Are legume‐feeding herbivores buffered against direct effects of elevated carbon dioxide on host plants? A test with the sulfur butterfly,Colias philodice

When grown under elevated atmospheric carbon dioxide (CO₂), leaf nitrogen content decreases less for legumes than for nonlegume C₃ plants. Given that elevated CO₂ adversely affects insect herbivores primarily through dilution of plant nitrogen, it is reasonable to expect that legume-feeding herbivores will be relatively buffered against CO₂-induced reduction in performance. However, despite their ecological and economic importance, very few studies have addressed the effects of elevated CO₂ on legume-feeding herbivores. Unlike the responses of the vast majority of nonlegume C₃ plants, when the legumes Trifolium pratense and Melilotus alba were grown under elevated (742 ppm) CO₂, leaf nitrogen and carbon contents and C : N ratios did not change. For Colias philodice larvae fed T. pratense , elevated CO₂ had little or no effect on consumption, digestion, or conversion of whole food or nitrogen and, consequently, no effect on growth rate, instar duration, or pupal weight. For larvae fed M. alba , elevated CO₂ had little or no effect on consumption of whole food or nitrogen, increased digestion but decreased conversion of both and, consequently, had no effect on growth rate, instar duration or pupal weight. These results suggest that, relative to herbivores of nonlegume C₃ plants, legume-feeding herbivores will be less affected as atmospheric CO₂ continues to rise.     

CO2-mediated changes in aspen chemistry: effects on gypsy moth performance and susceptibility to virus

We investigated the effects of long-term CO₂ enrichment on foliar chemistry of quaking aspen ( Populus tremuloides ) and the consequences of chemical changes for performance of the gypsy moth ( Lymantria dispar ) and susceptibility of the gypsy moth to a nucleopolyhedrosis virus (NPV). Foliage was collected from outdoor open-top chambers and fed to insects in a quarantine rearing facility. Under enriched CO₂, levels of leaf nitrogen declined marginally, levels of starch and phenolic glycosides did not change, and levels of condensed tannins increased. Long-term bioassays revealed reduced growth (especially females), prolonged development and increased consumption in larvae fed high-CO₂ foliage but no significant differences in final pupal weights or female fecundity. Short-term bioassays showed weaker, and sex-specific, effects of CO₂ treatment on larval performance. Correlation analyses revealed strong, negative associations between insect performance and phenolic glycoside concentrations, independent of CO₂ treatment. Larval susceptibility to NPV did not differ between CO₂ treatments, suggesting that effects of this natural enemy on gypsy moths are buffered from CO₂-induced changes in foliar chemistry. Our results emphasize that the impact of enriched CO₂ on plant–insect interactions will be determined not only by how concentrations of plant compounds are altered, but also by the relevance of particular compounds for insect fitness. This work also underscores the need for studies of genetic variation in plant responses to enriched CO₂ and long-term population-level responses of insects to CO₂-induced changes in host quality.     

Influence of soil O2 and CO2 on root respiration for Agave deserti

Respiration measured as CO₂ efflux was determined at various soil O₂ and CO₂ concentrations for individual, attached roots of a succulent perennial from the Sonoran Desert, Agave deserti Engelm. The respiration rate increased with increasing O₂ concentration up to about 16% O₂ for established roots and 5% O₂ for rain roots (fine branch roots on established roots induced by wetting of the soil) and then remained fairly constant up to 21% O₂. When O₂ was decreased from 21 to 0%, the respiration rates were similar to those obtained with increasing O₂ concentration. The CO₂ concentration in the root zone, which for the shallow-rooted A. deserti in the field was about 1 000 μl l^-1, did not affect root respiration at concentrations up to 2 000 μl l^-1, but higher concentrations reduced it, respiration being abolished at 20 000 μl l^-1 (2%) CO₂ for both established and rain roots. Upon lowering CO₂ to 1 000 μl l^-1 after exposure to concentrations up to 10000 μl l^-1 CO₂, inhibition of respiration was reversible. Uptake of the vital stain neutral red by root cortical cells was reduced to zero, indicating cell death, in about 4 h at 2% CO₂, substantiating the detrimental effects of high soil CO₂ concentrations on roots of A. deserti . This CO₂ response may explain why roots of desert succulents tend to occur in porous, well-aerated soils.     

