Individual growth rates do not predict aphid population densities under altered atmospheric conditions

• 1 Altered atmospheric composition, associated with climate change, can modify herbivore population dynamics through CO₂ and/or O₃‐mediated changes in plant quality.
• 2 Although pea aphid Acyrthosiphon pisum genotypes exhibit intraspecific variation in population growth in response to atmospheric composition, the proximate mechanisms underlying this variation are largely unknown.
• 3 By rearing single (green, pink) and mixed (green + pink) pea aphid genotypes on red clover Trifolium pratense at the Aspen Free Air CO₂ and O₃ Enrichment (Aspen FACE) site, we assessed whether: (i) elevated CO₂ and/or O₃ concentrations alter aphid growth and development and (ii) individual aphid growth rates predict aphid population densities.
• 4 We showed that growth and development of individual green and pink aphids were not influenced by CO₂ and/or O₃ concentrations when reared as individual or mixed genotypes. Individual growth rates, however, did not predict population densities.
• 5 Reared as a single genotype, green pea aphid populations decreased in response to elevated CO₂ concentrations, but not in response to elevated CO₂ + O₃ concentrations. Pink pea aphid populations reared as a single genotype were unaffected by augmented CO₂ or O₃. Populations of mixed genotypes, however, were reduced under elevated CO₂ concentrations, irrespective of O₃ concentrations.
• 6 Herbivore population sizes may not readily be predicted from growth rates of individual organisms under atmospheric conditions associated with global climate change.

     

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.     

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.     

CO2 and O3 effects on host plant preferences of the forest tent caterpillar (Malacosoma disstria)

Elevated levels of CO₂ and O₃ affect plant growth and phytochemistry, which in turn can alter physiological performance of associated herbivores. Little is known, however, about how generalist insect herbivores respond behaviorally to CO₂‐ and O₃‐mediated changes in their host plants. This research examined the effects of elevated CO₂ and O₃ levels on host plant preferences and consumption of forest tent caterpillar (FTC, Malacosoma disstria Hbn.) larvae. Dual choice feeding assays were performed with foliage from birch (Betula papyrifera Marsh.) and aspen (Populus tremuloides Michx., genotypes 216 and 259). Trees were grown at the Aspen Free Air CO₂ Enrichment (FACE) facility near Rhinelander, WI, USA, and had been exposed to ambient or elevated concentrations of CO₂ and/or O₃. Levels of nutritional and secondary compounds were quantified through phytochemical analyses. The results showed that elevated O₃ levels increased FTC larval preferences for birch compared with aspen, whereas elevated CO₂ levels had the opposite effect. In assays with the two aspen genotypes, addition of both CO₂ and O₃ caused a shift in feeding preferences from genotype 259 to genotype 216. Consumption was unaffected by experimental treatments in assays comparing aspen and birch, but were increased for larvae given high O₃ foliage in the aspen genotype assays. Elevated levels of CO₂ and O₃ altered tree phytochemistry, but did not explain shifts in feeding preferences. The results demonstrate that increased levels of CO₂ and O₃ can alter insect host plant preferences both between and within tree species. Also, consequences of altered host quality (e.g., compensatory consumption) may be buffered by partial host shifts in situations when alternative plant species are available. Environmentally induced changes in host plant preferences may have the potential to alter the distribution of herbivory across plant genotypes and species, as well as competitive interactions among them.     

Transgenerational phenotypic plasticity under future atmospheric conditions

Organisms often exhibit transgenerational phenotypic changes in response to an increased risk of parasitism or predation. Shifts in global atmospheric composition could modify these phenotypic effects through changes in either nutrient quantity/quality or altered interactions with higher trophic levels. Here we show that future atmospheric conditions alter a natural enemy‐induced wing polyphenism in aphids. Winged offspring production by Uroleucon nigrotuberculatum aphids on goldenrod (Solidago canadensis var. scabra) does not differ in enriched CO₂ and/or O₃ atmospheres. However, proportionally more winged offspring are produced in response to search cues from both coccinellid predators (Coccinella septempunctata) and hymenopteran parasitoids (Aphidius polygonaphis) relative to plants not searched by natural enemies. Moreover, the magnitude of this response differs under enriched CO₂ and O₃ environments. Aphids produce more winged offspring in response to predators under elevated CO₂, but produce more winged offspring in response to parasitoids under elevated O₃. Thus, global atmospheric changes influence natural enemy‐mediated phenotypic expression, with potentially far‐reaching consequences for trophic dynamics.     

