Forest biomass,productivity and carbon cycling along a rainfall gradient in West Africa

Net Primary Productivity (NPP) is one of the most important parameters in describing the functioning of any ecosystem and yet it arguably remains a poorly quantified and understood component of carbon cycling in tropical forests, especially outside of the Americas. We provide the first comprehensive analysis of NPP and its carbon allocation to woody, canopy and root growth components at contrasting lowland West African forests spanning a rainfall gradient. Using a standardized methodology to study evergreen (EF), semi‐deciduous (SDF), dry forests (DF) and woody savanna (WS), we find that (i) climate is more closely related with above and belowground C stocks than with NPP (ii) total NPP is highest in the SDF site, then the EF followed by the DF and WS and that (iii) different forest types have distinct carbon allocation patterns whereby SDF allocate in excess of 50% to canopy production and the DF and WS sites allocate 40%–50% to woody production. Furthermore, we find that (iv) compared with canopy and root growth rates the woody growth rate of these forests is a poor proxy for their overall productivity and that (v) residence time is the primary driver in the productivity‐allocation‐turnover chain for the observed spatial differences in woody, leaf and root biomass across the rainfall gradient. Through a systematic assessment of forest productivity we demonstrate the importance of directly measuring the main components of above and belowground NPP and encourage the establishment of more permanent carbon intensive monitoring plots across the tropics.     

Forest Productivity and Efficiency of Resource Use Across a Chronosequence of Tropical Montane Soils

We tested the hypothesis that plants adjust to nutrient availability by altering carbon allocation patterns and nutrient-use efficiency (NUE = net primary production [NPP] per unit nutrient uptake), but are constrained by a trade-off between NUE and light-use efficiency () = NPP per unit intercepted light). NPP, NUE and ) were measured in montane Metrosideros polymorpha forest across a 4.1 x 10⁶ yr space for time substitution chronosequence in which available soil N and P pools change with site age. Although the range of N and P availability across sites was broad, there was little difference in NPP between sites, and in contrast to theories of carbon allocation relative to limiting resources, we found no consistent relationships in production allocation to leaves, fine roots or wood. However, canopy nutrient pools and fluxes were correlated with the mass of fine roots per unit soil volume and there was a weak but positive correlation of NPP with LAI. Patterns of ) and NUE across the soil developmental sequence were opposite to each other. ) increased as nutrient availability and nutrient turnover increased, while NUE decreased in response to the same influences but reached its highest values where either N or P availability and turnover of both N and P were low. A negative correlation between ) and NUE supports the hypothesis that a trade-off exists between ) and leaf characteristics affecting NUE.     

What controls variation in carbon use efficiency among Amazonian tropical forests?

Why do some forests produce biomass more efficiently than others? Variations in Carbon Use Efficiency (CUE: total Net Primary Production (NPP)/ Gross Primary Production (GPP)) may be due to changes in wood residence time (Biomass/NPP_wood), temperature, or soil nutrient status. We tested these hypotheses in 14, one ha plots across Amazonian and Andean forests where we measured most key components of net primary production (NPP: wood, fine roots, and leaves) and autotrophic respiration (R_a; wood, rhizosphere, and leaf respiration). We found that lower fertility sites were less efficient at producing biomass and had higher rhizosphere respiration, indicating increased carbon allocation to belowground components. We then compared wood respiration to wood growth and rhizosphere respiration to fine root growth and found that forests with residence times <40 yrs had significantly lower maintenance respiration for both wood and fine roots than forests with residence times >40 yrs. A comparison of rhizosphere respiration to fine root growth showed that rhizosphere growth respiration was significantly greater at low fertility sites. Overall, we found that Amazonian forests produce biomass less efficiently in stands with residence times >40 yrs and in stands with lower fertility, but changes to long‐term mean annual temperatures do not impact CUE.     