Host plant effects on the performance of the aphid Aulacorthum solani (Kalt.) (Homoptera: Aphididae) at ambient and elevated CO2

In future elevated CO₂ environments, chewing insects are likely to perform less well than at present because of the effects of increased carbon fixation on their host plants. When the aphid, Aulacorthum solani was reared on bean ( Vicia faba ) and tansy ( Tanacetum vulgare ) plants under ambient and elevated CO_2, performance was enhanced on both hosts at elevated CO₂. The nature of the response was different on each plant species suggesting that feeding strategy may influence an insect's response to elevated CO₂. On bean, the daily rate of production of nymphs was increased by 16% but there was no difference in development time, whereas on tansy, development time was 10% shorter at elevated CO₂ but the rate of production of nymphs was not affected. The same aphid clone therefore responded differently to elevated CO₂ on different host plants. This increase in aphid performance could lead to larger populations of aphids in a future elevated CO₂ environment.     

Elevated CO2 and conifer roots: effects on growth, life span and turnover

Elevated CO₂ increases root growth and fine (diam. 2 mm) root growth across a range of species and experimental conditions. However, there is no clear evidence that elevated CO₂ changes the proportion of C allocated to root biomass, measured as either the root:shoot ratio or the fine root:needle ratio. Elevated CO₂ tends to increase mycorrhizal infection, colonization and the amount of extramatrical hyphae, supporting their key role in aiding the plant to more intensively exploit soil resources, providing a route for increased C sequestration. Only two studies have determined the effects of elevated CO₂ on conifer fine-root life span, and there is no clear trend. Elevated CO₂ increases the absolute fine-root turnover rates; however, the standing crop root biomass is also greater, and the effect of elevated CO₂ on relative turnover rates (turnover:biomass) ranges from an increase to a decrease. At the ecosystem level these changes could lead to increased C storage in roots. Increased fine-root production coupled with increased absolute turnover rates could also lead to increases in soil organic C as greater amounts of fine roots die and decompose. Although CO₂ can stimulate fine-root growth, it is not known if this stimulation persists over time. Modeling studies suggest that a doubling of the atmospheric CO₂ concentration initially increases biomass, but this stimulation declines with the response to elevated CO₂ because increases in assimilation are not matched by increases in nutrient supply.     

Mitigation of adverse effects of rising CO2 on a planktonic herbivore by mixed algal diets

Putative future increase in atmospheric CO₂ is expected to adversely affect herbivore growth due to decrease in contents of key nutrients such as nitrogen and phosphorus (P) relative to carbon in primary producers including plant and algal species. However, as many herbivores are polyphagous and as the response of primary producers to elevated CO₂ is highly species-specific, effects of elevated CO₂ on herbivore growth may differ between feeding conditions with monospecific and multiproducer diets. To examine this possibility, we performed CO₂ manipulation experiments under a P-limited condition with a planktonic herbivore, Daphnia , and three algal species, Scenedesmus obliquus (green algae), Cyclotella sp. (diatoms) and Synechococcus sp. (cyanobacteria). Semibatch cultures with single algal species (monocultures) and multiple algal species (mixed cultures) were grown at ambient (360 ppm) and high CO₂ levels (2000 ppm) that were within the natural range in lakes. Both in the mono- and mixed cultures, algal steady state abundance increased but algal P : C and N : C ratios decreased when they were grown at high CO₂. As expected, Daphnia fed monospecific algae cultured at high CO₂ had decreased growth rates despite increased algal abundance. However, when fed mixed algae cultured at high CO₂, especially consisting of diatoms and cyanobacteria or the three algal species, Daphnia maintained high growth rates despite lowered P and N contents relative to C in the algal diets. These results imply that algal diets composed of multiple species can mitigate the adverse effects of elevated CO₂ on herbivore performance, although the magnitude of this mitigation depends on the composition of algal species involved in the diets.     

Culicoides responses to 1-octen-3-ol and carbon dioxide in salt marshes near Sea Island, Georgia, U.S.A.

