Responses of leaf photosynthesis,pigments and chlorophyll fluorescence within canopy position in a boreal grass (<Emphasis Type="Italic">Phalaris arundinacea</Emphasis> L.) to elevated temperature and CO<Subscript>2</Subscript> under varying water regimes

The effects of elevated growth temperature (ambient + 3.5°C) and CO₂ (700 μmol mol⁻¹) on leaf photosynthesis, pigments and chlorophyll fluorescence of a boreal perennial grass (Phalaris arundinacea L.) under different water regimes (well watered to water shortage) were investigated. Layer-specific measurements were conducted on the top (younger leaf) and low (older leaf) canopy positions of the plants after anthesis. During the early development stages, elevated temperature enhanced the maximum rate of photosynthesis (P _max) of the top layer leaves and the aboveground biomass, which resulted in earlier senescence and lower photosynthesis and biomass at the later periods. At the stage of plant maturity, the content of chlorophyll (Chl), leaf nitrogen (N_L), and light response of effective photochemical efficiency (Φ_PSII) and electron transport rate (ETR) was significantly lower under elevated temperature than ambient temperature in leaves at both layers. CO₂ enrichment enhanced the photosynthesis but led to a decline of N_L and Chl content, as well as lower fluorescence parameters of Φ_PSII and ETR in leaves at both layers. In addition, the down-regulation by CO₂ elevation was significant at the low canopy position. Regardless of climate treatment, the water shortage had a strongly negative effect on the photosynthesis, biomass growth, and fluorescence parameters, particularly in the leaves from the low canopy position. Elevated temperature exacerbated the impact of water shortage, while CO₂ enrichment slightly alleviated the drought-induced adverse effects on P _max. We suggest that the light response of Φ_PSII and ETR, being more sensitive to leaf-age classes, reflect the photosynthetic responses to climatic treatments and drought stress better than the fluorescence parameters under dark adaptation.     

Effects of elevated CO2 and temperature on leaf characteristics,photosynthesis and carbon storage in aboveground biomass of a boreal bioenergy crop (Phalaris arundinacea L.) under varying water regimes

This work examined the effects of elevated CO₂ and temperature and water regimes, alone and in interaction, on the leaf characteristics [leaf area (LA), specific leaf weight (SLW), leaf nitrogen content (N_L) based on LA], photosynthesis (light‐saturated net carbon fixation rate, P_sat) and carbon storage in aboveground biomass of leaves (C_l) and stem (C_s) for a perennial reed canary grass (Phalaris arundinacea L., Finnish local cultivar). For this purpose, plants were grown under different water regimes (ranging from high to low soil moisture) in climate‐controlled growth chambers under the elevated CO₂ and/or temperature (following a factorial design) over a whole growing season (May–September in 2009). The results showed that the elevated temperature increased the leaf growth, photosynthesis and carbon storage of aboveground biomass the most in the early growing periods, compared with ambient temperature. However, the plant growth declined rapidly thereafter with a lower carbon storage at the end of growing season. This was related to the accelerated phenology regulation and consequent earlier growth senescence. Consequently, the elevation of CO₂ increased the P_sat, LA and SLW during the growing season, with a significant concurrent increase in the carbon storage in aboveground biomass. Low soil moisture decreased the P_sat, leaf stomatal conductance, LA and carbon storage in above ground biomass compared with high and normal soil moisture. This water stress effect was the largest under the elevated temperature. The elevated CO₂ partially mitigated the adverse effects of high temperature and low soil moisture. However, the combination of elevated temperature and CO₂ did not significantly increase the carbon storage in aboveground biomass of the plants.     

Changes in growth and photosynthetic patterns of oil palm (<Emphasis Type="Italic">Elaeis guineensis</Emphasis> Jacq.) seedlings exposed to short-term CO<Subscript>2</Subscript> enrichment in a closed top chamber