A historical and comparative review of 50 years of root data collection in Puerto Rico

Fine roots play an important role in plant nutrition, as well as in carbon, water, and nutrient cycling. Fine roots account for a third of terrestrial net primary production (NPP), and inclusion of their structure and function in global carbon models should improve predictions of ecosystem responses to climate change. However, studies focusing on underground plant components are much less frequent than those on aboveground structure. This is more marked in the tropics, where one-third of the planet's terrestrial NPP is produced. Some tropical forests have been more represented in the literature than others, as demonstrated in the collective studies in Puerto Rico. This Caribbean island's biodiversity, frequency of natural disturbances, ease of access to forests, and long-term plots have created an ideal place for the study of tropical ecological processes. This literature review emphasizes 50 years of root research and patterns revealed around Puerto Rico. The data in this review were compiled from scientific publications, conference reports, symposiums, and raw data shared by some researches. Emergent patterns include the shallow distribution of fine roots, the great variation in root biomass among different forest types, little variation in root phosphorus concentrations, the slow recovery of root biomass after Hurricane Hugo, and the fact that most data on roots come from the wet tropical Luquillo Experimental Forest, causing other habitat types to be underrepresented. This review also shows the gaps in knowledge about fine roots in the island's ecosystems, which should be used to promote and guide future studies.     

Tree and grass rooting patterns in an African humid savanna

Abstract. Spatial and temporal soil partitioning between roots of the two savanna plant components, i.e. trees and grasses, were investigated in a West African humid savanna. Vertical root phytomass distribution was described for grass roots, large (> 2 mm) and fine (< 2 mm) tree roots, in open sites and beneath tree canopies. These profiles were established monthly over one year of vegetation growth. Natural ¹³C abundance measurement was used to determine the woody/herbaceous phytomass ratio in root samples. Tree and grass root distributions widely overlapped and both were mostly located in the top 20 cm of the soil. Grass root phytomass decreased with depth whereas woody root phytomass peaked at about 10 cm depth. No time partitioning was detected. These structural results do not support the hypothesis of soil resource partitioning between trees and grasses and are thus consistent with functional results previously reported.     

树木根系碳分配格局及其影响因子

根系作为树木提供养分和水分的“源”和消耗C的“汇”,在陆地生态系统C平衡研究中具有重要的理论意义。尽管20多年来的研究已经认识到根系消耗净初级生产力占总净初级生产力较大的比例,但是,根系（尤其是细根）消耗C的机理以及C分配的去向一直没有研究清楚。主要原因是细根消耗光合产物的生理生态过程相当复杂,准确估计各个组分消耗的C具有很大的不确定性,常常受树种和环境空间和时间异质性、以及研究方法的限制。综述了分配到地下的C主要去向,即细根生产和周转、呼吸及养分吸收与同化、分泌有机物、土壤植食动物,及有关林木地下碳分配机理的几种假说,分析了地下碳分配估计中存在的不确定性。目的是在全球变化C循环研究中对生态系统地下部分根系消耗的C以及分配格局引起重视。     

Toward an allocation scheme for global terrestrial carbon models

The distribution of assimilated carbon among the plant parts has a profound effect on plant growth, and at a larger scale, on terrestrial biogeochemistry. Although important progress has been made in modelling photosynthesis, less effort has been spent on understanding the carbon allocation, especially at large spatial scales. Whereas several individual-level models of plant growth include an allocation scheme, most global terrestrial models still assume constant allocation of net primary production (NPP) among plant parts, without any environmental coupling. Here, we use the CASA biosphere model as a platform for exploring a new global allocation scheme that estimates allocation of photosynthesis products among leaves, stems, and roots depending on resource availability. The philosophy underlying the model is that allocation patterns result from evolved responses that adjust carbon investments to facilitate capture of the most limiting resources, i.e. light, water, and mineral nitrogen. In addition, we allow allocation of NPP to vary in response to changes in atmospheric CO₂. The relative magnitudes of changes in NPP and resource-use efficiency control the response of root:shoot allocation. For ambient CO₂, the model produces realistic changes in above-ground allocation along productivity gradients. In comparison to the CASA standard estimate using fixed allocation ratios, the new allocation scheme tends to favour root allocation, leading to a 10% lower global biomass. Elevated CO₂, which alters the balance between growth and available resources, generally leads to reduced water stress and consequently, decreased root:shoot ratio. The major exception is forest ecosystems, where increased nitrogen stress induces a larger root allocation.     