Abstract. The semiochemicals carbon dioxide (CO₂) and l-octen-3-ol (octenol) were evaluated as attractants at several release rates, alone and in combination, for the major coastal species of biting midges in Georgia: Culicoides furens, C.hollensis and C.melleus (Diptera: Ceratopogonidae). The data show that CO₂ is an effective attractant for the females of all three species, but each has a different response pattern. In contrast, octenol was an effective attractant only for C.furens , either alone or in combination with CO₂; octenol had a repellent effect on the other two species. The combination of octenol and C0₂ had a synergistic effect on the collections of C.furens.     

Effect of elevated CO2 on carbon partitioning and exudate release from Plantago lanceolata seedlings

Plantago lanceolata L. seedlings were grown in sand microcosm units over a 43‐day experimental period under two CO₂ regimes (800 or 400 µmol mol⁻¹) to investigate the effect of elevated atmospheric CO₂ concentration on carbon partitioning and exudate release. Total organic carbon (TOC) content of the collected exudate material was measured throughout the experimental period. After 42 days growth the seedlings were labelled with [¹⁴C]‐CO₂ and the fate of the label within the plant and its release by the roots monitored. Elevated CO₂ significantly (P ≤ 0.001) enhanced shoot, root and total dry matter production although the R:S ratio was unaltered, suggesting no alteration in gross carbon partitioning. The cumulative release of TOC (in mg C) over 0‐42 days was unaltered by CO₂ treatment however, when expressed as a percentage of net assimilated C, ambient‐grown plants released a significantly (P≤ 0.001) higher percentage from their roots compared to elevated CO₂‐grown plants (i.e. 8 vs 3%). The distribution of ¹⁴C‐label was markedly altered by CO₂ treatment with significantly (P≤ 0.001) greater per cent label partitioned to the roots under elevated CO₂. This indicates increased partitioning of recent assimilate below‐ground under elevated CO₂ treatment although there was no significant difference in the percentage of ¹⁴C‐label released by the roots. Comparison of plant C budgets based on ¹⁴C‐pulse‐chase methodology and TOC measurements is discussed.     

Carbon dioxide assimilation during postharvest removal of astringency from persimmon fruit

Application of anaerobic conditions with CO₂ or N₂ atmospheres to remove astringency from harvested persimmon fruit ( Diospryros kaki L. cv. Triumph), caused production of more acetaldehyde under CO₂ than under N₂, ¹⁴CO₂ applied in a 100% CO₂ atmosphere, for 48 h to astringent persimmon fruits was incorporated mainly into malate and very little into other metabolites, such as carbohydrate or amino acids. Application of malate or pyruvate to pulp discs of astringent persimmons caused an immediate rise in acetaldehyde production. The higher levels of acetaldehyde produced by whole fruits held in a CO₂ atmosphere, than by fruits held in a N₂ atmosphere, can be explained through fixation of atmospheric CO₂ into malate, leading to acetaldehyde production.     

Effect of enhanced atmospheric CO2 on mycorrhizal colonization by Glomus mosseae in Plantago lanceolata and Trifolium repens

Plantago lanceolata L. and Trifolium repens L. were grown for 16 wk in ambient (360 μmol mol⁻¹) and elevated (610 μmol mol⁻¹) atmospheric CO₂. Plants were inoculated with the arbuscular mycorrhizal (AM) fungus Glomus mosseae (Nicol. & Gerd.) Gerdemann & Trappe and given a phosphorus supply in the form of bonemeal, which would not be immediately available to the plants. Seven sequential harvests were taken to determine whether the effect of elevated CO₂ on mycorrhizal colonization was independent of the effect of CO₂ on plant growth. Plant growth analysis showed that both species grew faster in elevated CO₂ and that P. lanceolata had increased carbon allocation towards the roots. Elevated CO₂ did not affect the percentage of root length colonized (RLC); although total colonized root length was greater, when plant size was taken into account this effect disappeared. This finding was also true for root length colonized by arbuscules. No CO₂ effect was found on hyphal density (colonization intensity) in roots. The P content of plants was increased at elevated CO₂, although both shoot and root tissue P concentration were unchanged. This was again as a result of bigger plants at elevated CO₂. Phosphorus inflow was unaffected by CO₂ concentrations. It is concluded that there is no direct permanent effect of elevated CO₂ on mycorrhizal functioning, as internal mycorrhizal development and the mycorrhizal P uptake mechanism are unaffected. The importance of sequential harvests in experiments is discussed. The direction for future research is highlighted, especially in relation to C storage in the soil.     