Three varieties of oil palm seedlings (Deli Yangambi, Deli Urt, Deli AVROS) were exposed to three levels of CO₂ (400, 800, 1,200 μmol/mol) in split plot design to determine growth (net assimilation rate, NAR; relative growth rate, RGR) and photosynthetic patterns of the seedlings under short-term CO₂ exposure of 15 weeks. Increasing CO₂ from 400 to 800 and 1,200 μmol/mol significantly enhanced total biomass and leaf area, net photosynthesis (A) and water use efficiency (WUE) especially from weeks 9 to 15. By the end of week 15, total biomass increased by 113%, and A and WUE by one- and fivefold, respectively, while specific leaf area decreased by 37%. Both enhanced biomass and A under elevated CO₂ were effective in modifying NAR and RGR as shown by high correlation coefficient values (r ² = 0.68 and 0.72; r ² = 0.63 and 0.67, respectively), although WUE seemed to have more influence over the NAR (r ² = 0.97) and RGR (r ² = 0.93). Neither interspecific preference nor its interaction with CO₂ imposed any significant effect on parameters observed. Growth improvement with CO₂ seemed able to produce healthy, bigger and vigorous oil palm seedlings, and the technique may have potential to be developed for use to reduce nursery period.     

Carbon balance in a monospecific stand of an annual herb <Emphasis Type="Italic">Chenopodium album</Emphasis> at an elevated CO<Subscript>2</Subscript> concentration

Elevated CO₂ enhances carbon uptake of a plant stand, but the magnitude of the increase varies among growth stages. We studied the relative contribution of structural and physiological factors to the CO₂ effect on the carbon balance during stand development. Stands of an annual herb Chenopodium album were established in open-top chambers at ambient and elevated CO₂ concentrations (370 and 700 μmol mol⁻¹). Plant biomass growth, canopy structural traits (leaf area, leaf nitrogen distribution, and light gradient in the canopy), and physiological characteristics (leaf photosynthesis and respiration of organs) were studied through the growing season. CO₂ exchange of the stand was estimated with a canopy photosynthesis model. Rates of light-saturated photosynthesis and dark respiration of leaves as related with nitrogen content per unit leaf area and time-dependent reduction in specific respiration rates of stems and roots were incorporated into the model. Daily canopy carbon balance, calculated as an integration of leaf photosynthesis minus stem and root respiration, well explained biomass growth determined by harvests (r ² = 0.98). The increase of canopy photosynthesis with elevated CO₂ was 80% at an early stage and decreased to 55% at flowering. Sensitivity analyses suggested that an alteration in leaf photosynthetic traits enhanced canopy photosynthesis by 40–60% throughout the experiment period, whereas altered canopy structure contributed to the increase at the early stage only. Thus, both physiological and structural factors are involved in the increase of carbon balance and growth rate of C. album stands at elevated CO₂. However, their contributions were not constant, but changed with stand development.     

Innate and evolutionarily increased advantages of invasive <Emphasis Type="Italic">Eupatorium adenophorum</Emphasis> over native <Emphasis Type="Italic">E. japonicum</Emphasis> under ambient and doubled atmospheric CO<Subscript>2</Subscript> concentrations

Both innate and evolutionarily increased ecophysiological advantages can contribute to vigorous growth, and eventually to invasiveness of alien plants. Little effort has been made to explore the roles of innate factors of alien plants in invasiveness and the effects of CO₂ enrichment on alien plant invasions. To address these problems, we compared invasive Eupatorium adenophorum, its native conspecific, and a native congener (E. japonicum) under ambient and doubled atmospheric CO₂ concentrations. Native E. adenophorum from Mexico grew slower than invasive E. adenophorum but faster than native E. japonicum under both CO₂ concentrations. The faster growth rate of invasive E. adenophorum was associated with higher photosynthetic capacity and leaf area ratio. For invasive E. adenophorum, the higher photosynthetic capacity was associated with higher nitrogen (N) allocation to photosynthesis, which was related to lower leaf mass per area; the higher leaf area ratio was due to lower leaf mass per area and higher leaf mass fraction. Tradeoff between N allocations to photosynthesis versus defenses was found. CO₂ enrichment significantly increased relative growth rate and biomass accumulation by increasing actual photosynthetic rate for all studied materials. However, the relative increase in growth was not significantly different among them. CO₂ enrichment did not influence N allocation to photosynthesis, but increased N allocation to cell walls. The reduced leaf N content decreased N content in photosynthesis, explaining the down-regulation of photosynthetic capacity under prolonged elevated CO₂ concentration. Our results indicate that both innate and evolutionary advantages in growth and related ecophysiological traits contribute to invasiveness of invasive E. adenophorum, and CO₂ enrichment may not aggravate E. adenophroum’s invasion in the future.     