Insects and fungi on a C3 sedge and a C4 grass exposed to elevated atmospheric CO2 concentrations in open-top chambers in the field

The effects of elevated atmospheric CO₂ concentration on plant-fungi and plant-insect interactions were studied in an emergent marsh in the Chesapeake Bay. Stands of the C₃ sedge Scirpus olneyi Grey, and the C₄ grass Spartina patens (Ait.) Muhl. have been exposed to elevated atmospheric CO₂ concentrations during each growing season since 1987. In August 1991 the severities of fungal infections and insect infestations were quantified. Shoot nitrogen concentration ([N]) and water content (WC) were determined. In elevated concentrations of atmospheric CO₂, 32% fewer S. olneyi plants were infested by insects, and there was a 37% reduction in the severity of a pathogenic fungal infection, compared with plants grown in ambient CO₂ concentrations. S. olneyi also had reduced [N], which correlated positively with the severities of fungal infections and insect infestations. Conversely, S. patens had increased WC but unchanged [N] in elevated concentrations of atmospheric CO₂ and the severity of fungal infection increased. Elevated atmospheric CO₂ concentration increased or decreased the severity of fungal infection depending on at least two interacting factors, [N] and WC; but it did not change the number of plants that were infected with fungi. In contrast, the major results for insects were that the number of plants infected with insects decreased, and that the amount of tissue that each insect ate also decreased.     

Elevated CO2 decreases seed germination in Arabidopsis thaliana

The impact of elevated [CO₂] on seed germination was studied in different genotypes of Arabidopsis thaliana from natural populations. Two generations of seeds were studied: the maternal generation was produced in the greenhouse (present-day conditions), the offspring generation was produced in two chambers where the CO₂ concentration was either the present atmospheric concentration (about 350 ppm) or elevated (700 ppm). The seeds were tested for proportion of germinated seeds and mean germination time in both chambers to study the impact of elevated [CO₂] during seed production and germination. Elevated [CO₂] during maturation of seeds on the mother-plants decreased the proportion of germinated seeds, while elevated [CO₂] during germination had no effect on the proportion of germinated seeds. However, when seeds were both produced and germinated under elevated [CO₂] (situation expected by the end of next century), germination was slow and low. Moreover, the effect of the [CO₂] treatment differs among genotypes of Arabidopsis: there is a strong treatment × genotype interaction. This means that there is ample genetic variance for a selective response modiying the effects of high levels of [CO₂] in natural populations of Arabidopsis thaliana. The outcome at the community level will depend on what seeds are available, when they germinate and the resulting competition following germination.     

Dependence of the Post-Illumination Burst of CO2 on Temperature, Light, CO2, and O2 Concentration in Wheat (Triticum aestivum)

