turnover, such as evergreens. However, longer term data are necessary for a thorough evaluation of this hypothesis. The success of the model suggests that the principle of maximization of net growth to control growth and allocation could serve as a basis for simplification and generalization of larger scale forest and ecosystem models, for example by removing the need to specify parameters for relative foliage/stem/root allocation.     

A model of the long-term response of carbon allocation and productivity of forests to increased CO2concentration and nitrogen deposition

We present a simple theoretical analysis of the long term response of forest growth and carbon allocation to increased atmospheric [CO₂] and N deposition. Our analysis is based on a recent model which predicts that plant light-use efficiency increases with [CO₂] but is independent of plant N supply. We combine that model with simple assumptions for nitrogen fluxes in the soil. A quasi-equilibrium analysis of the short term tree and soil pools is then used to develop a simple graphical depiction of the long term carbon and nitrogen supply constraints on total growth, stem growth and foliar allocation. Our results suggest that long-term growth responses to [CO₂] and N deposition depend strongly on the extent to which stem allocation and foliage allocation are coupled. At one extreme (‘no coupling’), when stem allocation is fixed and independent of foliage allocation, there is no response of total growth or stem growth to increased [CO₂] unless N deposition increases. At the other extreme (‘linear coupling’), when stem allocation is proportional to foliage allocation, there is a significant long-term increase in total growth following a doubling of [CO₂], even when N deposition is unchanged, but stem growth decreases because of a long-term decrease in foliage allocation. For both types of coupling, total growth and stem growth increase with increasing N deposition. In the case of linear coupling, however, the N deposition response of stem growth is significantly larger than that of total growth, because of a long-term increase in foliage allocation. We compare our results with those obtained previously from an alternative model of canopy light-use efficiency involving a dependence on the foliar N:C ratio in addition to [CO₂]. Our results highlight the need for more experimental information on (i) the extent to which canopy light-use efficiency is independent of N supply, and (ii) the relationship between foliage allocation and stem allocation.     

Net mineralization of N at deeper soil depths as a potential mechanism for sustained forest production under elevated [CO2]

Elevated atmospheric carbon dioxide concentrations [CO₂] is projected to increase forest production, which could increase ecosystem carbon (C) storage. This study contributes to our broad goal of understanding the causes and consequences of increased fine‐root production and mortality under elevated [CO₂] by examining potential gross nitrogen (N) cycling rates throughout the soil profile. Our study was conducted in a CO₂‐enriched sweetgum (Liquidambar styraciflua L.) plantation in Oak Ridge, TN, USA. We used ¹⁵N isotope pool dilution methodology to measure potential gross N cycling rates in laboratory incubations of soil from four depth increments to 60 cm. Our objectives were twofold: (1) to determine whether N is available for root acquisition in deeper soil and (2) to determine whether elevated [CO₂], which has increased inputs of labile C resulting from greater fine‐root mortality at depth, has altered N cycling rates. Although gross N fluxes declined with soil depth, we found that N is potentially available for roots to access, especially below 15 cm depth where rates of microbial consumption of mineral N were reduced relative to production. Overall, up to 60% of potential gross N mineralization and 100% of potential net N mineralization occurred below 15 cm depth at this site. This finding was supported by in situ measurements from ion‐exchange resins, where total inorganic N availability at 55 cm depth was equal to or greater than N availability at 15 cm depth. While it is likely that trees grown under elevated [CO₂] are accessing a larger pool of inorganic N by mining deeper soil, we found no effect of elevated [CO₂] on potential gross or net N cycling rates. Thus, increased root exploration of the soil volume under elevated [CO₂] may be more important than changes in potential gross N cycling rates in sustaining forest responses to rising atmospheric CO₂.     

