Carbon dioxide‐ and temperature‐mediated changes in plant defensive compounds alter food utilization of herbivores

Although the impact of elevated carbon dioxide and rising temperature on plants and animals has been extensively documented recently, only limited understanding exists regarding their combined effects. The objective of this research was to address the consequences of using combinations of elevated CO₂ and elevated temperature on a plant's defensive chemistry, and subsequent utilization of the plant as insect food. Our results indicated that elevated CO₂ and increased temperature, for the most part, act independently on the production of defensive compounds in broccoli leaves (Brassica oleracea L. var. italica). CO₂ concentrations had significant effects on the foliar water content, total phenolic compounds, polyphenol oxidase and trypsin inhibitor concentrations. The herbivore Spodoptera litura (Fabricius; Lepidoptera: Noctuidae) responded to changes in the plant secondary chemistry, with larvae consuming more plant materials that had been exposed to elevated CO₂. The food utilization efficiencies of second‐instar larvae were more sensitive to CO₂‐treated foliage than those of the third‐ and fourth‐instar larvae. Temperature did exert a significant effect on food utilization (ECD) by the larvae. Our study will provide important information in future predictions on plant–insect interactions as a result of climate change. The study also demonstrated that since various larval stages might respond differently to climate change, this possibility needs to be considered in future forecasting and monitoring.     

Rice grain yield and quality responses to free‐air CO2 enrichment combined with soil and water warming

Rising air temperatures are projected to reduce rice yield and quality, whereas increasing atmospheric CO₂ concentrations ([CO₂]) can increase grain yield. For irrigated rice, ponded water is an important temperature environment, but few open‐field evaluations are available on the combined effects of temperature and [CO₂], which limits our ability to predict future rice production. We conducted free‐air CO₂ enrichment and soil and water warming experiments, for three growing seasons to determine the yield and quality response to elevated [CO₂] (+200 μmol mol^?1, E‐[CO₂]) and soil and water temperatures (+2 °C, E‐T). E‐[CO₂] significantly increased biomass and grain yield by approximately 14% averaged over 3 years, mainly because of increased panicle and spikelet density. E‐T significantly increased biomass but had no significant effect on the grain yield. E‐T decreased days from transplanting to heading by approximately 1%, but days to the maximum tiller number (MTN) stage were reduced by approximately 8%, which limited the panicle density and therefore sink capacity. On the other hand, E‐[CO₂] increased days to the MTN stage by approximately 4%, leading to a greater number of tillers. Grain appearance quality was decreased by both treatments, but E‐[CO₂] showed a much larger effect than did E‐T. The significant decrease in undamaged grains (UDG) by E‐[CO₂] was mainly the result of an increased percentage of white‐base grains (WBSG), which were negatively correlated with grain protein content. A significant decrease in grain protein content by E‐[CO₂] accounted in part for the increased WBSG. The dependence of WBSG on grain protein content, however, was different among years; the slope and intercept of the relationship were positively correlated with a heat dose above 26 °C. Year‐to‐year variation in the response of grain appearance quality demonstrated that E‐[CO₂] and rising air temperatures synergistically reduce grain appearance quality of rice.     

Will chemical defenses become more effective against specialist herbivores under elevated CO2?

