Forest responses to simulated elevated CO2 under alternate hypotheses of size‐ and age‐dependent mortality

Elevated atmospheric carbon dioxide (eCO₂) is predicted to increase growth rates of forest trees. The extent to which increased growth translates to changes in biomass is dependent on the turnover time of the carbon, and thus tree mortality rates. Size‐ or age‐dependent mortality combined with increased growth rates could result in either decreased carbon turnover from a speeding up of tree life cycles, or increased biomass from trees reaching larger sizes, respectively. However, most vegetation models currently lack any representation of size‐ or age‐dependent mortality and the effect of eCO₂ on changes in biomass and carbon turnover times is thus a major source of uncertainty in predictions of future vegetation dynamics. Using a reduced‐complexity form of the vegetation demographic model the Functionally Assembled Terrestrial Ecosystem Simulator to simulate an idealised tropical forest, we find increases in biomass despite reductions in carbon turnover time in both size‐ and age‐dependent mortality scenarios in response to a hypothetical eCO₂‐driven 25% increase in woody net primary productivity (wNPP). Carbon turnover times decreased by 9.6% in size‐dependent mortality scenarios due to a speeding up of tree life cycles, but also by 2.0% when mortality was age‐dependent, as larger crowns led to increased light competition. Increases in aboveground biomass (AGB) were much larger when mortality was age‐dependent (24.3%) compared with size‐dependent (13.4%) as trees reached larger sizes before death. In simulations with a constant background mortality rate, carbon turnover time decreased by 2.1% and AGB increased by 24.0%, however, absolute values of AGB and carbon turnover were higher than in either size‐ or age‐dependent mortality scenario. The extent to which AGB increases and carbon turnover decreases will thus depend on the mechanisms of large tree mortality: if increased size itself results in elevated mortality rates, then this could reduce by about half the increase in AGB relative to the increase in wNPP.     

林木分化对兴安落叶松异速生长方程和生物量分配的影响

林木因对资源竞争而产生分化,从而影响林木的异速生长方程和生物量分配,但其影响程度还不清楚。采用林木相对直径法将38株兴安落叶松(Larix gmelinii)样木在林分中的分化等级分为优势木、中等木和被压木,量化林木分化对林木异速生长方程和生物量分配的影响。结果显示:生物量组分异速生长方程多以胸径(DBH)为自变量为好,但以枝下高处的树干直径为自变量估测其枝、叶生物量时更精确。在一定的胸径范围内,同一胸径下不同林木分化等级的地下部分各组分生物量没有显著差异(P0.05),但优势木分配更多的生物量给枝和叶,中等木比优势木分配更多的生物量给树干,中等木比被压木分配更多的生物量给地上部分,而且被压木和中等木的树高显著高于优势木。除根茎生物量之外,不同林木分化等级的生物量组分(包括枝、叶、树干和根系)的相对分配比例无显著差异(P0.05),根冠比保持相对稳定。这些结果表明,主要由竞争而引起的林木分化改变了兴安落叶松地上生物量组分的异速生长和分配,但其相对分配格局较为保守。     

Nitrogen and phosphorus dynamics of a tallgrass prairie ecosystem exposed to elevated carbon dioxide

