The influence of elevated O3 and CO2 concentrations on secondary metabolites of Scots pine (Pinus sylvestris L. ) seedlings

Terpene, resin acid and total phenolic concentrations in five‐year‐old Scots pine (Pinus sylvestris L.) seedlings were analysed after exposure to ambient and realistically elevated (2 × ambient) O₃ and CO₂ concentrations and their combination in open‐top chambers during two growing seasons. Under O₃ exposure, limonene concentration in needles and isopimaric concentration in stems decreased significantly. As a response to elevated CO₂, α‐pinene and total phenolic concentrations in needles increased significantly, while bornyl acetate concentration in needles and palustric + levopimaric and neoabietic acid concentrations in stems decreased significantly. Some terpenes and resin acids were found at lower concentrations in the combined O₃ and CO₂ treatment than in O₃ exposure or elevated CO₂. A negative chamber effect was found: seedlings growing inside the chambers with ambient air had significantly lower concentrations of some terpenes and resin acids than seedlings growing outside the chambers. There was a lot of between‐tree variation in terpene and resin acid concentrations, which is typical of open‐pollinated populations. The results of this study suggest that, at least in short‐term experiments, Scots pine secondary metabolites are relatively insensitive to climate change factors. Total phenolics in the needles were the most responsive group showing about 25% increase in elevated CO₂, and O₃ exposure did not mitigate this CO₂ effect. Terpenes and resin acids were less responsive, although some individual compounds showed notable responses, e.g. α‐pinene in needles, which increased about 50% in response to elevated CO₂. As a consequence, although there were only slight effects on total pools of needle secondary metabolites, considerable O₃ and CO₂ effects on certain individual compounds might have ecological significance via trophic amplification, e.g. in decomposing processes of needle litter.     

Decomposition of soybean grown under elevated concentrations of CO2 and O3

A critical global climate change issue is how increasing concentrations of atmospheric CO₂ and ground‐level O₃ will affect agricultural productivity. This includes effects on decomposition of residues left in the field and availability of mineral nutrients to subsequent crops. To address questions about decomposition processes, a 2‐year experiment was conducted to determine the chemistry and decomposition rate of aboveground residues of soybean (Glycine max (L.) Merr.) grown under reciprocal combinations of low and high concentrations of CO₂ and O₃ in open‐top field chambers. The CO₂ treatments were ambient (370 μmol mol^?1) and elevated (714 μmol mol^?1) levels (daytime 12 h averages). Ozone treatments were charcoal‐filtered air (21 nmol mol^?1) and nonfiltered air plus 1.5 times ambient O₃ (74 nmol mol^?1) 12 h day^?1. Elevated CO₂ increased aboveground postharvest residue production by 28–56% while elevated O₃ suppressed it by 15–46%. In combination, inhibitory effects of added O₃ on biomass production were largely negated by elevated CO₂. Plant residue chemistry was generally unaffected by elevated CO₂, except for an increase in leaf residue lignin concentration. Leaf residues from the elevated O₃ treatments had lower concentrations of nonstructural carbohydrates, but higher N, fiber, and lignin levels. Chemical composition of petiole, stem, and pod husk residues was only marginally affected by the elevated gas treatments. Treatment effects on plant biomass production, however, influenced the content of chemical constituents on an areal basis. Elevated CO₂ increased the mass per square meter of nonstructural carbohydrates, phenolics, N, cellulose, and lignin by 24–46%. Elevated O₃ decreased the mass per square meter of these constituents by 30–48%, while elevated CO₂ largely ameliorated the added O₃ effect. Carbon mineralization rates of component residues from the elevated gas treatments were not significantly different from the control. However, N immobilization increased in soils containing petiole and stem residues from the elevated CO₂, O₃, and combined gas treatments. Mass loss of decomposing leaf residue from the added O₃ and combined gas treatments was 48% less than the control treatment after 20 weeks, while differences in decomposition of petiole, stem, and husk residues among treatments were minor. Decreased decomposition of leaf residues was correlated with lower starch and higher lignin levels. However, leaf residues only comprised about 20% of the total residue biomass assayed so treatment effects on mass loss of total aboveground residues were relatively small. The primary influence of elevated atmospheric CO₂ and O₃ concentrations on decomposition processes is apt to arise from effects on residue mass input, which is increased by elevated CO₂ and suppressed by O₃.     