The effect of environmental factors on the post-illumination burst of CO₂ (PIB) and O₂ inhibition of apparent photosynthesis (APS) in wheat (Triticum aestivum L.) was studied in an open gas exchange system utilizing the mathematics of non-steady-state systems. Two components of inhibition by O₂ are suggested: one is caused by photorespiration as measured from the maximum rate of the PIB, and the second is direct inhibition as taken as APS_2%O2— (APS_x%O2+ PIB_x%O2) where X is the oxygen concentration. A primary PIB which occurred from 16–28 s after the darkening of the foliage was attributed to photorespiration. No primary PIB was observed at 2% O₂. At a CO₂ concentration of 100 μ/1 in the atmosphere (about 2.5 μM based on leaf intercellular concentration) and at 30°C and 145 nE/cm² nE/cm²·s, APS decreased curve-linearly with increasing O₂ and reached an O₂ compensation point of 560 μM (48% by volume), above which there was a net loss of CO₂ in the light. The PIB increased with increasing O₂ and became saturated at about 500 μM O₂ but decreased above 900 μM O₂. Direct inhibition of photosynthesis by O₂ increased with increasing O₂ concentration. Decreasing CO₂ concentration had an effect on the magnitude of the PIB similar to that of increasing O₂. At 30°C and 21% O₂, the PIB increased with decreasing CO₂ down to the CO₂ compensation point (I) of 1.4 μM (47 μM/l). Below Γ, both PIB and CO₂ evolution into the air in the light (at 21% O₂) increased and then decreased at CO₂ below 0.8 μM. The ratio of the PIB to APS_{2% o} O₂ increased linearly with increasing O₂/CO₂ ratio where O₂ was held constant at 21% and CO₂ was varied from 1.4 to 8.5 μM, while direct inhibition of photosynthesis expressed as a proportion of APS_2%O2 remained constant over this range. At low CO₂ concentration photorespiration as estimated by the PIB is the major part of O₂ photosynthesis, while at atmospheric CO₂ levels, direct inhibition is the major component. The PIB and APS at 2% and 21% O₂ increased hyperbolically with increasing irradiance and all became light-saturated at about 65 nE/cm₂ s. The percentage total O₂ inhibition of photosynthesis remained constant with increasing irradiance as did the relative contribution of direct O₂ inhibition or photorespiration (PIB) to total O₂ inhibition. The PIB and APS at 21% O₂ had similar temperature optima of 30°C when experimental conditions were adjusted to provide a constant internal O₂/CO₂ solubility ratio at varying temperatures. However, with a constant external CO₂ concentration, the temperature optimum for the PIB shifted upward to 35°C while that for APS at 21% O₂ remained at 30°C, which may be due to an increased O₂/CO₂ concentration in the leaf with increasing temperature.     

Soil CO2 efflux of two silver birch clones exposed to elevated CO2 and O3 levels during three growing seasons

In the present open‐top chamber experiment, two silver birch clones (Betula pendula Roth, clone 4 and clone 80) were exposed to elevated levels of carbon dioxide (CO₂) and ozone (O₃), singly and in combination, and soil CO₂ efflux was measured 14 times during three consecutive growing seasons (1999–2001). In the beginning of the experiment, all experimental trees were 7 years old and during the experiment the trees were growing in sandy field soil and fertilized regularly. In general, elevated O₃ caused soil CO₂ efflux stimulation during most measurement days and this stimulation enhanced towards the end of the experiment. The overall soil respiration response to CO₂ was dependent on the genotype, as the soil CO₂ efflux below clone 80 trees was enhanced and below clone 4 trees was decreased under elevated CO₂ treatments. Like the O₃ impact, this clonal difference in soil respiration response to CO₂ increased as the experiment progressed. Although the O₃ impact did not differ significantly between clones, a significant time × clone × CO₂× O₃ interaction revealed that the O₃‐induced stimulation of soil respiration was counteracted by elevated CO₂ in clone 4 on most measurement days, whereas in clone 80, the effect of elevated CO₂ and O₃ in combination was almost constantly additive during the 3‐year experiment. Altogether, the root or above‐ground biomass results were only partly parallel with the observed soil CO₂ efflux responses. In conclusion, our data show that O₃ impacts may appear first in the below‐ground processes and that relatively long‐term O₃ exposure had a cumulative effect on soil CO₂ efflux. Although the soil respiration response to elevated CO₂ depended on the tree genotype as a result of which the O₃ stress response might vary considerably within a single tree species under elevated CO₂, the present experiment nonetheless indicates that O₃ stress is a significant factor affecting the carbon cycling in northern forest ecosystems.     