A tallgrass prairie ecosystem was exposed to ambient and twice-ambient CO₂ concentrations in open-top chambers and compared to unchambered ambient CO₂ during the entire growing season from 1989 through 1991. Dominant species were Andropogon gerardii (C₄), A. scoparius (C₄), Sorghastrum nutans (C₄) and Poa pratensis (C₃). Nitrogen and phosphorus concentrations in A. gerardii, P. pratensis and dicotyledonous herbs above ground biomass were estimated by periodic sampling throughout the growing season in 1989 and 1990. In 1991, N and P concentrations in peak biomass were estimated by an early August harvest. N and P concentrations in root production as a function of treatment were estimated using root ingrowth bags that remained in place throughout the growing season. Total N and P in above- and belowground biomass were calculated as products of concentration and peak biomass by species groups. N concentration in A. gerardii and dicotyledonous herb aboveground biomass was lower and total N higher in elevated CO₂ plots than in ambient CO₂ plots. N concentration in P. pratensis aboveground biomass was lower in elevated CO₂ plots than in ambient, but total N did not differ among treatments in 2 out of 3 years. In 1990, N concentration in root ingrowth bag biomass was lower and total N greater in elevated CO₂ than in ambient CO₂ plots. Root ingrowth bag biomass N concentration did not differ among treatments in 1991, but total N was greater in elevated CO₂ plots than in ambient CO₂ plots. P concentration was lower under elevated CO₂ compared to ambient in 1989, but did not differ substantially among treatments in 1990 or 1991. In all years, total P in aboveground A. gerardii and root ingrowth bag biomass was greater under elevated CO₂ than ambient. P concentration and total P in P. pratensis was similar among treatments.     

CO2 enrichment in a maturing pine forest: are CO2 exchange and water status in the canopy affected?

A _net, leaf net CO₂ assimilation
c_a, CO₂ concentration of air surrounding a leaf
c_i, leaf intercellular CO₂ concentration
Δ, ¹³C isotope discrimination
δ¹³C, relative stable carbon isotope content
?, ratio of A_net at c_a = 560μmol mol^–1 to A_net at c_a = 360 μmol mol^–1
FACE, free-air CO₂ enrichment
g_w, stomatal conductance to water vapour
Π_i, initial leaf osmotic potential
R_t, relative water content at incipient turgor loss
Ψ_l, xylem water potential of leaves
Ψ_m, soil matric potential

Elevated CO₂ is expected to reduce forest water use as a result of CO₂-induced stomatal closure, which has implications for ecosystem-scale phenomena controlled by water availability. Leaf-level CO₂ and H₂O exchange responses and plant and soil water relations were examined in a maturing loblolly pine (Pinus taeda L.) stand in a free-air CO₂ enrichment (FACE) experiment in North Carolina, USA to test if these parameters were affected by elevated CO₂. Current-year foliage in the canopy was continuously exposed to elevated CO₂ (ambient CO₂+200μmol mol^–1) in free-air during needle growth and development for up to 400 d. Photosynthesis in upper canopy foliage was stimulated by 50–60% by elevated CO₂ compared with ambient controls. This enhancement was similar in current-year, ambient-grown foliage temporarily measured at elevated CO₂ compared with long-term elevated CO₂ grown foliage. Significant photosynthetic enhancement by CO₂ was maintained over a range of conditions except during peak drought. There was no evidence of water savings in elevated CO₂ plots in FACE compared to ambient plots under drought and non-drought conditions. This was supported by evidence from three independent measures. First, stomatal conductance was not significantly different in elevated CO₂ versus ambient trees of P. taeda. Calculations of time-integrated c_i/c_a ratios from analysis of foliar δ¹³C showed that these ratios were maintained in foliage under elevated CO₂. Second, soil moisture was not significantly different between ambient and elevated CO₂ plots during drought. Third, pre-dawn and mid-day leaf water potentials were also unaffected by the seasonal CO₂ exposure, as were tissue osmotic potentials and turgor loss points. Together the results strongly support the hypothesis that maturing P. taeda trees have low stomatal responsiveness to elevated CO₂. Elevated CO₂ effects on water relations in loblolly pine-dominated forest ecosystems may be absent or small apart from those mediated by leaf area. Large photosynthetic enhancements in the upper canopy of P. taeda by elevated CO₂ indicate that this maturing forest may have a large carbon sequestration capacity with limiting water supply.     