Effects of elevated O3 and CO2 concentrations on photosynthesis and stomatal conductance in Scots pine

Naturally regenerated Scots pines (Pinus sylvestris L.), aged 28–30 years old, were grown in open-top chambers and subjected in situ to three ozone (O₃) regimes, two concentrations of CO₂, and a combination of O₃ and CO₂ treatments From 15 April to 15 September for two growing seasons (1994 and 1995). The gas exchanges of current-year and 1-year-old shoots were measured, along with the nitrogen content of needles. In order to investigate the factors underlying modifications in photosynthesis, five parameters linked to photosynthetic performance and three to stomatal conductance were determined. Elevated O₃ concentrations led to a significant decline in the CO₂ compensation point (Г^*), maximum RuP₂-saturated rate of carboxylation (V_cmax), maximum rate of electron transport (J_max), maximum stomatal conductance (g_smax), and sensitivity of stomatal conductance to changes in leaf-to-air vapour pressure difference (?g_s/?D_v) in both shoot-age classes. However, the effect of elevated O₃ concentrations on the respiration rate in light (R_d) was dependent on shoot age. Elevated CO₂(700 μmol mol^?1) significantly decreased J_max and g_smax but increased R_d in 1-year-old shoots and the ?g_s/?D_v in both shoot-age classes. The interactive effects of O₃ and CO₂ on some key parameters (e.g. V_cmax and J_max) were significant. This may be closely related to regulation of the maximum stomatal conductance and stomatal sensitivity induced by elevated CO₂. As a consequence, the injury induced by O₃ was reduced through decreased ozone uptake in 1-year-old shoots, but not in the current-year shoots. Compared to ambient O₃ concentration, reduced O₃ concentrations (charcoal-filtered air) did not lead to significant changes in any of the measured parameters. Compared to the control treatment, calculations showed that elevated O₃ concentrations decreased the apparent quantum yield by 15% and by 18%, and the maximum rate of photosynthesis by 21% and by 29% in the current-year and 1-year-old shoots, respectively. Changes in the nitrogen content of needles resulting from the various treatments were associated with modifications in photosynthetic components.     

Decomposition of Betula papyrifera leaf litter under the independent and interactive effects of elevated CO2 and O3

Litter decay dynamics of paper birch (Betula papyrifera) were assessed at the Aspen free‐air CO₂ enrichment (FACE) facility in northern Wisconsin, USA. Leaf litter was decomposed for 12 months under factorial combinations of 360 vs. 560 μL CO₂ L^?1, crossed with 36 vs. 55 nL O₃ L^?1. To differentiate between substrate quality and environment effects, litterbags were placed in their Native Plots of origin or transplanted into the other treatments. CO₂ enrichment, regardless of O₃ concentration, produced poorer quality litter (high C/N, lignin/N and condensed tannins) than did ambient CO₂ (low C/N, lignin/N and condensed tannins). Substrate quality differences were reflected in the mass loss rates (k‐values), which were high for litter generated under ambient CO₂ (0.887 year^?1) and low for litter generated under elevated CO₂ (0.674 year^?1). The rate‐retarding effects of CO₂ enrichment were neither alleviated nor exacerbated by O₃ exposure. Decay rates varied, however, depending on whether litter was placed back into its plot of origin or transplanted to Common Gardens. The results of this study are species specific, but they have important implications for understanding the processes regulating storage of fixed C and the release of CO₂ from northern forest ecosystems.     

Variable photosynthetic acclimation in consecutive cohorts of Scots pine needles during 3 years of growth at elevated CO2 and elevated temperature