Growth, light interception and yield responses of spring wheat (Triticum aestivum L.) grown under elevated CO2 and O3 in open-top chambers

Spring wheat cv. Minaret was grown to maturity under three carbon dioxide (CO₂) and two ozone (O₃) concentrations in open-top chambers (OTC). Green leaf area index (LAI) was increased by elevated CO₂ under ambient O₃ conditions as a direct result of increases in tillering, rather than individual leaf areas. Yellow LAI was also greater in the 550 and 680 μmol mol^–1 CO₂ treatments than in the chambered ambient control; individual leaves on the main shoot senesced more rapidly under 550 μmol mol^–1 CO₂, but senescence was delayed at 680 μmol mol^–1 CO₂. Fractional light interception (f) during the vegetative period was up to 26% greater under 680 μmol mol^–1 CO₂ than in the control treatment, but seasonal accumulated intercepted radiation was only increased by 8%. As a result of greater carbon assimilation during canopy development, plants grown under elevated CO₂ were taller at anthesis and stem and ear biomass were 27 and 16% greater than in control plants. At maturity, yield was 30% greater in the 680 μmol mol^–1 CO₂ treatment, due to a combination of increases in the number of ears per m^–2, grain number per ear and individual grain weight (IGW). Exposure to a seasonal mean (7 h d^–1) of 84 nmol mol^–1 O₃ under ambient CO₂ decreased green LAI and increased yellow LAI, thereby reducing both f and accumulated intercepted radiation by ≈ 16%. Individual leaves senesced completely 7–28 days earlier than in control plants. At anthesis, the plants were shorter than controls and exhibited reductions in stem and ear biomass of 15 and 23%. Grain yield at maturity was decreased by 30% due to a combination of reductions in ear number m^–2, the numbers of grains per spikelet and per ear and IGW. The presence of elevated CO₂ reduced the rate of O₃-induced leaf senescence and resulted in the maintenance of a higher green LAI during vegetative growth under ambient CO₂ conditions. Grain yields at maturity were nevertheless lower than those obtained in the corresponding elevated CO₂ treatments in the absence of elevated O₃. Thus, although the presence of elevated CO₂ reduced the damaging impact of ozone on radiation interception and vegetative growth, substantial yield losses were nevertheless induced. These data suggest that spring wheat may be susceptible to O₃-induced injury during anthesis irrespective of the atmospheric CO₂ concentration. Possible deleterious mechanisms operating through effects on pollen viability, seed set and the duration of grain filling are discussed.     

Competition increasingly dominates the responsiveness of juvenile beech and spruce to elevated CO2 and/or O3 concentrations throughout two subsequent growing seasons

Saplings of Fagus sylvatica and Picea abies were grown in mono‐ and mixed cultures in a 2‐year phytotron study under all four combinations of ambient and elevated ozone (O₃) and carbon dioxide (CO₂) concentrations. The hypotheses tested were (1) that the competitiveness of beech rather than spruce is negatively affected by the exposure to enhanced O₃ concentrations, (2) spruce benefits from the increase of resource availability (elevated CO₂) in the mixed culture and (3) that the responsiveness of plants to CO₂ and O₃ depends on the type of competition (i.e. intra vs. interspecific). Beech displayed a competitive disadvantage when growing in mixture with spruce: after two growing seasons under interspecific competition, beech showed significant reductions in leaf gas exchange, biomass development and crown volume as compared with beech plants growing in monoculture. In competition with spruce, beech appeared to be nitrogen (N)‐limited, whereas spruce tended to benefit in terms of its plant N status. The responsiveness of the juvenile trees to the atmospheric treatments differed between species and was dominated by the type of competition: spruce growth benefited from elevated CO₂ concentrations, while beech growth suffered from the enhanced O₃ regime. In general, interspecific competition enhanced these atmospheric treatment effects, supporting our hypotheses. Significant differences in root : shoot biomass ratio between the type of competition under both elevated O₃ and CO₂ were not caused by readjustments of biomass partitioning, but were dependent on tree size. Our study stresses that competition is an important factor driving plant development, and suggests that the knowledge about responses of plants to elevated CO₂ and/or O₃, acquired from plants growing in monoculture, may not be transferred to plants grown under interspecific competition as typically found in the field.     