Soil fungal-arthropod responses to Populus tremuloides grown under enriched atmospheric CO2 under field conditions

We investigated the influence of elevated CO₂ and soil N availability on the growth of arbuscular mycorrhizal and non-mycorrhizal fungi, and on the number of mycophagous soil microarthropods associated with the roots of Populus tremuloides . CO₂ concentration did not significantly affect percentage infection of Populus roots by mycorrhizal or non-mycorrhizal fungi. However, the extra-radical hyphal network was altered both qualitatively and quantitatively, and there was a strong interaction between CO₂ and soil N availability. Under N-poor soil conditions, elevated CO₂ stimulated hyphal length by arbuscular mycorrhizal fungi, but depressed growth by non-mycorrhizal fungi. There was no CO₂ effect at high N availability. High N availability stimulated growth by opportunistic saprobic/pathogenic fungi. Soil mites were not affected by any treatment, but collembolan numbers were positively correlated with the increase in non-mycorrhizal fungi. Results indicate a strong interaction between CO₂ concentration and soil N availability on mycorrhizal functioning and on fungal-based soil food webs.     

The active uptake of carbon dioxide by the unicellular green algae Chlorella saccharophila and C. ellipsoidea

Abstract. Mass spectrometry has been used to measure the rates of CO₂ uptake of acid- and alkali-grown cells of the green algae Chlorella ellipsoidea (UTEX 20) and C. saccharophila (UTEX 27). The time course of CO₂ formation on addition of 100mmol m⁻³ K₂CO₃ to cells in the dark was used as an assay for external carbonic anhydrase (CA). No external CA was detected in acid-grown cells of either species or in alkali-grown cells of C. ellipsoidea but was present in alkali-grown C. saccharophila . In the absence of external CA, or when it was inhibited by 5mmol m⁻³ acetazolamide, cells of both species, on illumination, rapidly depleted the free CO₂ in the medium at pH 7.5 to near zero concentrations before maximum photosynthetic O₂ evolution rates were established. Addition of bovine CA rapidly restored the equilibrium CO₂ concentration in the medium, indicating that the cells were selectively taking up CO₂. Transfer of cells to the dark caused a rapid increase in the CO₂ concentration in the medium largely due to the efflux of inorganic carbon from the cells as CO₂. This rapid light-dependent CO₂ uptake takes place against pH and concentration gradients and, thus, has the characteristics of active transport.     

Elevated atmospheric CO2 influences the interaction between the parasitic angiosperm Orobanche minor and its host Trifolium repens

The influence of the root holoparasitic angiosperm Orobanche minor Sm. on the biomass, photosynthesis, carbohydrate and nitrogen content of Trifolium repens L. was determined for plants grown at two CO₂ concentrations (350 and 550 μmol mol⁻¹). Infected plants accumulated less biomass than their uninfected counterparts, although early in the association there was a transient stimulation of growth. Infection also influenced biomass allocation both between tissues (infected plants had lower root:shoot ratios) and within tissues:infected roots were considerably thicker before the point of parasite attachment and thinner below. Higher concentrations of starch were also found in roots above the point of attachment, particularly for plants grown in elevated CO₂. Elevated CO₂ stimulated the growth of T. repens only during the early stages of development. There was a significant interaction between infection and CO₂ on growth, with infected plants showing a greater response, such that elevated CO₂ partly alleviated the effects of the parasite on host growth. Elevated CO₂ did not affect total O. minor biomass per host, the number of individual parasites supported by each host, or their time of attachment to the host root system. Photosynthesis was stimulated by elevated CO₂ but was unaffected by O. minor . There was no evidence of down-regulation of photosynthesis in T. repens grown at elevated CO₂ in either infected or uninfected plants. The data are discussed with regard to the influence of elevated CO₂ on other parasitic angiosperm-host associations and factors which control plant responses to elevated CO₂.     