In this experiment, the photosynthetic acclimation of successive needle cohorts of Scots pine were studied during 3 years of growth at elevated CO₂ and temperature. Naturally regenerated Scots pine (Pinus sylvestris L.) trees were subjected to elevated CO₂ concentration (+CO₂, 700 p.p.m), elevated temperature (+T, ambient +2 to +6 °C) and to a combination of elevated CO₂ and temperature (+CO₂ + T) in closed‐top chambers, starting in August 1996. Trees growing in chambers with ambient CO₂ and ambient temperature served as controls (AmbC). Elevated CO₂ influenced the dark reactions more than the light reactions of photosynthesis, as in the 1996 and 1997 cohorts the carboxylation capacity of Rubisco was reduced in the first and second year of exposure, but there was no consistent change in chlorophyll fluorescence. Net photosynthesis measured at growth concentration of CO₂ was higher at +CO₂ than at AmbC on only one measuring occasion, was generally lower at +T and was not changed at +CO₂ + T. However, trees grown at +T tended to invest more nitrogen (N) in Rubisco, as Rubisco/chlorophyll and the proportion of the total needle N bound to Rubisco occasionally increased. The interaction of +CO₂ and +T on Rubisco was mostly negative; consequently, in the second and third year of the experiment the carboxylation capacity decreased at +CO₂ + T. In the 1996, 1997 and 1998 cohorts, the structural N concentration of needles was lower at +CO₂ than at AmbC. Elevated CO₂ and elevated temperature generally had a positive interaction on N concentration; consequently, N concentration in needles decreased less at +CO₂ + T than at +CO₂. At +CO₂ + T, the acclimation response of needles varied between years and was more pronounced in the 1‐year‐old needles of the 1997 cohort than in those of the 1998 cohort. Thus, acclimation was not always greater in 1‐year‐old needles than in current‐year needles. In the +CO₂ + T treatment, elevated temperature had a greater effect on acclimation of needles than elevated CO₂.     

Wood properties of two silver birch clones exposed to elevated CO2 and O3

Effects of elevated carbon dioxide (CO₂) and ozone (O₃) on wood properties of two initially 7‐year‐old silver birch (Betula pendula Roth) clones were studied after a fumigation during three growing seasons. Forty trees, representing two fast‐growing clones (4 and 80), were exposed in open‐top chambers to the following treatments: outside control, chamber control, 2 × ambient [CO₂], 2 × ambient [O₃] and 2 × ambient [CO₂]+2 × ambient [O₃]. After the 3‐year exposure, the trees were felled and wood properties were analyzed. The treatments affected both stem wood structure and chemistry. Elevated [CO₂] increased annual ring width, and concentrations of extractives and starch, and decreased concentrations of cellulose and gravimetric lignin. Elevated O₃ decreased vessel percentage and increased cell wall percentage in clone 80. In vessel percentage, elevated CO₂ ameliorated the O₃‐induced decrease. In clone 4, elevated O₃ decreased nitrogen concentration of wood. The two clones had different wood properties. In clone 4, the concentrations of extractives, starch, soluble sugars and nitrogen were greater than in clone 80, while in clone 80 the concentrations of cellulose and acid‐soluble lignin were higher. Clone 4 also had slightly longer fibres, greater vessel lumen diameter and vessel percentage than clone 80, while in clone 80 cell wall percentage was greater. Our results show that wood properties of young silver birch trees were altered under elevated CO₂ in both clones, whereas the effects of O₃ depended on clone.     

Altered genotypic and phenotypic frequencies of aphid populations under enriched CO2 and O3 atmospheres

Environmental change is anticipated to negatively affect both plant and animal populations. As abiotic factors rapidly change habitat suitability, projections range from altered genetic diversity to wide-spread species loss. Here, we assess the degree to which changes in atmospheric composition associated with environmental change will influence not only the abundance, but also the genotypic/phenotypic diversity, of herbivore populations. Using free-air CO₂ and O₃ enrichment (FACE) technology, we assess numerical responses of pea aphids (Acyrthosiphon pisum) exhibiting a pink–green genetic polymorphism and an environmentally determined wing polyphenism on broad bean plants (Vicia faba) under enriched CO₂ and/or O₃ atmospheres, over multiple generations. We show that these two greenhouse gases alter not only aphid population sizes, but also genotypic and phenotypic frequencies. As the green genotype was positively influenced by elevated CO₂ levels, but the pink genotype was not, genotypic frequencies (pink morph : green morph) ranged from 1 : 1 to 9 : 1. These two genotypes also displayed marked differences in phenotypic frequencies. The pink genotype exhibited higher levels of wing induction under all atmospheric treatments, however, this polyphenism was negatively influenced by elevated O₃ levels. Resultantly, frequencies of winged phenotypes (pink morph : green morph) varied from 10 : 1 to 332 : 1. Thus, atmospheric conditions associated with environmental change may alter not just overall population sizes, but also genotypic and phenotypic frequencies of herbivore populations, thereby influencing community and ecosystem functioning.     