Optimal co-allocation of carbon and nitrogen in a forest stand at steady state

Nitrogen (N) is essential for plant production, but N uptake imposes carbon (C) costs through maintenance respiration and fine-root construction, suggesting that an optimal C:N balance can be found. Previous studies have elaborated this optimum under exponential growth; work on closed canopies has focused on foliage only. Here, the optimal co-allocation of C and N to foliage, fine roots and live wood is examined in a closed forest stand. Optimal co-allocation maximizes net primary productivity (NPP) as constrained by stand-level C and N balances and the pipe model. Photosynthesis and maintenance respiration increase with foliar nitrogen concentration ([N]), and stand-level photosynthesis and N uptake saturate at high foliage and fine-root density. Optimal NPP increases almost linearly from low to moderate N availability, saturating at high N. Where N availability is very low or very high, the system resembles a functional balance with a steady foliage [N]; in between, [N] increases with N availability. Carbon allocation to fine roots decreases, allocation to wood increases, and allocation to foliage remains stable with increasing N availability. The predicted relationships between biomass density and foliage [N] are in reasonable agreement with data from coniferous stands across Finland. All predictions agree with our qualitative understanding of N effects on growth.     

温度增高、CO2浓度升高、施氮对蒙古栎幼苗非结构碳水化合物积累及其分配的综合影响

蒙古栎(Quercus mongolica)是中国东北地区天然次生林重要组成树种, 研究该树种幼苗有机碳积累及碳库容对未来气候变化的响应, 可为预测未来气候变暖情景下蒙古栎林的天然更新及幼苗的培育提供科学参考。该文旨在探讨CO₂浓度和温度升高综合作用对蒙古栎幼苗非结构性碳水化合物(NSC)积累及其分配的影响。实验环境条件用人工气候箱控制, 控制条件如下: 1) CO₂浓度倍增(700 μmol·mol^-1), 温度升高4 ℃处理(HCHT); 2) CO₂浓度正常(400 μmol·mol^-1), 温度升高4 ℃处理(HT); 3) CO₂浓度和温度均正常, 即对照组(CK); 每个气候箱幼苗分别在3种氮素水平下生长: N2 (15 mmol·L^-1, 高氮), N1 (7.5 mmol·L^-1, 正常供氮)和N0 (不施氮), 一共为9个处理。研究结果表明, 1) HCHT共同作用对NSC积累无促进作用, 但改变了植物各器官中NSC的分配比例, 叶片中可溶性糖和淀粉的积累明显增加, HCHT下N2水平有利于NSC的积累。2) HT明显影响了蒙古栎一年生幼苗NCS的积累和分配。在N2水平下, HT明显促进NSC的积累, 并增加了在主根中的分配比例。3)植株各器官可溶性糖含量的动态变化因处理不同而异。主根淀粉含量随时间逐渐增加, 而细根淀粉含量随时间逐渐减少。在未来气候变暖的情况下, 土壤中大量的氮供给, 可能将促进蒙古栎幼苗的生长、增加其碳库容和抵御不良环境的能力, 进而提高其天然更新潜力。     

Elevated CO2 increases nitrogen fixation and decreases soil nitrogen mineralization in Florida scrub oak

We report changes in nitrogen cycling in Florida scrub oak in response to elevated atmospheric CO₂ during the first 14 months of experimental treatment. Elevated CO₂ stimulated above-ground growth, nitrogen mass, and root nodule production of the nitrogen-fixing vine, Galactia elliottii Nuttall. During this period, elevated CO₂ reduced rates of gross nitrogen mineralization in soil, and resulted in lower recovery of nitrate on resin lysimeters. Elevated CO₂ did not alter nitrogen in the soil microbial biomass, but increased the specific rate of ammonium immobilization (NH₄⁺ immobilized per unit microbial N) measured over a 24-h period. Increased carbon input to soil through greater root growth combined with a decrease in the quality of that carbon in elevated CO₂ best explains these changes. These results demonstrate that atmospheric CO₂ concentration influences both the internal cycling of nitrogen (mineralization, immobilization, and nitrification) as well as the processes that regulate total ecosystem nitrogen mass (nitrogen fixation and nitrate leaching) in Florida coastal scrub oak. If these changes in nitrogen cycling are sustained, they could cause long-term feedbacks to the growth responses of plants to elevated CO₂. Greater nitrogen fixation and reduced leaching could stimulate nitrogen-limited plant growth by increasing the mass of labile nitrogen in the ecosystem. By contrast, reduced nitrogen mineralization and increased immobilization will restrict the supply rate of plant-available nitrogen, potentially reducing plant growth. Thus, the net feedback to plant growth will depend on the balance of these effects through time.     