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.     

Responses of trembling aspen (Populus tremuloides) phytochemistry and aspen blotch leafminer (Phyllonorycter tremuloidiella) performance to elevated levels of atmospheric CO2 and O3

1 This research was conducted at the Aspen FACE (Free Air CO₂ Enrichment) site located in northern Wisconsin, U.S.A. where trembling aspen (Populus tremuloides Michaux) trees were exposed to one of four atmospheric treatments: elevated carbon dioxide (CO₂; 560 µL/L), elevated ozone (O₃; ambient × 1.5), elevated CO₂ and O₃, or ambient air. We evaluated the effects of these fumigants on aspen foliar quality and the performance of aspen blotch leafminer (Phyllonorycter tremuloidiella Braun). 2 CO₂ and O₃ each affected foliar quality, with the major changes consisting of an 11% reduction in nitrogen under elevated CO₂ and a 20% reduction in tremulacin under elevated O₃. In the CO₂ + O₃ treatment, nitrogen levels were reduced by 15% and CO₂ ameliorated the O₃‐mediated reduction in tremulacin levels. 3 Phyllonorycter tremuloidiella were allowed to colonize trees naturally. Elevated CO₂ and O₃ reduced colonization rates by 42 and 49% relative to ambient CO₂ and O₃, respectively. The only effect of fumigation treatments on larval performance occurred under elevated O₃, where male development time and larval consumption increased by 8 and 28%, respectively, over insects reared under ambient O₃. 4 These data demonstrate that the individual and combined effects of CO₂ and O₃ can alter aspen foliar chemistry and that these alterations in foliar chemistry produce little to no change in larval performance. However, both CO₂ and O₃ greatly reduced oviposition. In order to ascertain the full effects of CO₂ and O₃ on insect performance, future studies should address both population‐ and individual‐level characteristics.     

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.     

Natural CO2 springs in Italy: a resource for examining long-term response of vegetation to rising atmospheric CO2 concentrations

It is estimated that more than 100 geothermal CO₂ springs exist in central-western Italy. Eight springs were selected in which the atmospheric CO₂ concentrations were consistently observed to be above the current atmospheric average of 354μmol mol^-1. CO₂ concentration measurements at some of the springs are reported. The springs are described, and their major topographic and vegetational features are reported. Preliminary observations made on natural vegetation growing around the gas vents are then illustrated. An azonal pattern of vegetation distribution occurs around every CO₂ spring regardless of soil type and phytoclimatic areas. This is composed of pioneer populations of a Northern Eurasiatic species (Agrostis canina L.) which is often associated with Scirpus lacustris L. The potential of these sites for studying the long-term response of vegetation to rising atmospheric CO₂ concentrations is discussed.     

C4 characteristics of photosynthesis in the C3 alga Hydrodictyon africanum

Abstract Results obtained with Hydrodictyon africanum, and data from the literature, show that most green algae of the chlorophyte type (e.g. Chlorella, Chlamydomonas, Hydrodictyon) differ in their photosynthetic C fixation characteristics from most green algae of the charophyte type (e.g. Spirogyra, Chara) and from C₃ higher plants. The chlorophyte algae fix inorganic carbon by the photosynthetic carbon reduction cycle pathway, but have a low CO₂ compensation point in 250 μM O₂, a low inhibition of CO₂ fixation from 10 μM CO₂/250 μM O₂ when compared with 10 μM CO₂/zero O₂, and a low half-saturation constant for CO₂. These three characteristics are different from those of charophytes and C₃ higher plants, and resemble those of C₄ higher plants. It is suggested that these characteristics of chlorophyte algae are the result of a ‘CO₂ concentrating mechanism’ which increases the CO₂/O₂ ratio at the site of ribulose bisphosphate carboxylase-oxygenase action in a similar way to that achieved by the C₄?C₃ acid cycle in C₄ plants. In the chlorophyte algae, however, CO₂ concentration probably involves active HCO₃^? transport at the inner membrane of the chloroplast envelope. Active HCO₃^? transport can occur at the plasmalemma of charophyte algae and submerged aquatic higher plants as well as chlorophyte algae, so it is unlikely to explain the differences between the two groups of aquatic green plants. Differences in the properties of ribulose bisphosphate carboxylase-oxygenase, and differences in CO₂ production in the light, also seem inadequate to account for the different photosynthetic characteristics. The chlorophyte type of ‘C0₂ concentrating mechanism’ appears to be common in other classes of eukaryotic algae, and in cyanophytes. Some of the ‘advanced’ members of these eukaryotic algal classes (including the chlorophytes) may lack the mechanism, while some ‘primitive’ charophytes may retain the mechanism which their ancestors presumably possessed.     