Nitrogen input mediates the effect of free-air CO2 enrichment on mycorrhizal fungal abundance

Plots containing Lolium perenne L., Trifolium repens L. or a mixture of both plant species were exposed to elevated atmospheric CO₂ (eCO₂) for 10 consecutive seasons using free‐air CO₂ enrichment technology at ETH Zürich, Switzerland. The CO₂ treatment was crossed with a two‐level nitrogen (N) fertilization treatment. In the tenth year, soil samples were collected on three occasions through the growing season to assess the impact of eCO₂ and N fertilization on mycorrhizal fungal abundance. Soil moisture content, which varied with harvest date, was linked to the vegetation type and was higher under eCO₂. Root weight density was affected by vegetation type: lower for clover, higher for grass. Root weight density was stimulated by eCO₂ and decreased by high N fertilization. The percent root length colonized by mycorrhizal fungi was lowest in the clover plots and highest in the grass plots. High N significantly decreased root length colonized. There was no overall effect of eCO₂ on root length colonized; however, there was a significant eCO₂× N interaction: eCO₂ increased root length colonized at high N, but decreased root length colonized at low N. Extraradical mycorrhizal hyphal density was linked to soil moisture content. Extraradical mycorrhizal hyphal density was not affected by eCO₂ or high N individually, but as for root length colonized, there was a significant eCO₂× N interaction: eCO₂ increased extraradical mycorrhizal hyphal density at low N but not at high N. These environmental effects on root colonization and external mycorrhizal hyphae were independent of soil moisture content and root weight density. This field study demonstrated a significant mediating effect of N fertilization on the response of arbuscular mycorrhizal fungi to eCO₂ irrespective of any change in root biomass.     

Biomass responses in a temperate European grassland through 17 years of elevated CO2

Future increase in atmospheric CO₂ concentrations will potentially enhance grassland biomass production and shift the functional group composition with consequences for ecosystem functioning. In the “GiFACE” experiment (Giessen Free Air Carbon dioxide Enrichment), fertilized grassland plots were fumigated with elevated CO₂ (eCO₂) year‐round during daylight hours since 1998, at a level of +20% relative to ambient concentrations (in 1998, aCO₂ was 364 ppm and eCO₂ 399 ppm; in 2014, aCO₂ was 397 ppm and eCO₂ 518 ppm). Harvests were conducted twice annually through 23 years including 17 years with eCO₂ (1998 to 2014). Biomass consisted of C3 grasses and forbs, with a small proportion of legumes. The total aboveground biomass (TAB) was significantly increased under eCO₂ (p = .045 and .025, at first and second harvest). The dominant plant functional group grasses responded positively at the start, but for forbs, the effect of eCO₂ started out as a negative response. The increase in TAB in response to eCO₂ was approximately 15% during the period from 2006 to 2014, suggesting that there was no attenuation of eCO₂ effects over time, tentatively a consequence of the fertilization management. Biomass and soil moisture responses were closely linked. The soil moisture surplus (c. 3%) in eCO₂ manifested in the latter years was associated with a positive biomass response of both functional groups. The direction of the biomass response of the functional group forbs changed over the experimental duration, intensified by extreme weather conditions, pointing to the need of long‐term field studies for obtaining reliable responses of perennial ecosystems to eCO₂ and as a basis for model development.     

Low phosphorus supply constrains plant responses to elevated CO2: A meta‐analysis

Phosphorus (P) is an essential macro‐nutrient required for plant metabolism and growth. Low P availability could potentially limit plant responses to elevated carbon dioxide (eCO₂), but consensus has yet to be reached on the extent of this limitation. Here, based on data from experiments that manipulated both CO₂ and P for young individuals of woody and non‐woody species, we present a meta‐analysis of P limitation impacts on plant growth, physiological, and morphological response to eCO₂. We show that low P availability attenuated plant photosynthetic response to eCO₂ by approximately one‐quarter, leading to a reduced, but still positive photosynthetic response to eCO₂ compared to those under high P availability. Furthermore, low P limited plant aboveground, belowground, and total biomass responses to eCO₂, by 14.7%, 14.3%, and 12.4%, respectively, equivalent to an approximate halving of the eCO₂ responses observed under high P availability. In comparison, low P availability did not significantly alter the eCO₂‐induced changes in plant tissue nutrient concentration, suggesting tissue nutrient flexibility is an important mechanism allowing biomass response to eCO₂ under low P availability. Low P significantly reduced the eCO₂‐induced increase in leaf area by 14.3%, mirroring the aboveground biomass response, but low P did not affect the eCO₂‐induced increase in root length. Woody plants exhibited stronger attenuation effect of low P on aboveground biomass response to eCO₂ than non‐woody plants, while plants with different mycorrhizal associations showed similar responses to low P and eCO₂ interaction. This meta‐analysis highlights crucial data gaps in capturing plant responses to eCO₂ and low P availability. Field‐based experiments with longer‐term exposure of both CO₂ and P manipulations are critically needed to provide ecosystem‐scale understanding. Taken together, our results provide a quantitative baseline to constrain model‐based hypotheses of plant responses to eCO₂ under P limitation, thereby improving projections of future global change impacts.     