Effects of Elevated CO2 and Temperature on Biomass Growth and Allocation in a Boreal Bioenergy Crop (Phalaris arundinacea L.) from Young and Old Cultivations

An auto-controlled climate system was used to study how a boreal bioenergy crop (reed canary grass, Phalaris arundinacea L., hereafter RCG) responded to a warming climate and elevated CO₂. Over one growing season (April–September of 2009), RCG from young and old cultivations (3 years [3-year] and 10 years [10-year]) was grown in closed chambers under ambient conditions (CON), elevated CO₂ (EC, approximately 700 μmol?mol^?1), elevated temperature (ET, ambient + approximately 3 °C) and elevated temperature and CO₂ (ETC). The treatments were replicated four times. Throughout the growing season, the above-ground (leaf and stem biomass) and below-ground biomasses were measured six times, representing various developmental stages (early stages: the first three stages, and late stages: the last three stages). Compared to the growth observed under CON, EC enhanced RCG biomass growth over the whole growing season (p?<?0.05), whereas ET increased RCG biomass growth in early stages but decreased growth in late stages, regardless of the cultivation age. However, the negative effect of ET later in the growing season was partially mitigated by CO₂ enrichment. Compared to CON plants, the final total biomass was 18 % higher for 3-year plants and 8 % higher for 10-year plants grown under EC. In comparison, for 3-year and 10-year plants, the biomass was 5 and 3 % lower under ET and 7 and 4 % greater under ETC, respectively. Under EC, the below-ground growth contributed more to the total biomass growth compared to the above-ground portion. The opposite situation was observed under ET and ETC. The climate-related changes in biomass growth were smaller in the old cultivation than in the young cultivation due to the lower net assimilation rate and lower specific leaf area in the old cultivation plants.     

Plant responses to elevated CO<Subscript>2</Subscript> concentration at different scales: leaf,whole plant,canopy, and population

Elevated CO₂ enhances photosynthesis and growth of plants, but the enhancement is strongly influenced by the availability of nitrogen. In this article, we summarise our studies on plant responses to elevated CO₂. The photosynthetic capacity of leaves depends not only on leaf nitrogen content but also on nitrogen partitioning within a leaf. In Polygonum cuspidatum, nitrogen partitioning among the photosynthetic components was not influenced by elevated CO₂ but changed between seasons. Since the alteration in nitrogen partitioning resulted in different CO₂-dependence of photosynthetic rates, enhancement of photosynthesis by elevated CO₂ was greater in autumn than in summer. Leaf mass per unit area (LMA) increases in plants grown at elevated CO₂. This increase was considered to have resulted from the accumulation of carbohydrates not used for plant growth. With a sensitive analysis of a growth model, however, we suggested that the increase in LMA is advantageous for growth at elevated CO₂ by compensating for the reduction in leaf nitrogen concentration per unit mass. Enhancement of reproductive yield by elevated CO₂ is often smaller than that expected from vegetative growth. In Xanthium canadense, elevated CO₂ did not increase seed production, though the vegetative growth increased by 53%. As nitrogen concentration of seeds remained constant at different CO₂ levels, we suggest that the availability of nitrogen limited seed production at elevated CO₂ levels. We found that leaf area development of plant canopy was strongly constrained by the availability of nitrogen rather than by CO₂. In a rice field cultivated at free-air CO₂ enrichment, the leaf area index (LAI) increased with an increase in nitrogen availability but did not change with CO₂ elevation. We determined optimal LAI to maximise canopy photosynthesis and demonstrated that enhancement of canopy photosynthesis by elevated CO₂ was larger at high than at low nitrogen availability. We also studied competitive asymmetry among individuals in an even-aged, monospecific stand at elevated CO₂. Light acquisition (acquired light per unit aboveground mass) and utilisation (photosynthesis per unit acquired light) were calculated for each individual in the stand. Elevated CO₂ enhanced photosynthesis and growth of tall dominants, which reduced the light availability for shorter subordinates and consequently increased size inequality in the stand.     