Elevated atmospheric CO₂ is known to affect plant–insect herbivore interactions. Elevated CO₂ causes leaf nitrogen to decrease, the ostensible cause of herbivore compensatory feeding. CO₂ may also affect herbivore consumption by altering chemical defenses via changes in plant hormones. We considered the effects of elevated CO₂, in conjunction with soil fertility and damage (simulated herbivory), on glucosinolate concentrations of mustard (Brassica nigra) and collard (B. oleracea var. acephala) and the effects of leaf nitrogen and glucosinolate groups on specialist Pieris rapae consumption. Elevated CO₂ affected B. oleracea but not B. nigra glucosinolates; responses to soil fertility and damage were also species‐specific. Soil fertility and damage also affected B. oleracea glucosinolates differently under elevated CO₂. Glucosinolates did not affect P. rapae consumption at either CO₂ concentration in B. nigra, but had CO₂‐specific effects on consumption in B. oleracea. At ambient CO₂, leaf nitrogen had strong effects on glucosinolate concentrations and P. rapae consumption but only gluconasturtiin was a feeding stimulant. At elevated CO₂, direct effects of leaf nitrogen were weaker, but glucosinolates had stronger effects on consumption. Gluconasturtiin and aliphatic glucosinolates were feeding stimulants and indole glucosinolates were feeding deterrents. These results do not support the compensatory feeding hypothesis as the sole driver of changes in P. rapae consumption under elevated CO₂. Support for hormone‐mediated CO₂ response (HMCR) was mixed; it explained few treatment effects on constitutive or induced glucosinolates, but did explain patterns in SEMs. Further, the novel feeding deterrent effect of indole glucosinolates under elevated CO₂ in B. oleracae underscores the importance of defensive chemistry in CO₂ response. We speculate that P. rapae indole glucosinolate detoxification mechanisms may have been overwhelmed under elevated CO₂ forcing slowed consumption. Specialists may have to contend with hosts with poorer nutritional quality and more effective chemical defenses under elevated CO₂.     

Growth responses of gypsy moth larvae to elevated CO2: the influence of methods of insect rearing

Abstract The effects of elevated CO₂ on foliar chemistry of two tree species (Populus pseudo‐simonii Kitag. and Betula platyphylla) and on growth of gypsy moth (Lymantria dispar L.) larvae were examined. Furthermore, we focused on the comparison of results on the growth responses of larvae obtained from two methods of insect rearing, the no‐choice feeding trial performed in the laboratory or in situ in open‐top chambers. On the whole, both primary and secondary metabolites in the leaves of the two tree species were significantly affected by main effects of time (sampling date), CO₂ and species. Elevated CO₂ significantly increased the C : N ratio and concentrations of the soluble sugar, starch, total nonstructural carbohydrates, total phenolics and condensed tannins, but significantly decreased the concentration of nitrogen. Higher contents of total phenolics and condensed tannins were detected in the frass of larvae reared in elevated CO₂ treatments. Overall, the growth of gypsy moth larvae were significantly inhibited by elevated CO₂ and CO₂‐induced changes in leaf quality. Our study did not indicate the two methods of insect rearing could influence the direction of effects of elevated CO₂ on the growth of individual insects; however, the magnitude of negative effects of elevated CO₂ on larval growth did differ between the two insect rearing methods, and it seems that the response magnitude was also mediated by larval age and host plant species.     

Sustained effects of atmospheric [CO2] and nitrogen availability on forest soil CO2 efflux

Soil CO₂ efflux (F_soil) is the largest source of carbon from forests and reflects primary productivity as well as how carbon is allocated within forest ecosystems. Through early stages of stand development, both elevated [CO₂] and availability of soil nitrogen (N; sum of mineralization, deposition, and fixation) have been shown to increase gross primary productivity, but the long‐term effects of these factors on F_soil are less clear. Expanding on previous studies at the Duke Free‐Air CO₂ Enrichment (FACE) site, we quantified the effects of elevated [CO₂] and N fertilization on F_soil using daily measurements from automated chambers over 10 years. Consistent with previous results, compared to ambient unfertilized plots, annual F_soil increased under elevated [CO₂] (ca. 17%) and decreased with N (ca. 21%). N fertilization under elevated [CO₂] reduced F_soil to values similar to untreated plots. Over the study period, base respiration rates increased with leaf productivity, but declined after productivity saturated. Despite treatment‐induced differences in aboveground biomass, soil temperature and water content were similar among treatments. Interannually, low soil water content decreased annual F_soil from potential values – estimated based on temperature alone assuming nonlimiting soil water content – by ca. 0.7% per 1.0% reduction in relative extractable water. This effect was only slightly ameliorated by elevated [CO₂]. Variability in soil N availability among plots accounted for the spatial variability in F_soil, showing a decrease of ca. 114 g C m^?2 yr^?1 per 1 g m^?2 increase in soil N availability, with consistently higher F_soil in elevated [CO₂] plots ca. 127 g C per 100 ppm [CO₂] over the +200 ppm enrichment. Altogether, reflecting increased belowground carbon partitioning in response to greater plant nutritional needs, the effects of elevated [CO₂] and N fertilization on F_soil in this stand are sustained beyond the early stages of stand development and through stabilization of annual foliage production.     