Belowground competition and the response of developing forest communities to atmospheric CO2 and O3

As human activity continues to increase CO₂ and O₃, broad expanses of north temperate forests will be simultaneously exposed to elevated concentrations of these trace gases. Although both CO₂ and O₃ are potent modifiers of plant growth, we do not understand the extent to which they alter competition for limiting soil nutrients, like nitrogen (N). We quantified the acquisition of soil N in two 8‐year‐old communities composed of trembling aspen genotypes (n= 5) and trembling aspen–paper birch which were exposed to factorial combinations of CO₂ (ambient and 560 μL L⁻¹) and O₃ (ambient = 30–40 vs. 50–60 nL L⁻¹). Tracer amount of ¹⁵NH₄⁺ were applied to soil to determine how these trace gases altered the competitive ability of genotypes and species to acquire soil N. One year after isotope addition, we assessed N acquisition by measuring the amount of ¹⁵N tracer contained in the plant canopy (i.e. recent N acquisition), as well as the total amount of canopy N (i.e. cumulative N acquisition). Exposure to elevated CO₂ differentially altered recent and cumulative N acquisition among aspen genotypes, changing the rank order in which they obtained soil N. Elevated O₃ also altered the rank order in which aspen genotypes obtained soil N by eliciting increases, decreases and no response among genotypes. If aspen genotypes respond similarly under field conditions, then rising concentrations of CO₂ and O₃ could alter the structure of aspen populations. In the aspen–birch community, elevated CO₂ increased recent N (i.e. ¹⁵N) acquisition in birch (68%) to a greater extent than aspen (19%), suggesting that, over the course of this experiment, birch had gained a competitive advantage over aspen. The response of genotypes and species to rising CO₂ and O₃ concentrations, and how these responses are modified by competitive interactions, has the potential to change the future composition and productivity of northern temperate forests.     

Response of quaking aspen genotypes to enriched CO2: foliar chemistry and tussock moth performance

Abstract

• 1 Genetic variation in the phytochemical responses of plants to CO₂ enrichment is likely to alter trophic dynamics, and to shift intraspecific selection pressures on plant populations. We evaluated the independent and interactive effects of atmospheric CO₂ and quaking aspen (Populus tremuloides Michx.) genotype on chemical composition of foliage and performance of the whitemarked tussock moth (Orgyia leucostigma J. E. Sm.).
• 2 This research was conducted at the Aspen FACE (Free Air CO₂ Enrichment) site in northern Wisconsin, U.S.A. Leaf samples were collected periodically from each of three genetically variable aspen genotypes growing under ambient and elevated CO₂, and analysed for levels of primary and secondary metabolites. Tussock moth larvae were reared in situ on experimental trees, and development times and pupal masses were recorded.
• 3 Foliar chemical composition varied among aspen genotypes and in response to CO₂ enrichment. However, chemical responses of trees to elevated CO₂ were generally consistent across genotypes.
• 4 Larval development times varied among host genotypes and increased slightly for insects on high‐CO₂ plants. Enriched CO₂ tended to reduce insect pupal masses, particularly for females on one of the three aspen genotypes.
• 5 CO₂ × genotype interactions observed for plant chemistry and insect performance in this study with a small number of genotypes are probably too few, and too weak, to shift selection pressures in aspen populations. These results differ, however, from earlier work in which more substantial CO₂ × genotype interactions were observed for plant chemistry.