Elevated atmospheric carbon dioxide impairs the performance of root‐feeding vine weevils by modifying root growth and secondary metabolites

Predicting how insect crop pests will respond to global climate change is an important part of increasing crop production for future food security, and will increasingly rely on empirically based evidence. The effects of atmospheric composition, especially elevated carbon dioxide (eCO₂), on insect herbivores have been well studied, but this research has focussed almost exclusively on aboveground insects. However, responses of root‐feeding insects to eCO₂ are unlikely to mirror these trends because of fundamental differences between aboveground and belowground habitats. Moreover, changes in secondary metabolites and defensive responses to insect attack under eCO₂ conditions are largely unexplored for root herbivore interactions. This study investigated how eCO₂ (700 μmol mol^?1) affected a root‐feeding herbivore via changes to plant growth and concentrations of carbon (C), nitrogen (N) and phenolics. This study used the root‐feeding vine weevil, Otiorhynchus sulcatus and the perennial crop, Ribes nigrum. Weevil populations decreased by 33% and body mass decreased by 23% (from 7.2 to 5.4 mg) in eCO₂. Root biomass decreased by 16% in eCO₂, which was strongly correlated with weevil performance. While root N concentrations fell by 8%, there were no significant effects of eCO₂ on root C and N concentrations. Weevils caused a sink in plants, resulting in 8–12% decreases in leaf C concentration following herbivory. There was an interactive effect of CO₂ and root herbivory on root phenolic concentrations, whereby weevils induced an increase at ambient CO₂, suggestive of defensive response, but caused a decrease under eCO₂. Contrary to predictions, there was a positive relationship between root phenolics and weevil performance. We conclude that impaired root‐growth underpinned the negative effects of eCO₂ on vine weevils and speculate that the plant's failure to mount a defensive response at eCO₂ may have intensified these negative effects.     

Positive climate feedbacks of soil microbial communities in a semi‐arid grassland

Soil microbial communities may be able to rapidly respond to changing environments in ways that change community structure and functioning, which could affect climate–carbon feedbacks. However, detecting microbial feedbacks to elevated CO₂ (eCO₂) or warming is hampered by concurrent changes in substrate availability and plant responses. Whether microbial communities can persistently feed back to climate change is still unknown. We overcame this problem by collecting microbial inocula at subfreezing conditions under eCO₂ and warming treatments in a semi‐arid grassland field experiment. The inoculant was incubated in a sterilised soil medium at constant conditions for 30 days. Microbes from eCO₂ exhibited an increased ability to decompose soil organic matter (SOM) compared with those from ambient CO₂ plots, and microbes from warmed plots exhibited increased thermal sensitivity for respiration. Microbes from the combined eCO₂ and warming plots had consistently enhanced microbial decomposition activity and thermal sensitivity. These persistent positive feedbacks of soil microbial communities to eCO₂ and warming may therefore stimulate soil C loss.     