Interactive effects of pre-industrial, current and future [CO2] and temperature on an insect herbivore of Eucalyptus

Both atmospheric [CO₂] and average surface temperatures are predicted to increase with potentially different, additive or opposing, effects on leaf quality and insect herbivore activity. Few studies have directly measured the interactive effects of concurrent changes in [CO₂] and temperature on insect herbivores. None have done so over the entire developmental period of a tree-feeding insect, and none have compared responses to low pre-industrial [CO₂] and present day [CO₂] to estimate responses to future increases. Eucalypt herbivores may be particularly sensitive to climate-driven shifts in plant chemistry, as eucalypt foliage is naturally low in [N]. In this study, we assessed the development of the eucalypt herbivore Doratifera quadriguttata exposed concurrently to variable [CO₂] (290, 400, 650 μmol mol^?1) and temperature (ambient, ambient +4 °C) on glasshouse-grown Eucalyptus tereticornis. Overall, insects performed best on foliage grown at pre-industrial [CO₂], indicating that modern insect herbivores have already experienced nutritional shifts since industrialisation. Rising [CO₂] increased specific leaf mass and leaf carbohydrate concentration, subsequently reducing leaf [N]. Lower leaf [N] induced compensatory feeding and impeded insect performance, particularly by prolonging larval development. Importantly, elevated temperature dampened the negative effects of rising [CO₂] on larval performance. Therefore, rising [CO₂] over the past 200 years may have reduced forage quality for eucalypt insects, but concurrent temperature increases may have partially compensated for this, and may continue to do so in the future. These results highlight the importance of assessing plant–insect interactions within the context of multiple climate-change factors because of the interactive and potentially opposing effects of different factors within and between trophic levels.     

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.     

Aphid and host‐plant genotype × genotype interactions under elevated CO2

1. Elevated CO₂ can alter plant physiology and morphology, and these changes are expected to impact diet quality for insect herbivores. While the plastic responses of insect herbivores have been well studied, less is known about the propensity of insects to adapt to such changes. Genetic variation in insect responses to elevated CO₂ and genetic interactions between insects and their host plants may exist and provide the necessary raw material for adaptation. 2. We used clonal lines of Rhopalosiphum padi (L.) aphids to examine genotype‐specific responses to elevated CO₂. We used the host plant Schedonorus arundinaceus (tall fescue; Schreb), which is capable of asexual reproduction, to investigate host plant genotype‐specific effects and possible host plant‐by‐insect genotype interactions. The abundance and density of three R. padi genotypes on three tall fescue genotypes under three concentrations of CO₂ (ambient, 700, and 1000 ppm) in a controlled greenhouse environment were examined. 3. Aphid abundance decreased in the 700 ppm CO₂ concentration, but increased in the 1000 ppm concentration relative to ambient. The effect of CO₂ on aphid density was dependent on host plant genotype; the density of aphids in high CO₂ decreased for two plant genotypes but was unchanged in one. No interaction between aphid genotype and elevated CO₂ was found, nor did we find significant genotype‐by‐genotype interactions. 4. This study suggests that the density of R. padi aphids feeding on tall fescue may decrease under elevated CO₂ for some plant genotypes. The likely impact of genotype‐specific responses on future changes in the genetic structure of plant and insect populations is discussed.     

Photosynthetic responses of two eucalypts to industrial‐age changes in atmospheric [CO2] and temperature

The unabated rise in atmospheric [CO₂] is associated with increased air temperature. Yet, few CO₂‐enrichment studies have considered pre‐industrial [CO₂] or warming. Consequently, we quantified the interactive effects of growth [CO₂] and temperature on photosynthesis of faster‐growing Eucalyptus saligna and slower‐growing E. sideroxylon. Well‐watered and ‐fertilized tree seedlings were grown in a glasshouse at three atmospheric [CO₂] (290, 400, and 650 µL L^?1), and ambient (26/18 °C, day/night) and high (ambient + 4 °C) air temperature. Despite differences in growth rate, both eucalypts responded similarly to [CO₂] and temperature treatments with few interactive effects. Light‐saturated photosynthesis (A_sat) and light‐ and [CO₂]‐saturated photosynthesis (A_max) increased by ～50% and ～10%, respectively, with each step‐increase in growth [CO₂], underpinned by a corresponding 6–11% up‐regulation of maximal electron transport rate (J_max). Maximal carboxylation rate (V_cmax) was not affected by growth [CO₂]. Thermal photosynthetic acclimation occurred such that A_sat and A_max were similar in ambient‐ and high‐temperature‐grown plants. At high temperature, the thermal optimum of A_sat increased by 2–7 °C across [CO₂] treatments. These results are the first to suggest that photosynthesis of well‐watered and ‐fertilized eucalypt seedlings will remain strongly responsive to increasing atmospheric [CO₂] in a future, warmer climate.     