Effects of elevated O3, alone and in combination with elevated CO2, on tree leaf chemistry and insect herbivore performance: a meta-analysis

We reviewed the effects of elevated ozone (O₃), alone and in combination with elevated carbon dioxide (CO₂) on primary and secondary metabolites of trees and performance of insect herbivores by means of meta‐analysis. Our database consisted of 63 studies conducted on 22 species of trees and published between 1990 and 2005. Ozone alone had no overall effect on concentrations of carbohydrates or nutrients, whereas in combination with CO₂, elevated O₃ reduced nutrient concentrations and increased carbohydrate concentrations. In contrast to primary metabolites, concentrations of phenolics and terpenes were significantly increased by 16% and 8%, respectively, in response to elevated O₃. Effects of ozone in combination with elevated CO₂ were weaker than those of ozone alone on phenolics, but stronger than those of ozone alone on terpenes. The magnitude of secondary metabolite responses depended on the type of ozone exposure facility and increased in the following order: indoor growth chamber 3 than gymnosperms, as shifts in concentrations of carbohydrate and phenolics were observed in the former, but not in the latter. Elevated O₃ had positive effects on some indices of insect performance: pupal mass increased and larval development time shortened, but these effects were counteracted by elevated CO₂. Therefore, despite the observed increase in secondary metabolites, elevated O₃ tends to increase tree foliage quality for herbivores, but elevated CO₂ may alleviate these effects. Our meta‐analysis clearly demonstrated that effects of elevated O₃ alone on leaf chemistry and some indices of insect performance differed from those of O₃+CO₂, and therefore, it is important to study effects of several factors of global climate change simultaneously.     

Species control variation in litter decomposition in a pine forest exposed to elevated CO2

Net primary production and the flux of dry matter and nutrients from vegetation to soils has increased following four years of exposure to elevated CO₂ in a southern pine forest in NC, USA. This has increased the demand for nutrients to support enhanced rates of NPP and altered the conditions for litter decomposition on the forest floor. We quantified the chemistry and decomposition dynamics of leaf litter produced by five of the most abundant tree species in this ecosystem during the third and fourth growing seasons under elevated CO₂. The objectives of this study were to determine (i) if there were systemic or species‐specific changes in leaf litter chemistry associated with a sustained enhancement of plant growth under elevated CO₂; and (ii) whether the process of litter decomposition was altered by increased inputs of energy and nutrients to the forest floor in the plots under elevated CO₂. Leaf litter chemistry, including various C fractions and N concentration, was virtually unchanged by elevated CO₂. With few exceptions, plant litter produced under elevated CO₂ lost mass or N at the same relative rate as that produced under ambient CO₂. The relationship between initial litter chemistry and decomposition was not altered by elevated CO₂. The greater forest floor mass and nutrient content in the plots under elevated CO₂ had no consistent or long‐term effect on litter decomposition. Thus, we found no evidence that plant and microbial processes under elevated CO₂ resulted in systemic changes in mass loss or N dynamics during decomposition. In contrast to the limited effects of elevated CO₂ on litter chemistry and decomposition, there were large differences among species in initial litter chemistry, mass loss and N dynamics during decomposition. If the species composition of this forest community is altered by elevated CO₂, the indirect effect of a change in species composition will exert greater control over the long‐term rate of nutrient cycling than the direct effect of elevated CO₂ on litter chemistry and decomposition dynamics alone.     