Seasonal soil‐surface carbon fluxes from the root systems of young Pinus radiata trees growing at ambient and elevated CO2 concentration

Elevated atmospheric CO₂ concentration may result in increased below‐ground carbon allocation by trees, thereby altering soil carbon cycling. Seasonal estimates of soil surface carbon flux were made to determine whether carbon losses from Pinus radiata trees growing at elevated CO₂ concentration were higher than those at ambient CO₂ concentration, and whether this was related to increased fine root growth. Monthly soil surface carbon flux density (f) measurements were made on plots with trees growing at ambient (350) and elevated (650 μmol mol^?1) CO₂ concentration in large open‐top chambers. Prior to planting the soil carbon concentration (0.1%) and f (0.28 μmol m^?2 s^?1 at 15 °C) were low. A function describing the radial pattern of f with distance from tree stems was used to estimate the annual carbon flux from tree plots. Seasonal estimates of fine root production were made from minirhizotrons and the radial distribution of roots compared with radial measurements of f. A one‐dimensional gas diffusion model was used to estimate f from soil CO₂ concentrations at four depths. For the second year of growth, the annual carbon flux from the plots was 1671 g y^?1 and 1895 g y^?1 at ambient and elevated CO₂ concentrations, respectively, although this was not a significant difference. Higher f at elevated CO₂ concentration was largely explained by increased fine root biomass. Fine root biomass and stem production were both positively related to f. Both root length density and f declined exponentially with distance from the stem, and had similar length scales. Diurnal changes in f were largely explained by changes in soil temperature at a depth of 0.05 m. Ignoring the change of f with increasing distance from tree stems when scaling to a unit ground area basis from measurements with individual trees could result in under‐ or overestimates of soil‐surface carbon fluxes, especially in young stands when fine roots are unevenly distributed.     

Response to CO2 enrichment of understory vegetation in the shade of forests

Responses of forest ecosystems to increased atmospheric CO₂ concentration have been studied in few free‐air CO₂ enrichment (FACE) experiments during last two decades. Most studies focused principally on the overstory trees with little attention given to understory vegetation. Despite its small contribution to total productivity of an ecosystem, understory vegetation plays an important role in predicting successional dynamics and future plant community composition. Thus, the response of understory vegetation in Pinus taeda plantation at the Duke Forest FACE site after 15–17 years of exposure to elevated CO₂, 6–13 of which with nitrogen (N) amendment, was examined. Aboveground biomass and density of the understory decreased across all treatments with increasing overstory leaf area index (LAI). However, the CO₂ and N treatments had no effect on aboveground biomass, tree density, community composition, and the fraction of shade‐tolerant species. The increases of overstory LAI (~28%) under elevated CO₂ resulted in a reduction of light available to the understory (~18%) sufficient to nullify the expected growth‐enhancing effect of elevated CO₂ on understory vegetation.     

Response of wheat growth,grain yield and water use to elevated CO2 under a Free‐Air CO2 Enrichment (FACE) experiment and modelling in a semi‐arid environment

The response of wheat crops to elevated CO₂ (eCO₂) was measured and modelled with the Australian Grains Free‐Air CO₂ Enrichment experiment, located at Horsham, Australia. Treatments included CO₂ by water, N and temperature. The location represents a semi‐arid environment with a seasonal VPD of around 0.5 kPa. Over 3 years, the observed mean biomass at anthesis and grain yield ranged from 4200 to 10 200 kg ha^?1 and 1600 to 3900 kg ha^?1, respectively, over various sowing times and irrigation regimes. The mean observed response to daytime eCO₂ (from 365 to 550 μmol mol^?1 CO₂) was relatively consistent for biomass at stem elongation and at anthesis and LAI at anthesis and grain yield with 21%, 23%, 21% and 26%, respectively. Seasonal water use was decreased from 320 to 301 mm (P = 0.10) by eCO₂, increasing water use efficiency for biomass and yield, 36% and 31%, respectively. The performance of six models (APSIM‐Wheat, APSIM‐Nwheat, CAT‐Wheat, CROPSYST, OLEARY‐CONNOR and SALUS) in simulating crop responses to eCO₂ was similar and within or close to the experimental error for accumulated biomass, yield and water use response, despite some variations in early growth and LAI. The primary mechanism of biomass accumulation via radiation use efficiency (RUE) or transpiration efficiency (TE) was not critical to define the overall response to eCO₂. However, under irrigation, the effect of late sowing on response to eCO₂ to biomass accumulation at DC65 was substantial in the observed data (~40%), but the simulated response was smaller, ranging from 17% to 28%. Simulated response from all six models under no water or nitrogen stress showed similar response to eCO₂ under irrigation, but the differences compared to the dryland treatment were small. Further experimental work on the interactive effects of eCO₂, water and temperature is required to resolve these model discrepancies.     