The effects of elevated CO2 on symbiotic N2 fixation: a link between the carbon and nitrogen cycles in grassland ecosystems

The response of plants to elevated CO₂ is dependent on the availability of nutrients, especially nitrogen. It is generally accepted that an increase in the atmospheric CO₂ concentration increases the C:N ratio of plant residues and exudates. This promotes temporary N-immobilization which might, in turn, reduce the availability of soil nitrogen. In addition, both a CO₂ stimulated increase in plant growth (thus requiring more nitrogen) and an increased N demand for the decomposition of soil residues with a large C:N will result under elevated CO₂ in a larger N-sink of the whole grassland ecosystem. One way to maintain the balance between the C and N cycles in elevated CO₂ would be to increase N-import to the grassland ecosystem through symbiotic N₂ fixation. Whether this might happen in the context of temperate ecosystems is discussed, by assessing the following hypothesis: i) symbiotic N₂ fixation in legumes will be enhanced under elevated CO₂, ii) this enhancement of N₂ fixation will result in a larger N-input to the grassland ecosystem, and iii) a larger N-input will allow the sequestration of additional carbon, either above or below-ground, into the ecosystem. Data from long-term experiments with model grassland ecosystems, consisting of monocultures or mixtures of perennial ryegrass and white clover, grown under elevated CO₂ under free-air or field-like conditions, supports the first two hypothesis, since: i) both the percentage and the amount of fixed N increases in white clover grown under elevated CO₂, ii) the contribution of fixed N to the nitrogen nutrition of the mixed grass also increases in elevated CO₂. Concerning the third hypothesis, an increased nitrogen input to the grassland ecosystem from N₂ fixation usually promotes shoot growth (above-ground C storage) in elevated CO₂. However, the consequences of this larger N input under elevated CO₂ on the below-ground carbon fluxes are not fully understood. On one hand, the positive effect of elevated CO₂ on the quantity of plant residues might be overwhelming and lead to an increased long-term below-ground C storage; on the other hand, the enhancement of the decomposition process by the N-rich legume material might favour carbon turn-over and, hence, limit the storage of below-ground carbon.     

Comparison of photosynthetic acclimation to elevated CO2 and limited nitrogen supply in soybean

Plants grown at elevated CO₂ often acclimate such that their photosynthetic capacities are reduced relative to ambient CO₂-grown plants. Reductions in synthesis of photosynthetic enzymes could result either from reduced photosynthetic gene expression or from reduced availability of nitrogen-containing substrates for enzyme synthesis. Increased carbohydrate concentrations resulting from increased photosynthetic carbon fixation at elevated CO₂ concentrations have been suggested to reduce the expression of photosynthetic genes. However, recent studies have also suggested that nitrogen uptake may be depressed by elevated CO₂, or at least that it is not increased enough to keep pace with increased carbohydrate production. This response could induce a nitrogen limitation in elevated-CO₂ plants that might account for the reduction in photosynthetic enzyme synthesis. If CO₂ acclimation were a response to limited nitrogen uptake, the effects of elevated CO₂ and limiting nitrogen supply on photosynthesis and nitrogen allocation should be similar. To test this hypothesis we grew non-nodulating soybeans at two levels each of nitrogen and CO₂ concentration and measured leaf nitrogen contents, photosynthetic capacities and Rubisco contents. Both low nitrogen and elevated CO₂ reduced nitrogen as a percentage of total leaf dry mass but only low nitrogen supply produced significant decreases in nitrogen as a percentage of leaf structural dry mass. The primary effect of elevated CO₂ was to increase non-structural carbohydrate storage rather than to decrease nitrogen content. Both low nitrogen supply and elevated CO₂ also decreased carboxylation capacity (V_cmax) and Rubisco content per unit leaf area. However, when V_cmax and Rubisco content were expressed per unit nitrogen, low nitrogen supply generally caused them to increase whereas elevated CO₂ generally caused them to decrease. Finally, elevated CO₂ significantly increased the ratio of RuBP regeneration capacity to V_cmax whereas neither nitrogen supply nor plant age had a significant effect on this parameter. We conclude that reductions in photosynthetic enzyme synthesis in elevated CO₂ appear not to result from limited nitrogen supply but instead may result from feedback inhibition by increased carbohydrate contents.     