Diverse mechanisms for CO2 effects on grassland litter decomposition

The ongoing increase in atmospheric CO₂ concentration ([CO₂]) can potentially alter litter decomposition rates by changing: (i) the litter quality of individual species, (ii) allocation patterns of individual species, (iii) the species composition of ecosystems (which could alter ecosystem‐level litter quality and allocation), (iv) patterns of soil moisture, and (v) the composition and size of microbial communities. To determine the relative importance of these mechanisms in a California annual grassland, we created four mixtures of litter that differed in species composition (the annual legume Lotus wrangelianus Fischer & C. Meyer comprised either 10% or 40% of the initial mass) and atmospheric [CO₂] during growth (ambient or double‐ambient). These mixtures decomposed for 33 weeks at three positions (above, on, and below the soil surface) in four types of grassland microcosms (fertilized and unfertilized microcosms exposed to elevated or ambient [CO₂]) and at a common field site. Initially, legume‐rich litter mixtures had higher nitrogen concentrations ([N]) than legume‐poor mixtures. In most positions and environments, the different litter mixtures decomposed at approximately the same rate. Fertilization and CO₂ enrichment of microcosms had no effect on mass loss of litter within them. However, mass loss was strongly related to litter position in both microcosms and the field. Nitrogen dynamics of litter were significantly related to the initial [N] of litter on the soil surface, but not in other positions. We conclude that changes in allocation patterns and species composition are likely to be the dominant mechanisms through which ecosystem‐level decomposition rates respond to increasing atmospheric [CO₂].     

Interannual climatic variation mediates elevated CO2 and O3 effects on forest growth

We analyzed growth data from model aspen (Populus tremuloides Michx.) forest ecosystems grown in elevated atmospheric carbon dioxide ([CO₂]; 518 μL L^?1) and ozone concentrations ([O₃]; 1.5 × background of 30–40 nL L^?1 during daylight hours) for 7 years using free‐air CO₂ enrichment technology to determine how interannual variability in present‐day climate might affect growth responses to either gas. We also tested whether growth effects of those gasses were sustained over time. Elevated [CO₂] increased tree heights, diameters, and main stem volumes by 11%, 16%, and 20%, respectively, whereas elevated ozone [O₃] decreased them by 11%, 8%, and 29%, respectively. Responses similar to these were found for stand volume and basal area. There were no growth responses to the combination of elevated [CO₂+O₃]. The elevated [CO₂] growth stimulation was found to be decreasing, but relative growth rates varied considerably from year to year. Neither the variation in annual relative growth rates nor the apparent decline in CO₂ growth response could be explained in terms of nitrogen or water limitations. Instead, growth responses to elevated [CO₂] and [O₃] interacted strongly with present‐day interannual variability in climatic conditions. The amount of photosynthetically active radiation and temperature during specific times of the year coinciding with growth phenology explained 20–63% of the annual variation in growth response to elevated [CO₂] and [O₃]. Years with higher photosynthetic photon flux (PPF) during the month of July resulted in more positive growth responses to elevated [CO₂] and more negative growth responses to elevated [O₃]. Mean daily temperatures during the month of October affected growth in a similar fashion the following year. These results indicate that a several‐year trend of increasingly cloudy summers and cool autumns were responsible for the decrease in CO₂ growth response.     

Soil CO2 efflux in a boreal pine forest under atmospheric CO2 enrichment and air warming

The response of forest soil CO₂ efflux to the elevation of two climatic factors, the atmospheric concentration of CO₂ (↑CO₂ of 700 μmol mol⁻¹) and air temperature (↑ T with average annual increase of 5°C), and their combination (↑CO₂+↑ T ) was investigated in a 4-year, full-factorial field experiment consisting of closed chambers built around 20-year-old Scots pines ( Pinus sylvestris L.) in the boreal zone of Finland. Mean soil CO₂ efflux in May–October increased with elevated CO₂ by 23–37%, with elevated temperature by 27–43%, and with the combined treatment by 35–59%. Temperature elevation was a significant factor in the combined 4-year efflux data, whereas the effect of elevated CO₂ was not as evident. Elevated temperature had the most pronounced impact early and late in the season, while the influence of elevated CO₂ alone was especially notable late in the season. Needle area was found to be a significant predictor of soil CO₂ efflux, particularly in August, a month of high root growth, thus supporting the assumption of a close link between whole-tree physiology and soil CO₂ emissions. The decrease in the temperature sensitivity of soil CO₂ efflux observed in the elevated temperature treatments in the second year nevertheless suggests the existence of soil response mechanisms that may be independent of the assimilating component of the forest ecosystem. In conclusion, elevated atmospheric CO₂ and air temperature consistently increased forest soil CO₂ efflux over the 4-year period, their combined effect being additive, with no apparent interaction.     