Comparing global models of terrestrial net primary productivity (NPP): the importance of water availability

Given that neither absolute measures nor direct model validations of global terrestrial net primary productivity (NPP) are feasible, intercomparison of global NPP models provides an effective tool to check model consistency. For this study, we tested the assumption that water availability is the primary limiting factor of NPP in global terrestrial biospheric models. We compared a water balance coefficient (WBC), calculated as the difference of mean annual precipitation and potential evapotranspiration to NPP for each grid cell (0.5° × 0.5° longitude/latitude) in each of 14 models. We also evaluated different approaches used for introducing water budget limitations on NPP: (1) direct physiological control on evapotranspiration through canopy conductance; (2) climatological computation of constraints from supply/demand for ecosystem productivity; and (3) water limitation inferred from satellite data alone. Plots of NPP vs. WBC showed comparable patterns for the models using the same method for water balance limitation on NPP. While correlation plots revealed similar patterns for most global models, other environmental controls on NPP introduced substantial variability.     

Allometric constraints on,and trade‐offs in,belowground carbon allocation and their control of soil respiration across global forest ecosystems

To fully understand how soil respiration is partitioned among its component fluxes and responds to climate, it is essential to relate it to belowground carbon allocation, the ultimate carbon source for soil respiration. This remains one of the largest gaps in knowledge of terrestrial carbon cycling. Here, we synthesize data on gross and net primary production and their components, and soil respiration and its components, from a global forest database, to determine mechanisms governing belowground carbon allocation and their relationship with soil respiration partitioning and soil respiration responses to climatic factors across global forest ecosystems. Our results revealed that there are three independent mechanisms controlling belowground carbon allocation and which influence soil respiration and its partitioning: an allometric constraint; a fine‐root production vs. root respiration trade‐off; and an above‐ vs. belowground trade‐off in plant carbon. Global patterns in soil respiration and its partitioning are constrained primarily by the allometric allocation, which explains some of the previously ambiguous results reported in the literature. Responses of soil respiration and its components to mean annual temperature, precipitation, and nitrogen deposition can be mediated by changes in belowground carbon allocation. Soil respiration responds to mean annual temperature overwhelmingly through an increasing belowground carbon input as a result of extending total day length of growing season, but not by temperature‐driven acceleration of soil carbon decomposition, which argues against the possibility of a strong positive feedback between global warming and soil carbon loss. Different nitrogen loads can trigger distinct belowground carbon allocation mechanisms, which are responsible for different responses of soil respiration to nitrogen addition that have been observed. These results provide new insights into belowground carbon allocation, partitioning of soil respiration, and its responses to climate in forest ecosystems and are, therefore, valuable for terrestrial carbon simulations and projections.     

A cyclical but asynchronous pattern of fine root and woody biomass production in a hardwood forest of southern Quebec and its relationships with annual variation of temperature and nutrient availability