Soil carbon and nitrogen cycling and storage throughout the soil profile in a sweetgum plantation after 11 years of CO2‐enrichment

Increased partitioning of carbon (C) to fine roots under elevated [CO₂], especially deep in the soil profile, could alter soil C and nitrogen (N) cycling in forests. After more than 11 years of free‐air CO₂ enrichment in a Liquidambar styraciflua L. (sweetgum) plantation in Oak Ridge, TN, USA, greater inputs of fine roots resulted in the incorporation of new C (i.e., C with a depleted δ¹³C) into root‐derived particulate organic matter (POM) pools to 90‐cm depth. Even though production in the sweetgum stand was limited by soil N availability, soil C and N contents were greater throughout the soil profile under elevated [CO₂] at the conclusion of the experiment. Greater C inputs from fine‐root detritus under elevated [CO₂] did not result in increased net N immobilization or C mineralization rates in long‐term laboratory incubations, possibly because microbial biomass was lower in the CO₂‐enriched plots. Furthermore, the δ¹³CO₂ of the C mineralized from the incubated soil closely tracked the δ¹³C of the labile POM pool in the elevated [CO₂] treatment, especially in shallower soil, and did not indicate significant priming of the decomposition of pre‐experiment soil organic matter (SOM). Although potential C mineralization rates were positively and linearly related to total SOM C content in the top 30 cm of soil, this relationship did not hold in deeper soil. Taken together with an increased mean residence time of C in deeper soil pools, these findings indicate that C inputs from relatively deep roots under elevated [CO₂] may increase the potential for long‐term soil C storage. However, C in deeper soil is likely to take many years to accrue to a significant fraction of total soil C given relatively smaller root inputs at depth. Expanded representation of biogeochemical cycling throughout the soil profile may improve model projections of future forest responses to rising atmospheric [CO₂].     

Combined effects of elevated CO2 and soil drought on carbon and nitrogen allocation of the desert shrub Caragana intermedia

Impacts of either elevated CO₂ or drought stress on plant growth have been studied extensively, but interactive effects of these on plant carbon and nitrogen allocation is inadequately understood yet. In this study the response of the dominant desert shrub, Caragana intermedia Kuanget H.c.Fu, to the interaction of elevated CO₂ (700 ± 20 μmol mol⁻¹) and soil drought were determined in two large environmental growth chambers (18 m²). Elevated CO₂ increased the allocation of biomass and carbon into roots and the ratio of carbon to nitrogen (C:N) as well as the leaf soluble sugar content, but decreased the allocation of biomass and carbon into leaves, leaf nitrogen and leaf soluble protein concentrations. Elevated CO₂ significantly decreased the partitioning of nitrogen into leaves, but increased that into roots, especially under soil drought. Elevated CO₂ significantly decreased the carbon isotope discrimination (Δ) in leaves, but increased them in roots, and the ratio of Δ values between root and leaf, indicating an increased allocation into below-ground parts. It is concluded that stimulation of plant growth by CO₂ enrichment may be negated under soil drought, and under the future environment, elevated CO₂ may partially offset the negative effects of enhanced drought by regulating the partitioning of carbon and nitrogen.     