Aphid individual performance may not predict population responses to elevated CO2 or O3

Changes in atmospheric composition affect plant quality and herbivore performance. We used the Aspen Free Air CO₂ Enrichment (FACE) facility to investigate the impacts of elevated carbon dioxide (CO₂) and ozone (O₃) on the performance of the aphid Cepegillettea betulaefoliae Granovsky feeding on paper birch (Betula papyrifera Marsh.). In Year 1, we simultaneously measured individual performance and population growth rates, and in Year 2 we surveyed natural aphid, predator and parasitoid populations throughout the growing season. Aphid growth and development (relative growth rate (RGR), development time, adult weight, embryo number and the birth weight of newborn nymphs) were unaffected by CO₂ and O₃. Aphid fecundity decreased on trees grown at elevated CO₂, O₃ and CO₂+O₃. Neither nymphal performance nor adult size were reliable indicators of future fecundity at elevated CO₂ and/or O₃. Aphid populations protected from natural enemies were unaffected by elevated CO₂, but increased significantly at elevated O₃. Individual fecundity in elevated CO₂ and O₃ atmospheres did not predict population growth rates, probably because of changes in the strength of intraspecific competition or the ability of the aphids to induce nutrient sinks. Natural aphid, predator and parasitoids populations (Year 2) showed few significant responses to CO₂ and O₃, although CO₂ and O₃ did affect the timing of aphid and natural enemy peak abundance. Elevated CO₂ and O₃ affected aphid and natural enemy populations independently: no CO₂× O₃ interactions were observed. We conclude that: (1) aphid individual performance did not predict population responses to CO₂ and O₃ and (2) elevated CO₂ and O₃ atmospheres are unlikely to affect C. betulaefoliae populations in the presence of natural enemy communities.     

The impact of elevated CO2 on yield loss from a C3 and C4 weed in field-grown soybean

Soybean (Glycine max) was grown at ambient and enhanced carbon dioxide (CO₂, + 250 μL L^?1 above ambient) with and without the presence of a C3 weed (lambsquarters, Chenopodium album L.) and a C4 weed (redroot pigweed, Amaranthus retroflexus L.), in order to evaluate the impact of rising atmospheric carbon dioxide concentration [CO₂] on crop production losses due to weeds. Weeds of a given species were sown at a density of two per metre of row. A significant reduction in soybean seed yield was observed with either weed species relative to the weed‐free control at either [CO₂]. However, for lambsquarters the reduction in soybean seed yield relative to the weed‐free condition increased from 28 to 39% as CO₂ increased, with a 65% increase in the average dry weight of lambsquarters at enhanced [CO₂]. Conversely, for pigweed, soybean seed yield losses diminished with increasing [CO₂] from 45 to 30%, with no change in the average dry weight of pigweed. In a weed‐free environment, elevated [CO₂] resulted in a significant increase in vegetative dry weight and seed yield at maturity for soybean (33 and 24%, respectively) compared to the ambient CO₂ condition. Interestingly, the presence of either weed negated the ability of soybean to respond either vegetatively or reproductively to enhanced [CO₂]. Results from this experiment suggest: (i) that rising [CO₂] could alter current yield losses associated with competition from weeds; and (ii) that weed control will be crucial in realizing any potential increase in economic yield of agronomic crops such as soybean as atmospheric [CO₂] increases.     

Effects of elevated CO2 and temperature-grown red and sugar maple on gypsy moth performance

Few studies have investigated how tree species grown under elevated CO₂ and elevated temperature alter the performance of leaf‐feeding insects. The indirect effects of an elevated CO₂ concentration and temperature on leaf phytochemistry, along with potential direct effects on insect growth and consumption, may independently or interactively affect insects. To investigate this, we bagged larvae of the gypsy moth on leaves of red and sugar maple growing in open‐top chambers in four CO₂/temperature treatment combinations: (i) ambient temperature, ambient CO₂; (ii) ambient temperature, elevated CO₂ (+ 300 μL L^?1 CO₂); (iii) elevated temperature (+ 3.5°C), ambient CO₂; and (iv) elevated temperature, elevated CO₂. For both tree species, leaves grown at elevated CO₂ concentration were significantly reduced in leaf nitrogen concentration and increased in C: N ratio, while neither temperature nor its interaction with CO₂ concentration had any effect. Depending on the tree species, leaf water content declined (red maple) and carbon‐based phenolics increased (sugar maple) on plants grown in an enriched CO₂ atmosphere. The only observed effect of elevated temperature on leaf phytochemistry was a reduction in leaf water content of sugar maple leaves. Gypsy moth larval responses were dependent on tree species. Larvae feeding on elevated CO₂‐grown red maple leaves had reduced growth, while temperature had no effect on the growth or consumption of larvae. No significant effects of either temperature or CO₂ concentration were observed for larvae feeding on sugar maple leaves. Our data demonstrate strong effects of CO₂ enrichment on leaf phytochemical constituents important to folivorous insects, while an elevated temperature largely has little effect. We conclude that alterations in leaf chemistry due to an elevated CO₂ atmosphere are more important in this plant–folivorous insect system than either the direct short‐term effects of temperature on insect performance or its indirect effects on leaf chemistry.     