Elevated Atmospheric CO<Subscript>2</Subscript> Increases Root Exudation of Carbon in Wetlands: Results from the First Free-Air CO<Subscript>2</Subscript> Enrichment Facility (FACE) in a Marshland

Experiments employing free-air CO₂ enrichment (FACE) facilities have indicated that elevated atmospheric carbon dioxide (eCO₂) stimulates growth in diverse terrestrial ecosystems. Studies of the effects of eCO₂ on wetland plants have indicated a similar response, but these studies were mostly performed in growth chambers. We conducted a 2-year FACE experiment [CO₂ ≈ 582 µmol mol^?1] in a marsh in Spain to test whether the common reed (Phragmites australis) responds to carbon enrichment, as previously reported in other macrophytes. More specifically, we tested the effect of eCO₂ on P. australis growth, photosynthesis, transpiration, and biomass, its effect on modifying plant and soil ratios of carbon, nitrogen, and phosphorus, and whether the strong environmental variability of this wetland modulates these responses. Our findings show that effects of eCO₂ in this wetland environment are more complex than previously believed, probably due to hydrological effects. The effects of eCO₂ on reed plants were cumulative and manifested at the end of the growing season as increased 38–44% instantaneous transpiration efficiency (ratio of net photosynthesis to transpiration), which was dependent on plant age. However, this increase did not result in a significant increase in biomass, because of excessive root exudation of carbon. These observations contrast with previous observations of wetland plants to increased atmospheric CO₂ in growth chambers and shed new light on the role of wetland plants as a carbon sink in the face of global climate change. The combined effects of water stress, eCO₂, and soil carbon processes must be considered when assessing the function of wetlands as a carbon sink under global change scenarios.     

The impact of elevated CO<Subscript>2</Subscript> and water deficit stress on growth and photosynthesis of juvenile cacao (<Emphasis Type="Italic">Theobroma cacao</Emphasis> L.).

Atmospheric CO₂ concentration continues to rise and is predicted to reach approximately 700 ppm by 2100. Some predictions suggest that the dry season in West Africa could be extended with climate change. This study examined the effects of elevated CO₂ concentration and water deficit on growth and photosynthesis of juvenile cacao. Light-saturated photosynthesis (P_max), quantum efficiency, and intrinsic water-use efficiency increased significantly in response to elevated CO₂, as did a range of growth and development responses (e.g. leaf area and leaf number), but the magnitude of the increase was dependent on the water treatment. Stomatal index was significantly greater in the elevated CO₂ treatment; an atypical response which may be a reflection of the environment in which cacao evolved. This study shows a positive effect of elevated CO₂ on juvenile cacao which may help to alleviate some of the negative impacts of water deficit stress.     

Development of gypsy moth larvae feeding on red maple saplings at elevated CO<Subscript>2</Subscript> and temperature

Predicted increases in atmospheric CO₂ and global mean temperature may alter important plant-insect associations due to the direct effects of temperature on insect development and the indirect effects of elevated temperature and CO₂ enrichment on phytochemicals important for insect success. We investigated the effects of CO₂ and temperature on the interaction between gypsy moth (Lymantria dispar L.) larvae and red maple (Acer rubrum L.) saplings by bagging first instar larvae within open-top chambers at four CO₂/temperature treatments: (1) ambient temperature, ambient CO₂, (2) ambient temperature, elevated CO₂ (+300 l l^-1 CO₂), (3) elevated temperature (+3.5°C), ambient CO₂, and (4) elevated temperature, elevated CO₂. Larvae were reared to pupation and leaf samples taken biweekly to determine levels of total N, water, non-structural carbohydrates, and an estimate of defensive phenolic compounds in three age classes of foliage: (1) immature, (2) mid-mature and (3) mature. Elevated growth temperature marginally reduced (P <0.1) leaf N and significantly reduced (P <0.05) leaf water across CO₂ treatments in mature leaves, whereas leaves grown at elevated CO₂ concentration had a significant decrease in leaf N and a significant increase in the ratio of starch:N and total non-structural carbohydrates:N. Leaf N and water decreased and starch:N and total non-structural carbohydrates:N ratios increased as leaves aged. Phenolics were unaffected by CO₂ or temperature treatment. There were no interactive effects of CO₂ and temperature on any phytochemical measure. Gypsy moth larvae reached pupation earlier at the elevated temperature (female =8 days, P <0.07; male =7.5 days, P <0.03), whereas mortality and pupal fresh weight of insects were unrelated to either CO₂, temperature or their interaction. Our data show that CO₂ or temperature-induced alterations in leaf constituents had no effect on insect performance; instead, the long-term exposure to a 3.5°C increase in temperature shortened insect development but had no effect on pupal weight. It appears that in some tree-herbivorous insect systems the direct effects of an increased global mean temperature may have greater consequences for altering plant-insect interactions than the indirect effects of an increased temperature or CO₂ concentration on leaf constituents.     