In contrast to the well-documented seasonal variation in growth of below- and above-ground components of trees, the annual variation in below- and aboveground production is not well understood. In this study, we report on the monitoring of an unmanaged hardwood forest ecosystem in a small watershed of southern Quebec between 1993 and 1999. Below- and above-ground biomass production, leaf and soil solution chemistry, and air temperature were measured at different regular intervals and are reported on an annual basis. The objective of the study was to describe the annual dynamics of carbon partitioning between below- and above-ground tree components and to gain a better understanding of the soil and climatic factors that govern it. Fine root production peaked one year earlier than woody biomass production and years with high production of fine roots had low woody biomass production. All models that included May temperature in the calculation of the predicting/independent variables were significant predictors of total tree biomass production (r > 0.87). Fine root production was associated negatively with the previous year average growing season temperature (r < -0.84). Soil solution NO₃ ^–, NH₄ ⁺ and NO₃ ^– + NH₄ ⁺ concentrations were positively correlated with fine root production (r > 0.72) and negatively correlated with woody biomass production (r < -0.84). Leaf N and P concentrations were negatively correlated (r = -0.99 and r = -0.98, respectively) with fine root production for the period of 1994–1998. Our results suggest that a cool growing season, and in particular a cool month of October, is likely to result in low fine root production and nutrient uptake the following year and, therefore, to increase soil N availability and decrease leaf N. This initial response is thought to be the first step of a feedback loop involving plant N nutrition, soil N availability, fine root growth and aboveground biomass production that led to a cyclical (3–4 years) but asynchronous production of fine roots and aboveground biomass production.     

基于模型数据融合的长白山阔叶红松林碳循环模拟

 充分、有效地利用各种陆地生态系统碳观测数据改善陆地生态系统模型, 是当前我国陆地生态系统碳循环研究领域亟待解决的重要问题之一。该研究以2003~2005年长白山阔叶红松林的6组生物计量观测数据和涡度相关技术测定的碳通量数据为基础, 利用马尔可夫链-蒙特卡罗方法对陆地生态系统模型的关键参数(即碳滞留时间)进行了反演, 进而预测了长白山阔叶红松林生态系统碳库、碳通量及其不确定性。反演结果表明, 长白山阔叶红松林叶凋落物和微生物碳的平均滞留时间最短, 为2~6个月; 其次是叶和细根生物量碳, 二者的平均滞留时间为1~2 a; 慢性土壤有机碳的平均滞留时间为8~16 a; 碳在木质生物量和惰性土壤有机质库中的滞留时间最长, 平均滞留时间分别为77~109 a和409~1 879 a。模拟结果显示, 碳库和累积碳通量模拟值的不确定性将随着模拟时间的延长而增大。当气温升高10%和20%时, 长白山阔叶红松林总初级生产力年总量将分别增加6.5%和9.9%, 净生态系统生产力(NEP)年总量的变化取决于土壤温度的变化。若土壤温度保持不变, NEP年总量将分别增加11.4%~21.9%和17.6%~33.1%; 若土壤温度也相应升高10%和20%, NEP年总量的增幅反而下降甚至低于原来的水平。假设气候和植被保持在2003~2005年的状态, 2020年长白山阔叶红松林NEP年总量为(163±12) g C·m^–2·a^–1, 土壤呼吸年总量为(721±14) g C·m^–2·a^–1。马尔可夫链-蒙特卡罗方法是反演模型参数、优化模拟结果和评估模拟结果不确定性的有效方法, 但今后仍需在惰性土壤碳滞留时间的估计、驱动数据和模型结构的不确定性分析、模型数据融合方法方面进行深入研究, 以进一步提高碳循环模拟的准确性。     

Fine root branch orders respond differentially to carbon source-sink manipulations in a longleaf pine forest