Carbon-based secondary metabolites and internal nitrogen pools in Populus nigra under Free Air CO2 Enrichment (FACE) and nitrogen fertilisation

The goal of this study was to investigate whether increased nitrogen use efficiency found in Populus nigra L. (Jean Pourtet) under elevated CO₂ would correlate with changes in the production of carbon-based secondary compounds (CBSCs). Using Free-Air CO₂ Enrichment (FACE) technology, a poplar plantation was exposed to either ambient (about 370 μmol mol⁻¹ CO₂) or elevated (about 550 μmol mol⁻¹ CO₂) [CO₂] for 5 years. After three growing seasons, the plantation was coppiced and half of the experimental plots were fertilized with nitrogen. CBSCs, total nitrogen and lignin-bound nitrogen were quantified in secondary sprouts in seasons of active growth and dormancy during 2 years after coppicing. Neither elevated CO₂ nor nitrogen fertilisation alone or in combination influenced lignin concentrations in wood. Soluble phenolics and soluble proteins in wood decreased slightly in response to elevated CO₂. Higher nitrogen supply stimulated formation of CBSCs and increased protein concentrations. Lignin-bound nitrogen in wood ranged from 0.37–1.01 mg N g⁻¹ dry mass accounting for 17–26% of total nitrogen in wood, thus, forming a sizeable nitrogen fraction resistant to chemical degradation. The concentration of this nitrogen fraction was significantly decreased by elevated CO₂, increased in response to nitrogen fertilisation and showed a significant CO₂ × fertilisation interaction. Seasonal changes markedly affected the internal nitrogen pools. Soluble proteins in wood were 52–143% higher in the dormant than in the growth season. Positive correlations existed between the biosynthesis of proteins and CBSCs. The limited responses to elevated CO₂ and nitrogen fertilisation indicate that growth and defence are well orchestrated in P. nigra and that changes in the balance of both resources—nitrogen and C—have only marginal effects on wood chemistry. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Fine root chemistry and decomposition in model communities of north-temperate tree species show little response to elevated atmospheric CO<Subscript>2</Subscript> and varying soil resource availability

Rising atmospheric [CO₂] has the potential to alter soil carbon (C) cycling by increasing the content of recalcitrant constituents in plant litter, thereby decreasing rates of decomposition. Because fine root turnover constitutes a large fraction of annual NPP, changes in fine root decomposition are especially important. These responses will likely be affected by soil resource availability and the life history characteristics of the dominant tree species. We evaluated the effects of elevated atmospheric [CO₂] and soil resource availability on the production and chemistry, mycorrhizal colonization, and decomposition of fine roots in an early- and late-successional tree species that are economically and ecologically important in north temperate forests. Open-top chambers were used to expose young trembling aspen (Populus tremuloides) and sugar maple (Acer saccharum) trees to ambient (36 Pa) and elevated (56 Pa) atmospheric CO₂. Soil resource availability was composed of two treatments that bracketed the range found in the Upper Lake States, USA. After 2.5 years of growth, sugar maple had greater fine root standing crop due to relatively greater allocation to fine roots (30% of total root biomass) relative to aspen (7% total root biomass). Relative to the low soil resources treatment, aspen fine root biomass increased 76% with increased soil resource availability, but only under elevated [CO₂]. Sugar maple fine root biomass increased 26% with increased soil resource availability (relative to the low soil resources treatment), and showed little response to elevated [CO₂]. Concentrations of N and soluble phenolics, and C/N ratio in roots were similar for the two species, but aspen had slightly higher lignin and lower condensed tannins contents compared to sugar maple. As predicted by source-sink models of carbon allocation, pooled constituents (C/N ratio, soluble phenolics) increased in response to increased relative carbon availability (elevated [CO₂]/low soil resource availability), however, biosynthetically distinct compounds (lignin, starch, condensed tannins) did not always respond as predicted. We found that mycorrhizal colonization of fine roots was not strongly affected by atmospheric [CO₂] or soil resource availability, as indicated by root ergosterol contents. Overall, absolute changes in root chemical composition in response to increases in C and soil resource availability were small and had no effect on soil fungal biomass or specific rates of fine root decomposition. We conclude that root contributions to soil carbon cycling will mainly be influenced by fine root production and turnover responses to rising atmospheric [CO₂], rather than changes in substrate chemistry.     