Effect of panicle removal on photosynthetic acclimation under elevated CO<Subscript>2</Subscript> in rice

To examine the role of sink size on photosynthetic acclimation under elevated atmospheric CO₂ concentrations ([CO₂]), we tested the effects of panicle-removal (PR) treatment on photosynthesis in rice (Oryza sativa L.). Rice was grown at two [CO₂] levels (ambient and ambient + 200 μmol mol⁻¹) throughout the growing season, and at full-heading stage, at half the plants, a sink-limitation treatment was imposed by the removal of the panicles. The PR treatment alleviated the reduction of green leaf area, the contents of chlorophyll (Chl) and Rubisco after the full-heading stage, suggesting delay of senescence. Nonetheless, elevated [CO₂] decreased photosynthesis (measured at current [CO₂]) of plants exposed to the PR treatment. No significant [CO₂] × PR interaction on photosynthesis was observed. The decrease of photosynthesis by elevated [CO₂] of plants was associated with decreased leaf Rubisco content and N content. Leaf glucose content was increased by the PR treatment and also by elevated [CO₂]. In conclusion, a sink-limitation in rice improved N status in the leaves, but this did not prevent the photosynthetic down-regulation under elevated [CO₂].     

Compensatory effects of elevated CO<Subscript>2</Subscript> concentration on the inhibitory effects of high temperature and irradiance on photosynthetic gas exchange in carrots

We determined the interactive effects of irradiance, elevated CO₂ concentration (EC), and temperature in carrot (Daucus carota var. sativus). Plants of the cv. Red Core Chantenay (RCC) were grown in a controlled environmental plant growth room and exposed to 3 levels of photosynthetically active radiation (PAR) (400, 800, 1 200 μmol m⁻² s⁻¹), 3 leaf chamber temperatures (15, 20, 30 °C), and 2 external CO₂ concentrations (C _a), AC and EC (350 and 750 μmol mol⁻¹, respectively). Rates of net photosynthesis (P _N) and transpiration (E) and stomatal conductance (g _s ) were measured, along with water use efficiency (WUE) and ratio of internal and external CO₂ concentrations (C _i/C _a). P _N revealed an interactive effect between PAR and C _a. As PAR increased so did P _N under both C _a regimes. The g _s showed no interactive effects between the three parameters but had singular effects of temperature and PAR. E was strongly influenced by the combination of PAR and temperature. WUE was interactively affected by all three parameters. Maximum WUE occurred at 15 °C and 1 200 μmol m⁻² s^{− 1} PAR under EC. The C _i /C _a was influenced independently by temperature and C _a. Hence photosynthetic responses are interactively affected by changes in irradiance, external CO₂ concentration, and temperature. EC significantly compensates the inhibitory effects of high temperature and irradiance on P _N and WUE.     

Responses of tropical native and invader C<Subscript>4</Subscript> grasses to water stress,clipping and increased atmospheric CO<Subscript>2</Subscript> concentration

The invasion of African grasses into Neotropical savannas has altered savanna composition, structure and function. The projected increase in atmospheric CO₂ concentration has the potential to further alter the competitive relationship between native and invader grasses. The objective of this study was to quantify the responses of two populations of a widespread native C₄ grass (Trachypogon plumosus) and two African C₄ grass invaders (Hyparrhenia rufa and Melinis minutiflora) to high CO₂ concentration interacting with two primary savanna stressors: drought and herbivory. Elevated CO₂ increased the competitive potential of invader grasses in several ways. Germination and seedling size was promoted in introduced grasses. Under high CO₂, the relative growth rate of young introduced grasses was twice that of native grass (0.58 g g⁻¹ week⁻¹ vs 0.25 g g⁻¹ week⁻¹). This initial growth advantage was maintained throughout the course of the study. Well-watered and unstressed African grasses also responded more to high CO₂ than did the native grass (biomass increases of 21–47% compared with decreases of 13–51%). Observed higher water and nitrogen use efficiency of invader grasses may aid their establishment and competitive strength in unfertile sites, specially if the climate becomes drier. In addition, high CO₂ promoted lower leaf N content more in the invader grasses. The more intensive land use, predicted to occur in this region, may interact with high CO₂ to fincreasesavor the African grasses, as they generally recovered faster after simulated herbivory. The superiority of invader grasses under high CO₂ suggests further in their competitive strength and a potential increased rate of displacement of the native savannas in the future by grasslands dominated by introduced African species.     