Do elevated atmospheric CO2 and O3 affect food quality and performance of folivorous insects on silver birch?

The individual and combined effects of elevated CO₂ and O₃ on the foliar chemistry of silver birch (Betula pendula Roth) and on the performance of five potential birch‐defoliating insect herbivore species (two geometrid moths, one lymantrid moth and two weevils) were examined. Elevated CO₂ decreased the water concentration in both short‐ and long‐shoot leaves, but the effect of CO₂ on the concentration of nitrogen and individual phenolic compounds was mediated by O₃ treatment, tree genotype and leaf type. Elevated O₃ increased the total carbon concentration only in short‐shoot leaves. Bioassays showed that elevated CO₂ increased the food consumption rate of juvenile Epirrita autumnata and Rheumaptera hastata larvae fed with short‐ and long‐shoot leaves in spring and mid‐summer, respectively, but had no effect on the growth of larvae. The contribution of leaf quality variables to the observed CO₂ effects indicate that insect compensatory consumption may be related to leaf age. Elevated CO₂ increased the food preference of only two tested species: Phyllobius argentatus (CO₂ alone) and R. hastata (CO₂ combined with O₃). The observed stimulus was dependent on tree genotype and the measured leaf quality variables explained only a portion of the stimulus. Elevated O₃ decreased the growth of flush‐feeding young E. autumnata larvae, irrespective of CO₂ concentration, apparently via reductions in general food quality. Therefore, the increasing tropospheric O₃ concentration could pose a health risk for juvenile early‐season birch folivores in future. In conclusion, the effects of elevated O₃ were found to be detrimental to the performance of early‐season insect herbivores in birch whereas elevated CO₂ had only minor effects on insect performance despite changes in food quality related foliar chemistry.     

The effect of heat waves,elevated [CO2] and low soil water availability on northern red oak (Quercus rubra L.) seedlings

The frequency and intensity of heat waves are predicted to increase. This study investigates whether heat waves would have the same impact as a constant increase in temperature with the same heat sum, and whether there would be any interactive effects of elevated [CO₂] and soil moisture content. We grew Quercus rubra seedlings in treatment chambers maintained at either ambient or elevated [CO₂] (380 or 700 μmol CO₂ mol^?1) with temperature treatments of ambient, ambient +3 °C, moderate heat wave (+6 °C every other week) or severe heat wave (+12 °C every fourth week) temperatures. Averaged over a 4‐week period, and the entire growing season, the three elevated temperature treatments had the same average temperature and heat sum. Half the seedlings were watered to a soil water content near field capacity, half to about 50% of this value. Foliar gas exchange measurements were performed morning and afternoon (9:00 and 15:00 hours) before, during and after an applied heat wave in August 2010. Biomass accumulation was measured after five heat wave cycles. Under ambient [CO₂] and well‐watered conditions, biomass accumulation was highest in the +3 °C treatment, intermediate in the +6 °C heat wave and lowest in the +12 °C heat wave treatment. This response was mitigated by elevated [CO₂]. Low soil moisture significantly decreased net photosynthesis (A_net) and biomass in all [CO₂] and temperature treatments. The +12 °C heat wave reduced afternoon A_net by 23% in ambient [CO₂]. Although this reduction was relatively greater under elevated [CO₂], A_net values during this heat wave were still 34% higher than under ambient [CO₂]. We concluded that heat waves affected biomass growth differently than the same amount of heat applied uniformly over the growing season, and that the plant response to heat waves also depends on [CO₂] and soil moisture conditions.     