Interaction of enriched CO2 and water stress on the physiology of and biomass production in sweet potato grown in open-top chambers

Abstract. The objective of this study was to investigate the effects of water stress in sweet potato ( Ipomoea batatas L. [Lam] 'Georgia Jet') on biomass production and plant-water relationships in an enriched CO₂ atmosphere. Plants were grown in pots containing sandy loam soil (Typic Paleudult) at two concentrations of elevated CO₂ and two water regimes in open-top field chambers. During the first 12 d of water stress, leaf xylem potentials were higher in plants grown in a CO₂ concentration of 438 and 666 μmol mol⁻¹ than in plants grown at 364 μmol mol⁻¹. The 364 μmol mol⁻¹ CO₂ grown plants had to be rewatered 2 d earlier than the high CO₂-grown plants in response to water stress. For plants grown under water stress, the yield of storage roots and root: shoot ratio were greater at high CO₂ than at 364 μmol mol⁻¹; the increase, however, was not linear with increasing CO₂ concentrations. In well-watered plants, biomass production and storage root yield increased at elevated CO₂, and these were greater as compared to water-stressed plants grown at the same CO₂ concentration.     

The effects of increased atmospheric carbon dioxide and temperature on carbon partitioning, source-sink relations and respiration

Abstract. Herbaceous C₃ plants grown in elevated CO₂ show increases in carbon assimilation and carbohydrate accumulation (particularly starch) within source leaves. Although changes in the partitioning of biomass between root and shoot occur, the proportion of this extra assimilate made available for sink growth is not known. Root:shoot ratios tend to increase for CO₂-enriched herbaceous plants and decrease for CO₂-enriched trees. Root:shoot ratios for cereals tend to remain constant. In contrast, elevated temperatures decrease carbohydrate accumulation within source and sink regions of a plant and decrease root:shoot ratios. Allometric analysis of at least two species showing changes in root: shoot ratios due to elevated CO₂ show no alteration in the whole-plant partitioning of biomass. Little information is available for interactions between temperature and CO₂. Cold-adapted plants show little response to elevated levels of CO₂, with some species showing a decline in biomass accumulation. In general though, increasing temperature will increase sucrose synthesis, transport and utilization for CO₂-enriched plants and decrease carbohydrate accumulation within the leaf. Literature reports are discussed in relation to the hypothesis that sucrose is a major factor in the control of plant carbon partitioning. A model is presented in support.     

Heterotrophic (Dark) CO2 Fixation by Euglena gracilis. Possible regulation by tricarboxylic acid cycle intermediates

SYNOPSIS. Heterotrophic (dark) CO₂ fixation by Euglena gracilis strain Z varies with phase of batch culture and mode of nutrition. Dark CO₂ fixation increased transiently during the growth of cells under photoautotrophic (CO₂, light) and heterotrophic (glucose, dark) conditions. Cells grown heterotrophically with acetate or ethanol had no transient increase in fixation. The addition of acetate to a heterotrophically growing culture during the period of increasing dark CO₂ fixation resulted in rapid elimination of this fixation. The results suggest that dark CO₂ fixation in Euglena functions in anaplerotic feeding of the tricarboxylic acid cycle, drained by biosyntheses during growth. Induction of the glyoxylate cycle by acetate may provide an alternate source of tricarboxylic cycle intermediates, obviating the requirement for dark CO₂ fixation as a source of the intermediates.     

Responses of carrot fly larvae, Psila rosae, to the odorous and contact-chemostimulatory metabolites of host and non-host plants

ABSTRACT. The responses of third instar carrot fly larvae, Psila rosae , to plant metabolites were studied in a moving air olfactometer. Directed responses over distances greater than 7.5 cm were obtained to both host and non-host plant odours, to CO₂ and methyl eugenol, a volatile secondary metabolite of carrot roots. Larvae also oriented to gradients of contact-chemostimuli. The role of both olfactory and contact-chemostimuli in the host plant finding behaviour of P. rosae larvae is discussed, and observational evidence for the behavioural mechanisms involved presented.     