Elevated CO2 alters birch resistance to Lagomorpha herbivores

We studied the three‐way interaction of elevated CO₂, nitrogen (N), and temperature (T), and the two‐way interaction of elevated CO₂ and early‐season defoliation on the secondary chemistry and resistance of Eurasian silver birch (Betula pendula) and North American paper birch (B. papyrifera) against the Eurasian hare (Lepus timidus) and the North American eastern cottontail rabbit (Sylvilagus floridanus), respectively. Elevated CO₂ decreased the palatability of winter‐dormant silver and paper birch stems to both hares and rabbits, respectively. But the effect on hares was only apparent at intermediate levels of N fertilization. Elevated T had no effect on palatability. The effects of elevated CO₂, N, and T on levels of silver birch bark phenolics and terpenoids were dominated by two‐way interactions between N and CO₂, and N and T. Generally, however, N amendments elicited a parabolic response in carbon partitioning to most biosynthetic classes of silver birch phenolics (i.e. highest concentrations occurring at intermediate N). CO₂ elevation was most enhancing at highest levels of N. On the other hand, T increases, more often than not, elicited reductions in phenolics, but especially so at the highest N level. In the case of B. papyrifera, elevated CO₂ increased carbon partitioning to Folin‐Denis stem and branch phenolics and condensed tannins. Early‐season defoliation, on the other hand, had no effect on phenolics and tannins but lowered both N and energy levels of branches. Elevated CO₂ substantially ameliorated the negative effects of severe defoliation on tree growth. These results support the hypothesis that continuing anthropogenic alterations of the atmosphere may trigger significant changes in plant phenotypic resistance to mammalian herbivores owing to an increasing net carbon balance between the highly vagile supply and demand capacities of plant carbon sources and sinks.     

The influence of elevated CO2 and O3 on fine roots and mycorrhizas of naturally growing young Scots pine trees during three exposure years

Young Scots pine trees naturally established at a pine heath were exposed to two concentrations of CO₂ (ambient and doubled ambient) and two O₃ regimes (ambient and doubled ambient) and their combination in open-top field chambers during growing seasons 1994, 1995 and 1996 (late May to 15 September). Filtered ozone treatment and chamberless control trees were also included in the treatment comparisons. Root ingrowth cores were inserted to the undisturbed soil below the branch projection of each tree at the beginning of the fumigation period in 1994 and were harvested at the end of the fumigation periods in 1995 and 1996. Root biomasses were determined from different soil layers in the ingrowth cores, and the infection levels of different mycorrhizal types were calculated. Elevated O₃ and CO₂ did not have significant effects on the biomass production of Scots pine coarse (Ø > 2 mm) or fine roots (Ø < 2 mm) and roots of grasses and dwarf shrubs. Elevated O₃ caused a transient stimulation, observable in 1995, in the proportion of tuber-like mycorrhizas, total mycorrhizas and total short roots but this stimulation disappeared during the last study year. Elevated CO₂ did not enhance carbon allocation to root growth or mycorrhiza formation, although a diminishing trend in the mycorrhiza formation was observed. In the combination treatment increased CO₂ inhibited the transient stimulating effect of ozone, and a significant increase of old mycorrhizas was observed. Our conclusion is that doubled CO₂ is not able to increase carbon allocation to growth of fine roots or mycorrhizas in nutrient poor forest sites and realistically elevated ozone does not cause a measurable limitation to roots within a period of three exposure years.