Lower responsiveness of canopy evapotranspiration rate than of leaf stomatal conductance to open‐air CO2 elevation in rice

An elevated atmospheric CO₂ concentration ([CO₂]) can reduce stomatal conductance of leaves for most plant species, including rice (Oryza sativa L.). However, few studies have quantified seasonal changes in the effects of elevated [CO₂] on canopy evapotranspiration, which integrates the response of stomatal conductance of individual leaves with other responses, such as leaf area expansion, changes in leaf surface temperature, and changes in developmental stages, in field conditions. We conducted a field experiment to measure seasonal changes in stomatal conductance of the uppermost leaves and in the evapotranspiration, transpiration, and evaporation rates using a lysimeter method. The study was conducted for flooded rice under open‐air CO₂ elevation. Stomatal conductance decreased by 27% under elevated [CO₂], averaged throughout the growing season, and evapotranspiration decreased by an average of 5% during the same period. The decrease in daily evapotranspiration caused by elevated [CO₂] was more significantly correlated with air temperature and leaf area index (LAI) rather than with other parameters of solar radiation, days after transplanting, vapor‐pressure deficit and FAO reference evapotranspiration. This indicates that higher air temperatures, within the range from 16 to 27 °C, and a larger LAI, within the range from 0 to 4 m² m^?2, can increase the magnitude of the decrease in evapotranspiration rate caused by elevated [CO₂]. The crop coefficient (i.e. the evapotranspiration rate divided by the FAO reference evapotranspiration rate) was 1.24 at ambient [CO₂] and 1.17 at elevated [CO₂]. This study provides the first direct measurement of the effects of elevated [CO₂] on rice canopy evapotranspiration under open‐air conditions using the lysimeter method, and the results will improve future predictions of water use in rice fields.     

Canopy radiation‐ and water‐use efficiencies as affected by elevated [CO2]

This study used an environmentally controlled plant growth facility, EcoCELLs, to measure canopy gas exchanges directly and to examine the effects of elevated [CO₂] on canopy radiation‐ and water‐use efficiencies. Sunflowers (Helianthus annus var. Mammoth) were grown at ambient (399 μmol mol^?1) and elevated [CO₂] (746 μmol mol^?1) for 53 days in EcoCELLs. Whole canopy carbon‐ and water‐fluxes were measured continuously during the period of the experiment. The results indicated that elevated [CO₂] enhanced daily total canopy carbon‐ and water‐fluxes by 53% and 11%, respectively, on a ground‐area basis, resulting in a 54% increase in radiation‐use efficiency (RUE) based on intercepted photosynthetic active radiation and a 26% increase in water‐use efficiency (WUE) by the end of the experiment. Canopy carbon‐ and water‐fluxes at both CO₂ treatments varied with canopy development. They were small at 22 days after planting (DAP) and gradually increased to the maxima at 46 DAP. When canopy carbon‐ and water‐fluxes were expressed on a leaf‐area basis, no effect of CO₂ was found for canopy water‐flux while elevated [CO₂] still enhanced canopy carbon‐flux by 29%, on average. Night‐time canopy carbon‐flux was 32% higher at elevated than at ambient [CO₂]. In addition, RUE and WUE displayed strong diurnal variations, high at noon and low in the morning or afternoon for WUE but opposite for RUE. This study provided direct evidence that plant canopy may consume more, instead of less, water but utilize both water and radiation more efficiently at elevated than at ambient [CO₂], at least during the exponential growth period as illustrated in this experiment.     

Future carbon dioxide concentration decreases canopy evapotranspiration and soil water depletion by field‐grown maize