     

Soil nitrogen transformations under Populus tremuloides, Betula papyrifera and Acer saccharum following 3 years exposure to elevated CO2 and O3

Increases in atmospheric CO₂ and tropospheric O₃ may affect forest N cycling by altering plant litter production and the availability of substrates for microbial metabolism. Three years following the establishment of our free‐air CO₂–O₃ enrichment experiment, plant growth has been stimulated by elevated CO₂ resulting in greater substrate input to soil; elevated O₃ has counteracted this effect. We hypothesized that rates of soil N cycling would be enhanced by greater plant productivity under elevated CO₂, and that CO₂ effects would be dampened by O₃. We found that elevated CO₂ did not alter gross N transformation rates. Elevated O₃ significantly reduced gross N mineralization and microbial biomass N, and effects were consistent among species. We also observed significant interactions between CO₂ and O₃: (i) gross N mineralization was greater under elevated CO₂ (1.0 mg N kg^?1 day^?1) than in the presence of both CO₂ and O₃ (0.5 mg N kg^?1 day^?1) and (ii) gross NH₄⁺ immobilization was also greater under elevated CO₂ (0.8 mg N kg^?1 day^?1) than under CO₂ plus O₃ (0.4 mg N kg^?1 day^?1). We used a laboratory ¹⁵N tracer method to quantify transfer of inorganic N to organic pools. Elevated CO₂ led to greater recovery of NH₄⁺‐¹⁵N in microbial biomass and corresponding lower recovery in the extractable NO₃^? pool. Elevated CO₂ resulted in a substantial increase in NO₃^?‐¹⁵N recovery in soil organic matter. We observed no O₃ main effect and no CO₂ by O₃ interaction effect on ¹⁵N recovery in any soil pool. All of the above responses were most pronounced beneath Betula papyrifera and Populus tremuloides, which have grown more rapidly than Acer saccharum. Although elevated CO₂ has increased plant productivity, the resulting increase in plant litter production has yet to overcome the influence of the pre‐existing pool of soil organic matter on soil microbial activity and rates of N cycling. Ozone reduces plant litter inputs and also appears to affect the composition of plant litter in a way that reduces microbial biomass and activity.     

Atmospheric change alters foliar quality of host trees and performance of two outbreak insect species

This study examined the independent and interactive effects of elevated carbon dioxide (CO₂) and ozone (O₃) on the foliar quality of two deciduous trees species and the performance of two outbreak herbivore species. Trembling aspen (Populus tremuloides) and paper birch (Betula papyrifera) were grown at the Aspen FACE research site in northern Wisconsin, USA, under four combinations of ambient and elevated CO₂ and O₃. We measured the effects of elevated CO₂ and O₃ on aspen and birch phytochemistry and on gypsy moth (Lymantria dispar) and forest tent caterpillar (Malacosoma disstria) performance. Elevated CO₂ nominally affected foliar quality for both tree species. Elevated O₃ negatively affected aspen foliar quality, but only marginally influenced birch foliar quality. Elevated CO₂ slightly improved herbivore performance, while elevated O₃ decreased herbivore performance, and both responses were stronger on aspen than birch. Interestingly, elevated CO₂ largely offset decreased herbivore performance under elevated O₃. Nitrogen, lignin, and C:N were identified as having strong influences on herbivore performance when larvae were fed aspen, but no significant relationships were observed for insects fed birch. Our results support the notion that herbivore performance can be affected by atmospheric change through altered foliar quality, but how herbivores will respond will depend on interactions among CO₂, O₃, and tree species. An emergent finding from this study is that tree age and longevity of exposure to pollutants may influence the effects of elevated CO₂ and O₃ on plant–herbivore interactions, highlighting the need to continue long-term atmospheric change research.