Assessment of soil nitrogen and phosphorous availability under elevated CO2 and N-fertilization in a short rotation poplar plantation

Photosynthetic stimulation by elevated [CO₂] is largely regulated by nitrogen and phosphorus availability in the soil. During a 6 year Free Air CO₂ Enrichment (FACE) experiment with poplar trees in two short rotations, inorganic forms of soil nitrogen, extractable phosphorus, microbial and total nitrogen were assessed. Moreover, in situ and potential nitrogen mineralization, as well as enzymatic activities, were determined as measures of nutrient cycling. The aim of this study was to evaluate the effects of elevated [CO₂] and fertilization on: (1) N mineralization and immobilization processes; (2) soil nutrient availability; and (3) soil enzyme activity, as an indication of microbial and plant nutrient acquisition activity. Independent of any treatment, total soil N increased by 23% in the plantation after 6 years due to afforestation. Nitrification was the main process influencing inorganic N availability in soil, while ammonification being null or even negative. Ammonium was mostly affected by microbial immobilization and positively related to total N and microbial biomass N. Elevated [CO₂] negatively influenced nitrification under unfertilised treatment by 44% and consequently nitrate availability by 30% on average. Microbial N immobilization was stimulated by [CO₂] enrichment and probably enhanced the transformation of large amounts of N into organic forms less accessible to plants. The significant enhancement of enzyme activities under elevated [CO₂] reflected an increase in nutrient acquisition activity in the soil, as well as an increase of fungal population. Nitrogen fertilization did not influence N availability and cycling, but acted as a negative feed-back on phosphorus availability under elevated CO₂.     

Modelling forest response to increasing CO2 concentration under nutrient-limited conditions

The growth rates of woody plants depend on both the rate of photosynthetic carbon gain and the availability of essential nutrients. Instantaneous carbon gain is known to increase in response to increasing atmospheric CO₂ concentration, but it is uncertain whether this will translate into increased growth in the longer term under nutrient-limited conditions. An analytical model to address this question was developed by Comins & McMurtrie (1993, Ecological Applications 3, 666–681). Their model was further tested and analysed. Manipulation of various assumptions in the model revealed its key assumptions and allowed a more confident prediction of expected growth responses to CO₂ enrichment under nutrient-limited conditions. The analysis indicated that conclusions about the CO₂ sensitivity of production were strongly influenced by assumptions about the relationship between foliar and heartwood nitrogen concentrations. With heartwood nitrogen concentration proportional to foliar nitrogen concentration, the model predicted a strong response of plant productivity to increasing CO₂ concentration, whereas with heartwood nitrogen concentration set constant, the model predicted only a very slight growth response to changing CO₂ concentration. On the other hand, predictions were only slightly affected by: (1) assumptions about the extent of nitrogen retranslocation out of senescing roots and foliage or wood during heartwood formation; (2) the effects of nitrogen status on specific leaf area or (3) leaf longevity; (4) carbon allocation between different plant parts; or (5) changes in the N:C ratio of organic matter sequestered in the passive pool of soil organic matter. Modification of the effect of foliar nitrogen concentration on the light utilization coefficient had only a small effect on the CO₂ sensitivity for pines. However, this conclusion was strongly dependent on the chosen relationship between single-leaf photosynthesis and leaf nitrogen concentration. Overall, the analysis suggested that trees growing under nitrogen-limited conditions can respond to increasing atmospheric CO₂ concentration with considerable increases in growth.     