Maize, in rotation with soybean, forms the largest continuous ecosystem in temperate North America, therefore changes to the biosphere‐atmosphere exchange of water vapor and energy of these crops are likely to have an impact on the Midwestern US climate and hydrological cycle. As a C₄ crop, maize photosynthesis is already CO₂‐saturated at current CO₂ concentrations ([CO₂]) and the primary response of maize to elevated [CO₂] is decreased stomatal conductance (g_s). If maize photosynthesis is not stimulated in elevated [CO₂], then reduced g_s is not offset by greater canopy leaf area, which could potentially result in a greater ET reduction relative to that previously reported in soybean, a C₃ species. The objective of this study is to quantify the impact of elevated [CO₂] on canopy energy and water fluxes of maize (Zea mays). Maize was grown under ambient and elevated [CO₂] (550 μmol mol^?1 during 2004 and 2006 and 585 μmol mol^?1 during 2010) using Free Air Concentration Enrichment (FACE) technology at the SoyFACE facility in Urbana, Illinois. Maize ET was determined using a residual energy balance approach based on measurements of sensible (H) and soil heat fluxes, and net radiation. Relative to control, elevated [CO₂] decreased maize ET (7–11%; P < 0.01) along with lesser soil moisture depletion, while H increased (25–30 W m^?2; P < 0.01) along with higher canopy temperature (0.5–0.6 °C). This reduction in maize ET in elevated [CO₂] is approximately half that previously reported for soybean. A partitioning analysis showed that transpiration contributed less to total ET for maize compared to soybean, indicating a smaller role of stomata in dictating the ET response to elevated [CO₂]. Nonetheless, both maize and soybean had significantly decreased ET and increased H, highlighting the critical role of elevated [CO₂] in altering future hydrology and climate of the region that is extensively cropped with these species.     

Distinct seasonal dynamics of responses to elevated CO2 in two understorey grass species differing in shade‐tolerance

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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.     

Long‐term elevated air [CO2] strengthens photosynthetic functioning and mitigates the impact of supra‐optimal temperatures in tropical Coffea arabica and C. canephora species

The tropical coffee crop has been predicted to be threatened by future climate changes and global warming. However, the real biological effects of such changes remain unknown. Therefore, this work aims to link the physiological and biochemical responses of photosynthesis to elevated air [CO₂] and temperature in cultivated genotypes of Coffea arabica L. (cv. Icatu and IPR108) and Coffea canephora cv. Conilon CL153. Plants were grown for ca. 10 months at 25/20 °C (day/night) and 380 or 700 μl CO₂ l^?1 and then subjected to temperature increase (0.5 °C day^?1) to 42/34 °C. Leaf impacts related to stomatal traits, gas exchanges, C isotope composition, fluorescence parameters, thylakoid electron transport and enzyme activities were assessed at 25/20, 31/25, 37/30 and 42/34 °C. The results showed that (1) both species were remarkably heat tolerant up to 37/30 °C, but at 42/34 °C a threshold for irreversible nonstomatal deleterious effects was reached. Impairments were greater in C. arabica (especially in Icatu) and under normal [CO₂]. Photosystems and thylakoid electron transport were shown to be quite heat tolerant, contrasting to the enzymes related to energy metabolism, including RuBisCO, which were the most sensitive components. (2) Significant stomatal trait modifications were promoted almost exclusively by temperature and were species dependent. Elevated [CO₂], (3) strongly mitigated the impact of temperature on both species, particularly at 42/34 °C, modifying the response to supra‐optimal temperatures, (4) promoted higher water‐use efficiency under moderately higher temperature (31/25 °C) and (5) did not provoke photosynthetic downregulation. Instead, enhancements in [CO₂] strengthened photosynthetic photochemical efficiency, energy use and biochemical functioning at all temperatures. Our novel findings demonstrate a relevant heat resilience of coffee species and that elevated [CO₂] remarkably mitigated the impact of heat on coffee physiology, therefore playing a key role in this crop sustainability under future climate change scenarios.     

Elevated [CO2] does not ameliorate the negative effects of elevated temperature on drought‐induced mortality in Eucalyptus radiata seedlings

It has been reported that elevated temperature accelerates the time‐to‐mortality in plants exposed to prolonged drought, while elevated [CO₂] acts as a mitigating factor because it can reduce stomatal conductance and thereby reduce water loss. We examined the interactive effects of elevated [CO₂] and temperature on the inter‐dependent carbon and hydraulic characteristics associated with drought‐induced mortality in Eucalyptus radiata seedlings grown in two [CO₂] (400 and 640 μL L^?1) and two temperature (ambient and ambient +4 °C) treatments. Seedlings were exposed to two controlled drying and rewatering cycles, and then water was withheld until plants died. The extent of xylem cavitation was assessed as loss of stem hydraulic conductivity. Elevated temperature triggered more rapid mortality than ambient temperature through hydraulic failure, and was associated with larger water use, increased drought sensitivities of gas exchange traits and earlier occurrence of xylem cavitation. Elevated [CO₂] had a negligible effect on seedling response to drought, and did not ameliorate the negative effects of elevated temperature on drought. Our findings suggest that elevated temperature and consequent higher vapour pressure deficit, but not elevated [CO₂], may be the primary contributors to drought‐induced seedling mortality under future climates.     