A hierarchical analysis of the interactive effects of elevated CO<Subscript>2</Subscript> and water availability on the nitrogen and transpiration productivities of velvet mesquite seedlings

In this study we apply new extensions of classical growth analysis to assess the interactive effects of elevated CO₂ and differences in water availability on the leaf-nitrogen and transpiration productivities of velvet mesquite (Prosopis velutina Woot.) seedlings. The models relate transpiration productivity (biomass gained per mass of water transpired per day) and leaf-nitrogen productivity (biomass gain per unit leaf N per day) to whole-plant relative growth rate (RGR) and to each other, allowing a comprehensive hierarchical analysis of how physiological and morphological responses to the treatments interact with each other to affect plant growth. Elevated CO₂ led to highly significant increases in N and transpiration productivities but reduced leaf N per unit leaf area and transpiration per unit leaf area, resulting in no net effect of CO₂ on the RGR of seedlings. In contrast, higher water availability led to an increase in leaf-tissue thickness or density without affecting leaf N concentration, resulting in a higher leaf N per unit leaf area and consequently a higher assimilatory capacity per unit leaf area. The net effect was a marginal increase in seedling RGR. Perhaps most important from an ecological perspective was a 41% reduction in whole-plant water use due to elevated CO₂. These results demonstrate that even in the absence of CO₂ effects on integrative measures of plant growth such as RGR, highly significant effects may be observed at the physiological and morphological level that effectively cancel each other out. The quantitative framework presented here enables some of these tradeoffs to be identified and related directly to each other and to plant growth.     

Autumnal leaf abscission of sugar maple is not delayed by atmospheric CO<Subscript>2</Subscript> enrichment

To investigate the effects of atmospheric CO₂ enrichment on physiology and autumnal leaf phenology, we exposed 3-year-old sugar maple (Acer saccharum Marsh.) seedlings to 800 (A8), 600 (A6), and 400 μL(CO₂) L^–1 (AA) in nine continuous stirred tank reactor (CSTR) chambers during the growing season of 2014. Leaf abscission timing, abscised leaf area percentages, leaf number, light-saturated net photosynthetic rate (P_Nmax), leaf area, accumulative growth rates, and biomass were determined and assessed. The results suggested the following: (1) no significant differences were found in the timing of leaf abscission in the three CO₂-concentration treatments; (2) P_Nmax was continuously stimulated to the greatest extent in A8 at 319% and 160% in A6 until the end of the growing season, respectively; and (3) leaf number, leaf area, and accumulative height growth all significantly increased by elevated CO₂, which led to a 323% increase in A8 biomass and 235% in A6 biomass after 156-d fumigation. In summary, the results suggest, the timing of leaf abscission of sugar maple in fall was not modified by CO₂ enrichment, the increased carbon gain by elevated CO₂ was mainly due to increased leaf area, more leaves, and the continuously enhanced high photosynthesis throughout the growing season instead of the leaf life span.     

Performance of a generalist grasshopper on a C<Subscript>3</Subscript> and a C<Subscript>4</Subscript> grass: compensation for the effects of elevated CO<Subscript>2</Subscript> on plant nutritional quality