Fine roots are a key component of carbon (C) flow and nitrogen (N) cycling in forest ecosystems. However, the complexity and heterogeneity of the fine root branching system have hampered the assessment and prediction of C and N dynamics at ecosystem scales. We examined how root morphology, biomass, and chemistry differed with root branch orders (1–5 with root tips classified as first order roots) and how different root orders responded to increased C sink strength (via N fertilization) and reduced carbon source strength (via canopy scorching) in a longleaf pine (Pinus palustris L.) ecosystem. With increasing root order, the diameter and length of individual roots increased, whereas the specific root length decreased. Total root biomass on an areal basis was similar among the first four orders but increased for the fifth order roots. Consequently, total root length and total root surface area decreased systematically with increasing root order. Fine root N and lignin concentrations decreased, while total non-structural carbohydrate (TNC) and cellulose concentrations increased with increasing root order. N addition and canopy disturbance did not alter root morphology, but they did influence root chemistry. N fertilization increased fine root N concentration and content per unit area in all five orders, while canopy scorching decreased root N concentration. Moreover, TNC concentration and content in fifth order roots were also reduced by canopy scorching. Our results indicate that the small, fragile, and more easily overlooked first and second order roots may be disproportionately important in ecosystem scale C and N fluxes due to their large proportions of fine root biomass, high N concentrations, relatively short lifespans, and potentially high decomposition rates.Electronic Supplementary Material Supplementary material is available for this article at     

Comprehensive assessments of root biomass and production in a Kobresia humilis meadow on the Qinghai-Tibetan Plateau

Alpine meadow covers ca. 700,000 km² with an extreme altitude range from 3200 m to 5200 m. It is the most widely distributed vegetation on the vast Qinghai-Tibetan Plateau. Previous studies suggest that meadow ecosystems play the most important role in both uptake and storage of carbon in the plateau. The ecosystem has been considered currently as an active “CO₂ sink”, in which roots may contribute a very important part, because of the large root biomass, for storage and translocation of carbon to soil. To bridge the gap between the potential importance and few experimental data, root systems, root biomass, turnover rate, and net primary production were investigated in a Kobresia humilis meadow on the plateau during the growing season from May to September in 2008 and 2009. We hypothesized that BNPP/NPP of the alpine meadow would be more than 50%, and that small diameter roots sampled in ingrowth cores have a shorter lifespan than the lager diameter roots, moreover we expected that roots in surface soils would turn over more quickly than those in deeper soil layers. The mean root mass in the 0–20 cm soil layer, investigated by the sequential coring method, was 1995?±?479 g?m^?2 and 1595?±?254 g?m^?2 in growing season of 2008 and 2009, respectively. And the mean fine root biomass in ingrowth cores of the same soil layer was 119?±?37 g?m^?2 and 196?±?45 g?m^?2 in the 2 years. Annual total NPP was 12387 kg?ha^?1?year^?1, in which 53% was allocated to roots. In addition, fine roots accounted for 33% of belowground NPP and 18% of the total NPP, respectively. Root turnover rate was 0.52 year^?1 for bulk roots and 0.74 year^?1 for fine roots. Furthermore, roots turnover was faster in surface than in deeper soil layers. The results confirmed the important role of roots in carbon storage and turnover in the alpine meadow ecosystem. It also suggested the necessity of separating fine roots from the whole root system for a better understanding of root turnover rate and its response to environmental factors.     

Patterns of above- and belowground biomass allocation in China’s grasslands: Evidence from individual-level observations

Above- and belowground biomass allocation not only influences growth of individual plants, but also influences vegetation structures and functions, and consequently impacts soil carbon input as well as terrestrial ecosystem carbon cycling. However, due to sampling difficulties, a considerable amount of uncertainty remains about the root: shoot ratio (R/S), a key parameter for models of terrestrial ecosystem carbon cycling. We investigated biomass allocation patterns across a broad spatial scale. We collected data on individual plant biomass and systematically sampled along a transect across the temperate grasslands in Inner Mongolia as well as in the alpine grasslands on the Tibetan Plateau. Our results indicated that the median of R/S for herbaceous species was 0.78 in China’s grasslands as a whole. R/S was significantly higher in temperate grasslands than in alpine grasslands (0.84 vs. 0.65). The slope of the allometric relationship between above- and belowground biomass was steeper for temperate grasslands than for alpine. Our results did not support the hypothesis that aboveground biomass scales isometrically with belowground biomass. The R/S in China’s grasslands was not significantly correlated with mean annual temperature (MAT) or mean annual precipitation (MAP). Moreover, comparisons of our results with previous findings indicated a large difference between R/S data from individual plants and communities. This might be mainly caused by the underestimation of R/S at the individual level as a result of an inevitable loss of fine roots and the overestimation of R/S in community-level surveys due to grazing and difficulties in identifying dead roots. Our findings suggest that root biomass in grasslands tended to have been overestimated in previous reports of R/S.     