Sustained effects of atmospheric [CO2] and nitrogen availability on forest soil CO2 efflux

Soil CO₂ efflux (F_soil) is the largest source of carbon from forests and reflects primary productivity as well as how carbon is allocated within forest ecosystems. Through early stages of stand development, both elevated [CO₂] and availability of soil nitrogen (N; sum of mineralization, deposition, and fixation) have been shown to increase gross primary productivity, but the long‐term effects of these factors on F_soil are less clear. Expanding on previous studies at the Duke Free‐Air CO₂ Enrichment (FACE) site, we quantified the effects of elevated [CO₂] and N fertilization on F_soil using daily measurements from automated chambers over 10 years. Consistent with previous results, compared to ambient unfertilized plots, annual F_soil increased under elevated [CO₂] (ca. 17%) and decreased with N (ca. 21%). N fertilization under elevated [CO₂] reduced F_soil to values similar to untreated plots. Over the study period, base respiration rates increased with leaf productivity, but declined after productivity saturated. Despite treatment‐induced differences in aboveground biomass, soil temperature and water content were similar among treatments. Interannually, low soil water content decreased annual F_soil from potential values – estimated based on temperature alone assuming nonlimiting soil water content – by ca. 0.7% per 1.0% reduction in relative extractable water. This effect was only slightly ameliorated by elevated [CO₂]. Variability in soil N availability among plots accounted for the spatial variability in F_soil, showing a decrease of ca. 114 g C m^?2 yr^?1 per 1 g m^?2 increase in soil N availability, with consistently higher F_soil in elevated [CO₂] plots ca. 127 g C per 100 ppm [CO₂] over the +200 ppm enrichment. Altogether, reflecting increased belowground carbon partitioning in response to greater plant nutritional needs, the effects of elevated [CO₂] and N fertilization on F_soil in this stand are sustained beyond the early stages of stand development and through stabilization of annual foliage production.     

Effects of elevated CO<Subscript>2</Subscript> and nitrogen on wood structure related to water transport in seedlings of two deciduous broad-leaved tree species

Wood structure might be altered through the physiological responses to atmospheric carbon dioxide concentration ([CO₂]) and nitrogen (N) deposition. We investigated growth, water relations and wood structure of 1-year-old seedlings of two deciduous broad-leaved tree species, Quercus mongolica (oak, a ring-porous species) and Alnus hirsuta (alder, a diffuse-porous species and N₂–fixer), grown under a factorial combination of two levels of [CO₂] (36 and 72 Pa) and nitrogen supply (N; low and high) for 141 days in phytotron chambers. In oak, there was no significant effect of [CO₂] on wood structure, although elevated [CO₂] tended to decrease stomatal conductance (g _s) and increased water use efficiency regardless of the N treatment. However, high N supply increased root biomass and induced wider earlywood and larger vessels in the secondary xylem in stems, leading to increased hydraulic conductance. In alder, there was significant interactive effect of [CO₂] and N on vessel density, and high N supply increased the mean vessel area. Our results suggest that wood structures related to water transport were not markedly altered, although elevated [CO₂] induced changes in physiological parameters such as g _s and biomass allocation, and that N fertilization had more pronounced effects on non-N₂-fixing oak than on N₂-fixing alder.     