Effects of mycorrhizal colonization on biomass production and nitrogen fixation of black locust (Robinia pseudoacacia) seedlings grown under elevated atmospheric carbon dioxide

Interactive effects of elevated atmospheric CO₂ and arbuscular mycorrhizal (AM) fungi on biomass production and N₂ fixation were investigated using black locust ( Robinia pseudoacacia ). Seedlings were grown in growth chambers maintained at either 350 μmol mol⁻¹ or 710 μmol mol⁻¹ CO₂. Seedlings were inoculated with Rhizobium spp. and were grown with or without AM fungi. The ¹⁵N isotope dilution method was used to determine N source partitioning between N₂ fixation and inorganic fertilizer uptake. Elevated atmospheric CO₂ significantly increased the percentage of fine roots that were colonized by AM fungi. Mycorrhizal seedlings grown under elevated CO₂ had the greatest overall plant biomass production, nodulation, N and P content, and root N absorption. Additionally, elevated CO₂ levels enhanced nodule and root mass production, as well as N₂ fixation rates, of non- mycorrhizal seedlings. However, the relative response of biomass production to CO₂ enrichment was greater in non-mycorrhizal seedlings than in mycorrhizal seedlings. This study provides strong evidence that arbuscular mycorrhizal fungi play an important role in the extent to which plant nutrition of symbiotic N₂-fixing tree species is affected by enriched atmospheric CO₂.     

Rice production in a changing climate: a meta-analysis of responses to elevated carbon dioxide and elevated ozone concentration

Rice is arguably the most important food source on the planet and is consumed by over half of the world's population. Considerable increases in yield are required over this century to continue feeding the world's growing population. This meta-analysis synthesizes the research to date on rice responses to two elements of global change, rising atmospheric carbon dioxide concentration ([CO₂]) and rising tropospheric ozone concentration ([O₃]). On an average, elevated [CO₂] (627 ppm) increased rice yields by 23%. Modest increases in grain mass and larger increases in panicle and grain number contributed to this response. The response of rice to elevated [CO₂] varied with fumigation technique. The more closely the fumigation conditions mimicked field conditions, the smaller was the stimulation of yield by elevated [CO₂]. Free air concentration enrichment (FACE) experiments showed only a 12% increase in rice yield. The rise in atmospheric [CO₂] will be accompanied by increases in tropospheric O₃ and temperature. When compared with rice grown in charcoal-filtered air, rice exposed to 62 ppb O₃ showed a 14% decrease in yield. Many determinants of yield, including photosynthesis, biomass, leaf area index, grain number and grain mass, were reduced by elevated [O₃]. While there have been too few studies of the interaction of CO₂ and O₃ for meta-analysis, the interaction of temperature and CO₂ has been studied more widely. Elevated temperature treatments negated any enhancement in rice yield at elevated [CO₂], which suggests that identifying high temperature tolerant germplasm will be key to realizing yield benefits in the future.     

Ontogeny affects response of northern red oak seedlings to elevated CO2 and water stress: II. Recent photosynthate distribution and growth

Northern red oak in the western Lake States area of the USA exists on the most xeric edge of its distribution range. Future climate-change scenarios for this area predict decreased water availability along with increased atmospheric CO₂. We examined recent photosynthate distribution and growth in seedlings as a function of CO₂ mole fraction (400, 530 and 700 μmol mol⁻¹ CO₂), water regime (well watered and water-stressed), and ontogenic stage. Water stress effects on growth were largely offset by elevated CO₂.
Water stress increased root mass ratio without concurrently increasing allocation of recent photosynthate to the roots. However, apparent sink strength of water-stressed seedlings at the completion of the third growth stage tended to be greater than that of well watered seedlings, as shown by continued high export, which may contribute carbon reserves to support preferential root growth under water-stressed conditions.
Elevated CO₂ decreased apparent shoot sink strength associated with the rapid expansion of the third flush. Carbon resources for the observed enhanced growth under elevated CO₂ could be provided by enhanced photosynthetic rate over an increased leaf area (Anderson & Tomlinson, 1998, this volume).
Increased sink strength of LG seedlings under water-stressed conditions, together with decreased apparent shoot sink strength associated with growth in elevated CO₂ provide mechanisms for offsetting water stress effects by growth in elevated CO₂.
Careful control of ontogeny was necessary to discern these changes and provides further evidence of the need for such careful control in mechanistic studies.