Carbon allocation in forest ecosystems

Carbon allocation plays a critical role in forest ecosystem carbon cycling. We reviewed existing literature and compiled annual carbon budgets for forest ecosystems to test a series of hypotheses addressing the patterns, plasticity, and limits of three components of allocation: biomass, the amount of material present; flux, the flow of carbon to a component per unit time; and partitioning, the fraction of gross primary productivity (GPP) used by a component. Can annual carbon flux and partitioning be inferred from biomass? Our survey revealed that biomass was poorly related to carbon flux and to partitioning of photosynthetically derived carbon, and should not be used to infer either. Are component fluxes correlated? Carbon fluxes to foliage, wood, and belowground production and respiration all increased linearly with increasing GPP (a rising tide lifts all boats). Autotrophic respiration was strongly linked to production for foliage, wood and roots, and aboveground net primary productivity and total belowground carbon flux (TBCF) were positively correlated across a broad productivity gradient. How does carbon partitioning respond to variability in resources and environment? Within sites, partitioning to aboveground wood production and TBCF responded to changes in stand age and resource availability, but not to competition (tree density). Increasing resource supply and stand age, with one exception, resulted in increased partitioning to aboveground wood production and decreased partitioning to TBCF. Partitioning to foliage production was much less sensitive to changes in resources and environment. Overall, changes in partitioning within a site in response to resource supply and age were small (<15% of GPP), but much greater than those inferred from global relationships. Across all sites, foliage production plus respiration, and total autotrophic respiration appear to use relatively constant fractions of GPP – partitioning to both was conservative across a broad range of GPP – but values did vary across sites. Partitioning to aboveground wood production and to TBCF were the most variable – conditions that favored high GPP increased partitioning to aboveground wood production and decreased partitioning to TBCF. Do priorities exist for the products of photosynthesis? The available data do not support the concept of priorities for the products of photosynthesis, because increasing GPP increased all fluxes. All facets of carbon allocation are important to understanding carbon cycling in forest ecosystems. Terrestrial ecosystem models require information on partitioning, yet we found few studies that measured all components of the carbon budget to allow estimation of partitioning coefficients. Future studies that measure complete annual carbon budgets contribute the most to understanding carbon allocation.     

The age of fine-root carbon in three forests of the eastern United States measured by radiocarbon

Using a new approach involving one-time measurements of radiocarbon (¹⁴C) in fine (<2 mm diameter) root tissues we have directly measured the mean age of fine-root carbon. We find that the carbon making up the standing stock of fine roots in deciduous and coniferous forests of the eastern United States has a mean age of 3-18 years for live fine roots, 10-18 years for dead fine roots, and 3-18 years for mixed live+dead fine roots. These ¹⁴C-derived mean ages represent the time C was stored in the plant before being allocated for root growth, plus the average lifespan (for live roots), plus the average time for the root to decompose (for dead roots and mixtures). Comparison of the ¹⁴C content of roots known to have grown within 1 year with the ¹⁴C of atmospheric CO₂ for the same period shows that root tissues are derived from recently fixed carbon, and the storage time prior to allocation is <2 years and likely <1 year. Fine-root mean ages tend to increase with depth in the soil. Live roots in the organic horizons are made of C fixed 3-8 years ago compared with 11-18 years in the mineral B horizons. The mean age of C in roots increases with root diameter and also is related to branching order. Our results differ dramatically from previous estimates of fine-root mean ages made using mass balance approaches and root-viewing cameras, which generally report life spans (mean ages for live roots) of a few months to 1-2 years. Each method for estimating fine-root dynamics, including this new radiocarbon method, has biases. Root-viewing approaches tend to emphasize more rapidly cycling roots, while radiocarbon ages tend to reflect those components that persist longest in the soil. Our ¹⁴C-derived estimates of long mean ages can be reconciled with faster estimates only if fine-root populations have varying rates of root mortality and decomposition. Our results indicate that a standard definition of fine roots, as those with diameters of <2 mm, is inadequate to determine the most dynamic portion of the root population. Recognition of the variability in fine-root dynamics is necessary to obtain better estimates of belowground C inputs.     