Elevated atmospheric CO2 and feedback between carbon and nitrogen cycles

We tested a conceptual model describing the influence of elevated atmospheric CO₂ on plant production, soil microorganisms, and the cycling of C and N in the plant-soil system. Our model is based on the observation that in nutrient-poor soils, plants (C₃) grown in an elevated CO₂ atmosphere often increase production and allocation to belowground structures. We predicted that greater belowground C inputs at elevated CO₂ should elicit an increase in soil microbial biomass and increased rates of organic matter turnover and nitrogen availability. We measured photosynthesis, biomass production, and C allocation of Populus grandidentata Michx. grown in nutrient-poor soil for one field season at ambient and twice-ambient (i.e., elevated) atmospheric CO₂ concentrations. Plants were grown in a sandy subsurface soil i) at ambient CO₂ with no open top chamber, ii) at ambient CO₂ in an open top chamber, and iii) at twice-ambient CO₂ in an open top chamber. Plants were fertilized with 4.5 g N m⁻² over a 47 d period midway through the growing season. Following 152 d of growth, we quantified microbial biomass and the availabilities of C and N in rhizosphere and bulk soil. We tested for a significant CO₂ effect on plant growth and soil C and N dynamics by comparing the means of the chambered ambient and chambered elevated CO₂ treatments. Rates of photosynthesis in plants grown at elevated CO₂ were significantly greater than those measured under ambient conditions. The number of roots, root length, and root length increment were also substantially greater at elevated CO₂. Total and belowground biomass were significantly greater at elevated CO₂. Under N-limited conditions, plants allocated 50–70% of their biomass to roots. Labile C in the rhizosphere of elevated-grown plants was significantly greater than that measured in the ambient treatments; there were no significant differences between labile C pools in the bulk soil of ambient and elevated-grown plants. Microbial biomass C was significantly greater in the rhizosphere and bulk soil of plants grown at elevated CO₂ compared to that in the ambient treatment. Moreover, a short-term laboratory assay of N mineralization indicated that N availability was significantly greater in the bulk soil of the elevated-grown plants. Our results suggest that elevated atmospheric CO₂ concentrations can have a positive feedback effect on soil C and N dynamics producing greater N availability. Experiments conducted for longer periods of time will be necessary to test the potential for negative feedback due to altered leaf litter chemistry. ei]{gnH}{fnLambers} ei]{gnA C}{fnBorstlap}     

Soil [N] modulates soil C cycling in CO2‐fumigated tree stands: a meta‐analysis

Under elevated atmospheric CO₂ concentrations, soil carbon (C) inputs are typically enhanced, suggesting larger soil C sequestration potential. However, soil C losses also increase and progressive nitrogen (N) limitation to plant growth may reduce the CO₂ effect on soil C inputs with time. We compiled a data set from 131 manipulation experiments, and used meta‐analysis to test the hypotheses that: (1) elevated atmospheric CO₂ stimulates soil C inputs more than C losses, resulting in increasing soil C stocks; and (2) that these responses are modulated by N. Our results confirm that elevated CO₂ induces a C allocation shift towards below‐ground biomass compartments. However, the increased soil C inputs were offset by increased heterotrophic respiration (Rh), such that soil C content was not affected by elevated CO₂. Soil N concentration strongly interacted with CO₂ fumigation: the effect of elevated CO₂ on fine root biomass and –production and on microbial activity increased with increasing soil N concentration, while the effect on soil C content decreased with increasing soil N concentration. These results suggest that both plant growth and microbial activity responses to elevated CO₂ are modulated by N availability, and that it is essential to account for soil N concentration in C cycling analyses.     

Carbon allocation in shoots of alpine treeline conifers in a CO<Subscript>2</Subscript> enriched environment

With a new approach we assessed the relative contribution of stored and current carbon compounds to new shoot growth in alpine treeline conifers. Within a free air CO₂ enrichment experiment at the alpine treeline in Switzerland, ¹³C-depleted fossil CO₂ was used to trace new carbon in the two tree species Larix decidua L. and Pinus uncinata Ramond over two subsequent years. The deciduous L. decidua was found to supply new shoot growth (structural woody part) by 46% from storage. Surprisingly, the evergreen P. uncinata, assumed to use current-year photosynthates, also utilized a considerable fraction of storage (42%) for new wood growth. In contrast, the needles of P. uncinata were built up almost completely from current-year photosynthates. The isotopic composition of different wood carbon fractions revealed a similar relative allocation of current and stored assimilates to various carbon fractions. Elevated CO₂ influenced the composition of woody tissue in a species-specific way, e.g. the water soluble fraction decreased in pine in 2001 but increased in larch in 2002 compared to ambient CO₂. Heavy defoliation applied as an additional treatment factor in the second year of the experiment decreased the lipophilic fraction in current-year wood in both species compared to undefoliated trees. We conclude that storage may play an important role for new shoot growth in these treeline conifers and that altered carbon availability (elevated CO₂, defoliation) results in significant changes in the relative amount of mobile carbon fractions in woody tissue. In particular, stored carbon seems to be of greater importance in the evergreen P. uncinata than has been previously thought.