The increasing CO₂ concentration in Earths atmosphere is expected to cause a greater decline in the nutritional quality of C₃ than C₄ plants. As a compensatory response, herbivorous insects may increase their feeding disproportionately on C₃ plants. These hypotheses were tested by growing the grasses Lolium multiflorum C₃) and Bouteloua curtipendula C₄) at ambient (370 ppm) and elevated (740 ppm) CO₂ levels in open top chambers in the field, and comparing the growth and digestive efficiencies of the generalist grasshopper Melanoplus sanguinipes on each of the four plant × CO₂ treatment combinations. As expected, the nutritional quality of the C₃ grass declined to a greater extent than did that of the C₄ grass at elevated CO₂; protein levels declined in the C₃ grass, while levels of carbohydrates (sugar, fructan and starch) increased. However, M. sanguinipes did not significantly increase its consumption rate to compensate for the lower nutritional quality of the C₃ grass grown under elevated CO₂. Instead, these grasshoppers appear to use post-ingestive mechanisms to maintain their growth rates on the C₃ grass under elevated CO₂. Consumption rates of the C₃ and C₄ grasses were also similar, demonstrating a lack of compensatory feeding on the C₄ grass. We also examined the relative efficiencies of nutrient utilization from a C₃ and C₄ grass by M. sanguinipes to test the basis for the C₄ plant avoidance hypothesis. Contrary to this hypothesis, neither protein nor sugar was digested with a lower efficiency from the C₄ grass than from the C₃ grass. A novel finding of this study is that fructan, a potentially large carbohydrate source in C₃ grasses, is utilized by grasshoppers. Based on the higher nutrient levels in the C₃ grass and the better growth performance of M. sanguinipes on this grass at both CO₂ levels, we conclude that C₃ grasses are likely to remain better host plants than C₄ grasses in future CO₂ conditions.     

A meta-analysis of plant physiological and growth responses to temperature and elevated CO<Subscript>2</Subscript>

Atmospheric carbon dioxide (CO₂) and global mean temperature are expected to be significantly higher by the end of the 21st century. Elevated CO₂ (eCO₂) and higher temperature each affect plant physiology and growth, but their interactive effects have not been reviewed statistically with respect to higher chronic mean temperatures and abrupt heat stress. In this meta-analysis, we examined the effect of CO₂ on the physiology and growth of plants subjected to different temperature treatments. The CO₂ treatments were categorized into ambient (<400 ppm) or elevated (>560 ppm) levels, while temperature treatments were categorized into ambient temperature (AT), elevated temperature (ET; AT + 1.4–6°C), or heat stress (HS; AT + >8°C). Plant species were grouped according to photosynthetic pathways (C₃, C₄), functional types (legumes, non-legumes), growth forms (herbaceous, woody), and economic purposes (crop, non-crop). eCO₂ enhanced net photosynthesis at AT, ET, and HS in C₃ species (especially at the HS level), but in C₄ species, it had no effect at AT, a positive effect at ET, and a negative effect at HS. The positive effect of eCO₂ on net photosynthesis was greater for legumes than for non-legumes at HS, for non-crops than crops at ET, and for woody than herbaceous species at ET and HS. Total (W _T) and above- (W _AG) and below-ground (W _BG) biomass were increased by eCO₂ for most species groups at all temperatures, except for C₄ species and W _BG of legumes at HS. Hence, eCO₂ × heat effects on growth were often not explained by effects on net photosynthesis. Overall, the results show that eCO₂ effects on plant physiology and growth vary under different temperature regimes, among functional groups and photosynthetic pathways, and among response variables. These findings have important implications for biomass accumulation and ecosystem functioning in the future when the CO₂ level is higher and climate extremes, such as heat waves, become more frequent.     

The stomatal CO<Subscript>2</Subscript> proxy does not saturate at high atmospheric CO<Subscript>2</Subscript> concentrations: evidence from stomatal index responses of Araucariaceae conifers

The inverse relationship between the number of stomata on a leaf surface and the atmospheric carbon dioxide concentration ([CO₂]) in which the leaf developed allows plants to optimise water-use efficiency (WUE), but it also permits the use of fossil plants as proxies of palaeoatmospheric [CO₂]. The ancient conifer family Araucariaceae is often represented in fossil floras and may act as a suitable proxy of palaeo-[CO₂], yet little is known regarding the stomatal index (SI) responses of extant Araucariaceae to [CO₂]. Four Araucaria species (Araucaria columnaris, A. heterophylla, A. angustifolia and A. bidwillii) and Agathis australis displayed no significant relationship in SI to [CO₂] below current ambient levels (~380 ppm). However, representatives of the three extant genera within the Araucariaceae (A. bidwillii, A. australis and Wollemia nobilis) all exhibited significant reductions in SI when grown in atmospheres of elevated [CO₂] (1,500 ppm). Stomatal conductance was reduced and WUE increased when grown under elevated [CO₂]. Stomatal pore length did not increase alongside reduced stomatal density (SD) and SI in the three araucariacean conifers when grown at elevated [CO₂]. These pronounced SD and SI reductions occur at higher [CO₂] levels than in other species with more recent evolutionary origins, and may reflect an evolutionary legacy of the Araucariaceae in the high [CO₂] world of the Mesozoic Era. Araucariacean conifers may therefore be suitable stomatal proxies of palaeo-[CO₂] during periods of “greenhouse” climates and high [CO₂] in the Earth’s history.     