Improved allometric models to estimate the aboveground biomass of tropical trees

Terrestrial carbon stock mapping is important for the successful implementation of climate change mitigation policies. Its accuracy depends on the availability of reliable allometric models to infer oven‐dry aboveground biomass of trees from census data. The degree of uncertainty associated with previously published pantropical aboveground biomass allometries is large. We analyzed a global database of directly harvested trees at 58 sites, spanning a wide range of climatic conditions and vegetation types (4004 trees ≥ 5 cm trunk diameter). When trunk diameter, total tree height, and wood specific gravity were included in the aboveground biomass model as covariates, a single model was found to hold across tropical vegetation types, with no detectable effect of region or environmental factors. The mean percent bias and variance of this model was only slightly higher than that of locally fitted models. Wood specific gravity was an important predictor of aboveground biomass, especially when including a much broader range of vegetation types than previous studies. The generic tree diameter–height relationship depended linearly on a bioclimatic stress variable E, which compounds indices of temperature variability, precipitation variability, and drought intensity. For cases in which total tree height is unavailable for aboveground biomass estimation, a pantropical model incorporating wood density, trunk diameter, and the variable E outperformed previously published models without height. However, to minimize bias, the development of locally derived diameter–height relationships is advised whenever possible. Both new allometric models should contribute to improve the accuracy of biomass assessment protocols in tropical vegetation types, and to advancing our understanding of architectural and evolutionary constraints on woody plant development.     

Response of maize biomass and soil water fluxes on elevated CO2 and drought—From field experiments to process‐based simulations

The rising concentration of atmospheric carbon dioxide (CO₂) is known to increase the total aboveground biomass of several C3 crops, whereas C4 crops are reported to be hardly affected when water supply is sufficient. However, a free‐air carbon enrichment (FACE) experiment in Braunschweig, Germany, in 2007 and 2008 resulted in a 25% increased biomass of the C4 crop maize under restricted water conditions and elevated CO₂ (550 ppm). To project future yields of maize under climate change, an accurate representation of the effects of eCO₂ and drought on biomass and soil water conditions is essential. Current crop growth models reveal limitations in simulations of maize biomass under eCO₂ and limited water supply. We use the coupled process‐based hydrological‐plant growth model Catchment Modeling Framework‐Plant growth Modeling Framework to overcome this limitation. We apply the coupled model to the maize‐based FACE experiment in Braunschweig that provides robust data for the investigation of combined CO₂ and drought effects. We approve hypothesis I that CO₂ enrichment has a small direct‐fertilizing effect with regard to the total aboveground biomass of maize and hypothesis II that CO₂ enrichment decreases water stress and leads to higher yields of maize under restricted water conditions. Hypothesis III could partly be approved showing that CO₂ enrichment decreases the transpiration of maize, but does not raise soil moisture, while increasing evaporation. We emphasize the importance of plant‐specific CO₂ response factors derived by use of comprehensive FACE data. By now, only one FACE experiment on maize is accomplished applying different water levels. For the rigorous testing of plant growth models and their applicability in climate change studies, we call for datasets that go beyond single criteria (only yield response) and single effects (only elevated CO₂).     