Photosynthetic downregulation in leaves of the Japanese white birch grown under elevated CO2 concentration does not change their temperature‐dependent susceptibility to photoinhibition

To determine the effects of elevated CO₂ concentration ([CO₂]) on the temperature‐dependent photosynthetic properties, we measured gas exchange and chlorophyll fluorescence at various leaf temperatures (15, 20, 25, 30, 35 and 40°C) in 1‐year‐old seedlings of the Japanese white birch (Betula platyphylla var. japonica), grown in a phytotron under natural daylight at two [CO₂] levels (ambient: 400 µmol mol^?1 and elevated: 800 µmol mol^?1) and limited N availability (90 mg N plant^?1). Plants grown under elevated [CO₂] exhibited photosynthetic downregulation, indicated by a decrease in the carboxylation capacity of Rubisco. At temperatures above 30°C, the net photosynthetic rates of elevated‐CO₂‐grown plants exceeded those grown under ambient [CO₂] when compared at their growth [CO₂]. Electron transport rates were significantly lower in elevated‐CO₂‐grown plants than ambient‐CO₂‐grown ones at temperatures below 25°C. However, no significant difference was observed in the fraction of excess light energy [(1 ? q_P)× F_v′/F_m′] between CO₂ treatments across the temperature range. The quantum yield of regulated non‐photochemical energy loss was significantly higher in elevated‐CO₂‐grown plants than ambient, when compared at their respective growth [CO₂] below 25°C. These results suggest that elevated‐CO₂‐induced downregulation might not exacerbate the temperature‐dependent susceptibility to photoinhibition, because reduced energy consumption by electron transport was compensated for by increased thermal energy dissipation at low temperatures.     

Enhanced isoprene emission capacity and altered light responsiveness in aspen grown under elevated atmospheric CO2 concentration

Controversial evidence of CO₂‐responsiveness of isoprene emission has been reported in the literature with the response ranging from inhibition to enhancement, but the reasons for such differences are not understood. We studied isoprene emission characteristics of hybrid aspen (Populus tremula x P. tremuloides) grown under ambient (380 μmol mol^?1) and elevated (780 μmol mol^?1) [CO₂] to test the hypothesis that growth [CO₂] effects on isoprene emission are driven by modifications in substrate pool size, reflecting altered light use efficiency for isoprene synthesis. A novel in vivo method for estimation of the pool size of the immediate isoprene precursor, dimethylallyldiphosphate (DMADP) and the activity of isoprene synthase was used. Growth at elevated [CO₂] resulted in greater leaf thickness, more advanced development of mesophyll and moderately increased photosynthetic capacity due to morphological “upregulation”, but isoprene emission rate under growth light and temperature was not significantly different among ambient‐ and elevated‐[CO₂]‐grown plants independent of whether measured at 380 μmol mol^?1 or 780 μmol mol^?1 CO₂. However, DMADP pool size was significantly less in elevated‐[CO₂]‐grown plants, but this was compensated by increased isoprene synthase activity. Analysis of CO₂ and light response curves of isoprene emission demonstrated that the [CO₂] for maximum isoprene emission was shifted to lower [CO₂] in elevated‐[CO₂]‐grown plants. The light‐saturated isoprene emission rate (I_max,Q) was greater, but the quantum efficiency at given I_max,Q was less in elevated‐[CO₂]‐grown plants, especially at higher CO₂ measurement concentration, reflecting stronger DMADP limitation at lower light and higher [CO₂]. These results collectively demonstrate important shifts in light and CO₂‐responsiveness of isoprene emission in elevated‐[CO₂]‐acclimated plants that need consideration in modeling isoprene emissions in future climates.