Effects of elevated atmospheric CO<Subscript>2</Subscript> on the nutritional ecology of C<Subscript>3</Subscript> and C<Subscript>4</Subscript> grass-feeding caterpillars

It is plausible that the nutritional quality of C₃ plants will decline more under elevated atmospheric CO₂ than will the nutritional quality of C₄ plants, causing herbivorous insects to increase their feeding on C₃ plants relative to C₄ plants. We tested this hypothesis with a C₃ and C₄ grass and two caterpillar species with different diet breadths. Lolium multiflorum (C₃) and Bouteloua curtipendula (C₄) were grown in outdoor open top chambers at ambient (370 ppm) or elevated (740 ppm) CO₂. Bioassays compared the performance and digestive efficiencies of Pseudaletia unipuncta (a grass-specialist noctuid) and Spodoptera frugiperda (a generalist noctuid). As expected, the nutritional quality of L. multiflorum changed to a greater extent than did that of B. curtipendula when grown in elevated CO₂; levels of protein (considered growth limiting) declined in the C₃ grass, while levels of carbohydrates (sugar, starch and fructan) increased. However, neither insect species increased its feeding rate on the C₃ grass to compensate for its lower nutritional quality when grown in an elevated CO₂ atmosphere. Consumption rates of P. unipuncta and S. frugiperda were higher on the C₃ grass than the C₄ grass, the opposite of the result expected for a compensatory response to the lower nutritional quality of the C₄ grass. Although our results do not support the hypothesis that grass-specialist insects compensate for lower nutritional quality by increasing their consumption rates more than do generalist insects, the performance of the specialist was greater than that of the generalist on each grass species and at both CO₂ levels. Mechanisms other than compensatory feeding, such as increased nutrient assimilation efficiency, appear to determine the relative performance of these herbivores. Our results also provide further evidence against the hypothesis that C₄ grasses would be avoided by insect herbivores because a large fraction of their nutrients is unavailable to herbivores. Instead, our results are consistent with the hypothesis that C₄ grasses are poorer host plants primarily because of their lower nutrient levels, higher fiber levels, and greater toughness.     

An estimation of CO<Subscript>2</Subscript> fixation capacity in mangrove forest using two methods of CO<Subscript>2</Subscript> gas exchange and growth curve analysis

In many coastal areas of South-East Asia, attempts have been made to revive coastal ecosystem by initiating projects that encourage planting of mangrove trees. Compared to the terrestrial trees, mangrove trees possess a higher carbon fixation capacity. It becomes a very significant option for clean development mechanism (CDM) program. However, a reliable method to estimate CO₂ fixation capacity of mangrove trees has not been established. Acknowledging the above fact, we decided to set up an estimation method for the CDM program, using gas exchange analysis to estimate mangrove productivity, we put into consideration the net CO₂ fixation of reforested Kandelia candel (5-, 10-, and 15-year-old stand). This was estimated by gas exchange analysis and growth curve analysis. In growth curve analysis, we drew a growth curve of a single stand using data of above- and below-ground biomass. In the gas exchange analysis, we calculated CO₂ fixation capacity by (1) measuring respiration rate of each organ of stand and calculating respiratory CO₂ emission from above- to below-ground biomass. (2) Measuring the single-leaf photosynthetic rate in response to light intensity and calculating the photosynthetic CO₂ absorption. (3) We also developed a model for the diurnal changes in temperature, and monthly averages based on one-day estimation of CO₂ absorption and emission, which we corrected by this model in order to estimate the net CO₂ fixation capacity in response to temperature. Comparing the biomass accumulation of the two methods constructed for the same forest, the above-ground biomass accumulation of 10-year-old forest (34.3 ton ha⁻¹ yr⁻¹) estimated by gas exchange analysis was closely compared to those of growth curve analysis (26.6 ton ha⁻¹ yr⁻¹), suggesting that the gas exchange analysis was capable of estimating mangrove productivity. The validity of the estimated CO₂ fixation capacity by the gas exchange analysis and the growth curve analysis was also discussed.