Fertilization of boreal forest reduces both autotrophic and heterotrophic soil respiration

The boreal forest is expected to experience the greatest warming of all forest biomes, raising concerns that some of the large quantities of soil carbon in these systems may be added to the atmosphere as CO₂. However, nitrogen deposition or fertilization has the potential to increase boreal forest production and retard the decomposition of soil organic matter, hence increasing both tree stand and soil C storage. The major contributors to soil‐surface CO₂ effluxes are autotrophic and heterotrophic respiration. To evaluate the effect of nutrient additions on the relative contributions from autotrophic and heterotrophic respiration, a large‐scale girdling experiment was performed in a long‐term nutrient optimization experiment in a 40‐year‐old stand of Norway spruce in northern Sweden. Trees on three nonfertilized plots and three fertilized plots were girdled in early summer 2002, and three nonfertilized and three fertilized plots were used as control plots. Each plot was 0.1 ha and contained around 230 trees. Soil‐surface CO₂ fluxes, soil moisture, and soil temperature were monitored in both girdled and nongirdled plots. In late July, the time of the seasonal maximum in soil‐surface CO₂ efflux, the total soil‐CO₂ efflux in nongirdled plots was 40% lower in the fertilized than in the nonfertilized plots, while the efflux in girdled fertilized and nonfertilized plots was 50% and 60% lower, respectively, than in the corresponding nongirdled controls. We attribute these reductions to losses of the autotrophic component of the total soil‐surface CO₂ efflux. The estimates of autotrophic respiration are conservative as root starch reserves were depleted more rapidly in roots of girdled than in nongirdled trees. Thus, heterotrophic activity was overestimated. Calculated on a unit area basis, both the heterotrophic and autotrophic soil respiration was significantly lower in fertilized plots, which is especially noteworthy given that aboveground production was around three times higher in fertilized than in nonfertilized plots.     

Elevated CO2 and tree fecundity: the role of tree size, interannual variability, and population heterogeneity

Long‐term population effects of changes in atmospheric CO₂ will be largely determined by reproductive effort. Our research objectives were to quantify variability in seed production and rate of maturation among individual Pinus taeda L. (Pinaceae) trees growing in elevated CO₂ (ambient plus 200 μL L^?1) since 1996. Estimating tree fecundity in nature is frustrated by the difficulty of counting seeds from individual trees and the need for long‐term data. We have used a hierarchical Bayes approach to model individual tree fecundity, accounting for the complexity of experimentation in a natural setting over multiple years. The study presented here demonstrates large variability in natural fecundity rates and contributes to our understanding of how both interannual variation and population heterogeneity influence elevated CO₂ effects. We found that trees growing under elevated CO₂ matured earlier and produced more seeds and cones per unit basal area than ambient grown trees. By 2004, trees grown in high CO₂ had produced an average 300 more seeds per tree than ambient grown trees. Although there was a trend toward decreasing mean CO₂ effect (difference in fecundity between elevated and ambient treatments) over time, the hierarchical analysis indicates that this decrease comes from the emergence of a few highly fecund ambient grown trees by 2002, rather than acclimation or downregulation among the fumigated trees. The most important effect of increased CO₂ in forest ecosystems may be the increase in fecundity reported here. Although biomass responses can sometimes be large, the increase in fecundity can have long‐term impacts on forest dynamics that transcend the current generation.     

Soil warming and CO2 enrichment induce biomass shifts in alpine tree line vegetation

Responses of alpine tree line ecosystems to increasing atmospheric CO₂ concentrations and global warming are poorly understood. We used an experiment at the Swiss tree line to investigate changes in vegetation biomass after 9 years of free air CO₂ enrichment (+200 ppm; 2001–2009) and 6 years of soil warming (+4 °C; 2007–2012). The study contained two key tree line species, Larix decidua and Pinus uncinata, both approximately 40 years old, growing in heath vegetation dominated by dwarf shrubs. In 2012, we harvested and measured biomass of all trees (including root systems), above‐ground understorey vegetation and fine roots. Overall, soil warming had clearer effects on plant biomass than CO₂ enrichment, and there were no interactive effects between treatments. Total plant biomass increased in warmed plots containing Pinus but not in those with Larix. This response was driven by changes in tree mass (+50%), which contributed an average of 84% (5.7 kg m^?2) of total plant mass. Pinus coarse root mass was especially enhanced by warming (+100%), yielding an increased root mass fraction. Elevated CO₂ led to an increased relative growth rate of Larix stem basal area but no change in the final biomass of either tree species. Total understorey above‐ground mass was not altered by soil warming or elevated CO₂. However, Vaccinium myrtillus mass increased with both treatments, graminoid mass declined with warming, and forb and nonvascular plant (moss and lichen) mass decreased with both treatments. Fine roots showed a substantial reduction under soil warming (?40% for all roots <2 mm in diameter at 0–20 cm soil depth) but no change with CO₂ enrichment. Our findings suggest that enhanced overall productivity and shifts in biomass allocation will occur at the tree line, particularly with global warming. However, individual species and functional groups will respond differently to these environmental changes, with consequences for ecosystem structure and functioning.     