Inverse analysis of coupled carbon-nitrogen cycles against multiple datasets at ambient and elevated CO2

Aims Carbon (C) sequestration in terrestrial ecosystems is strongly regulated by nitrogen (N) processes. However, key parameters that determine the degree of N regulation on terrestrial C sequestration have not been well quantified.Methods Here, we used a Bayesian probabilistic inversion approach to estimate 14 target parameters related to ecosystem C and N interactions from 19 datasets obtained from Duke Forests under ambient and elevated carbon dioxide (CO₂).Important findings Our results indicated that 8 of the 14 target parameters, such as C:N ratios in most ecosystem compartments, plant N uptake and external N input, were well constrained by available datasets whereas the others, such as N allocation coefficients, N loss and the initial value of mineral N pool were poorly constrained. Our analysis showed that elevated CO₂ led to the increases in C:N ratios in foliage, fine roots and litter. Moreover, elevated CO₂ stimulated plant N uptake and increased ecosystem N capital in Duke Forests by 25.2 and 8.5%, respectively. In addition, elevated CO₂ resulted in the decrease of C exit rates (i.e. increases in C residence times) in foliage, woody biomass, structural litter and passive soil organic matter, but the increase of C exit rate in fine roots. Our results demonstrated that CO₂ enrichment substantially altered key parameters in determining terrestrial C and N interactions, which have profound implications for model improvement and predictions of future C sequestration in terrestrial ecosystems in response to global change.     

Elevated carbon dioxide and ozone alter productivity and ecosystem carbon content in northern temperate forests

Three young northern temperate forest communities in the north‐central United States were exposed to factorial combinations of elevated carbon dioxide (CO₂) and tropospheric ozone (O₃) for 11 years. Here, we report results from an extensive sampling of plant biomass and soil conducted at the conclusion of the experiment that enabled us to estimate ecosystem carbon (C) content and cumulative net primary productivity (NPP). Elevated CO₂ enhanced ecosystem C content by 11%, whereas elevated O₃ decreased ecosystem C content by 9%. There was little variation in treatment effects on C content across communities and no meaningful interactions between CO₂ and O₃. Treatment effects on ecosystem C content resulted primarily from changes in the near‐surface mineral soil and tree C, particularly differences in woody tissues. Excluding the mineral soil, cumulative NPP was a strong predictor of ecosystem C content (r² = 0.96). Elevated CO₂ enhanced cumulative NPP by 39%, a consequence of a 28% increase in canopy nitrogen (N) content (g N m^?2) and a 28% increase in N productivity (NPP/canopy N). In contrast, elevated O₃ lowered NPP by 10% because of a 21% decrease in canopy N, but did not impact N productivity. Consequently, as the marginal impact of canopy N on NPP (?NPP/?N) decreased through time with further canopy development, the O₃ effect on NPP dissipated. Within the mineral soil, there was less C in the top 0.1 m of soil under elevated O₃ and less soil C from 0.1 to 0.2 m in depth under elevated CO₂. Overall, these results suggest that elevated CO₂ may create a sustained increase in NPP, whereas the long‐term effect of elevated O₃ on NPP will be smaller than expected. However, changes in soil C are not well‐understood and limit our ability to predict changes in ecosystem C content.     

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.