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.     

Wood properties and ring width responses to long-term atmospheric CO2 enrichment in field-grown loblolly pine (Pinus taeda L.)

Loblolly pine (Pinus taeda L.) were grown in the field, under non-limiting nutrient conditions, in open-top chambers for 4 years at ambient CO₂ partial pressures (pCO₂) and with a CO₂-enriched atmosphere (+ 30 Pa pCO₂ compared to ambient concentration). A third replicate of trees were grown without chambers at ambient pCO₂. Wood anatomy, wood density and tree ring width were analysed using stem wood samples. No significant differences were observed in the cell wall to cell lumen ratio within the latewood of the third growth ring formed in 1994. No significant differences were observed in the density of resin canals or in the ratio of resin canal cross-sectional area to xylem area within the same growth ring. Ring widths were significantly wider in the CO₂-enrichment treatment for 3 of 4 years compared to the ambient chamber control treatment. Latewood in the 1995 growth ring was significantly wider than that in the ambient control and represented a larger percentage of the total growth-ring width. Carbon dioxide enrichment also significantly increased the total wood specific gravity (determined by displacement). However, when determined as total sample wood density by X-ray densitometry, the density of enriched samples was not significantly higher than that of the ambient chamber controls. Only the 1993 growth ring of enriched trees had a significantly higher maximum latewood density than that of trees grown on non-chambered plots or ambient chambered controls. No significant differences were observed in the minimum earlywood density of individual growth rings between chambered treatments. These results show that the most significant effect of CO₂ enrichment on wood production in loblolly pine is its influence on radial growth, measured as annual tree ring widths. This influence is most pronounced in the first year of growth and decreases with age.     

Elevated carbon dioxide concentrations indirectly affect plant fitness by altering plant tolerance to herbivory

Global environmental changes, such as rising atmospheric CO₂ concentrations, have a wide range of direct effects on plant physiology, growth, and fecundity. These environmental changes also can affect plants indirectly by altering interactions with other species. Therefore, the effects of global changes on a particular species may depend on the presence and abundance of other community members. We experimentally manipulated atmospheric CO₂ concentration and amounts of herbivore damage (natural insect folivory and clipping to simulate browsing) to examine: (1) how herbivores mediate the effects of elevated CO₂ (eCO₂) on the growth and fitness of Arabidopsis thaliana; and (2) how predicted changes in CO₂ concentration affect plant resistance to herbivores, which influences the amount of damage plants receive, and plant tolerance of herbivory, or the fitness consequences of damage. We found no evidence that CO₂ altered resistance, but plants grown in eCO₂ were less tolerant of herbivory—clipping reduced aboveground biomass and fruit production by 13 and 22%, respectively, when plants were reared under eCO₂, but plants fully compensated for clipping in ambient CO₂ (aCO₂) environments. Costs of tolerance in the form of reduced fitness of undamaged plants were detected in eCO₂ but not aCO₂ environments. Increased costs could reduce selection on tolerance in eCO₂ environments, potentially resulting in even larger fitness effects of clipping in predicted future eCO₂ conditions. Thus, environmental perturbations can indirectly affect both the ecology and evolution of plant populations by altering both the intensity of species interactions as well as the fitness consequences of those interactions.