Consequences of CO2 and light interactions for leaf phenology, growth, and senescence in Quercus rubra

We investigated how light and CO₂ levels interact to influence growth, phenology, and the physiological processes involved in leaf senescence in red oak (Quercus rubra) seedlings. We grew plants in high and low light and in elevated and ambient CO₂. At the end of three years of growth, shade plants showed greater biomass enhancement under elevated CO₂ than sun plants. We attribute this difference to an increase in leaf area ratio (LAR) in shade plants relative to sun plants, as well as to an ontogenetic effect: as plants increased in size, the LAR declined concomitant with a decline in biomass enhancement under elevated CO₂ Elevated CO₂ prolonged the carbon gain capacity of shade‐grown plants during autumnal senescence, thus increasing their functional leaf lifespan. The prolongation of carbon assimilation, however, did not account for the increased growth enhancement in shade plants under elevated CO₂. Elevated CO₂ did not significantly alter leaf phenology. Nitrogen concentrations in both green and senesced leaves were lower under elevated CO₂ and declined more rapidly in sun leaves than in shade leaves. Similar to nitrogen concentration, the initial slope of A/C_i curves indicated that Rubisco activity declined more rapidly in sun plants than in shade plants, particularly under elevated CO₂. Absolute levels of chlorophyll were affected by the interaction of CO₂ and light, and chlorophyll content declined to a minimal level in sun plants sooner than in shade plants. These declines in N concentration, in the initial slope of A/C_i curves, and in chlorophyll content were consistent with declining photosynthesis, such that elevated CO₂ accelerated senescence in sun plants and prolonged leaf function in shade plants. These results have implications for the carbon economy of seedlings and the regeneration of red oak under global change conditions.     

Nitrate supply and plant development influence nitrogen uptake and allocation under elevated CO<Subscript>2</Subscript> in durum wheat grown hydroponically

Growth in elevated CO₂ often leads to decreased plant nitrogen contents and down-regulation of photosynthetic capacity. Here, we investigated whether elevated CO₂ limits nitrogen uptake when nutrient movement to roots is unrestricted, and the dependence of this limitation on nitrogen supply and plant development in durum wheat (Triticum durum Desf.). Plants were grown hydroponically at two N supplies and ambient and elevated CO₂ concentrations. Elevated CO₂ decreased nitrate uptake per unit root mass with low N supply at early grain filling, but not at anthesis. This decrease was not associated with higher nitrate or amino acid, or lower non-structural carbohydrate contents in roots. At anthesis, elevated CO₂ decreased the nitrogen content of roots with both levels of N and that of aboveground organs with high N. With low N, elevated CO₂ increased N allocation to aboveground plant organs and nitrogen concentration per unit flag leaf area at anthesis, and per unit aboveground dry mass at both growth stages. The results from the hydroponic experiment suggest that elevated CO₂ restricts nitrate uptake late in development, high N supply overriding this restriction. Increased nitrogen allocation to young leaves at low N supply could alleviate photosynthetic acclimation to elevated CO₂.     

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 CO<Subscript>2</Subscript>, litter chemistry,and decomposition: a synthesis

The results of published and unpublished experiments investigating the impacts of elevated [CO₂] on the chemistry of leaf litter and decomposition of plant tissues are summarized. The data do not support the hypothesis that changes in leaf litter chemistry often associated with growing plants under elevated [CO₂] have an impact on decomposition processes. A meta-analysis of data from naturally senesced leaves in field experiments showed that the nitrogen (N) concentration in leaf litter was 7.1% lower in elevated [CO₂] compared to that in ambient [CO₂]. This statistically significant difference was: (1) usually not significant in individual experiments, (2) much less than that often observed in green leaves, and (3) less in leaves with an N concentration indicative of complete N resorption. Under ideal conditions, the efficiency with which N is resorbed during leaf senescence was found not to be altered by CO₂ enrichment, but other environmental influences on resorption inevitably increase the variability in litter N concentration. Nevertheless, the small but consistent decline in leaf litter N concentration in many experiments, coupled with a 6.5% increase in lignin concentration, would be predicted to result in a slower decomposition rate in CO₂-enriched litter. However, across the assembled data base, neither mass loss nor respiration rates from litter produced in elevated [CO₂] showed any consistent pattern or differences from litter grown in ambient [CO₂]. The effects of [CO₂] on litter chemistry or decomposition were usually smallest under experimental conditions similar to natural field conditions, including open-field exposure, plants free-rooted in the ground, and complete senescence. It is concluded that any changes in decomposition rates resulting from exposure of plants to elevated [CO₂] are small when compared to other potential impacts of elevated [CO₂] on carbon and N cycling. Reasons for experimental differences are considered, and recommendations for the design and execution of decomposition experiments using materials from CO₂-enrichment experiments are outlined.     

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].     

RETRACTED ARTICLE: CO<Subscript>2</Subscript> effects on plant nutrient concentration depend on plant functional group and available nitrogen: a meta-analysis

Elevated CO₂ is expected to lower plant nutrient concentrations via carbohydrate dilution and increased nutrient use efficiency. Elevated CO₂ consistently lowers plant foliar nitrogen, but there is no consensus on CO₂ effects across the range of plant nutrients. We used meta-analysis to quantify elevated CO₂ effects on leaf, stem, root, and seed concentrations of B, Ca, Cu, Fe, K, Mg, Mn, P, S, and Zn among four plant functional groups and two levels of N fertilization. CO₂ effects on plant nutrient concentration depended on the nutrient, plant group, tissue, and N status. CO₂ reduced B, Cu, Fe, and Mg, but increased Mn concentration in the leaves of N₂ fixers. Elevated CO₂ increased Cu, Fe, and Zn, but lowered Mn concentration in grass leaves. Tree leaf responses were strongly related to N status: CO₂ significantly decreased Cu, Fe, Mg, and S at high N, but only Fe at low N. Elevated CO₂ decreased Mg and Zn in crop leaves grown with high N, and Mn at low N. Nutrient concentrations in crop roots were not affected by CO₂ enrichment, but CO₂ decreased Ca, K, Mg and P in tree roots. Crop seeds had lower S under elevated CO₂. We also tested the validity of a “dilution model.” CO₂ reduced the concentration of plant nutrients 6.6% across nutrients and plant groups, but the reduction is less than expected (18.4%) from carbohydrate accumulation alone. We found that elevated CO₂ impacts plant nutrient status differently among the nutrient elements, plant functional groups, and among plant tissues. Our synthesis suggests that differences between plant groups and plant organs, N status, and differences in nutrient chemistry in soils preclude a universal hypothesis strictly related to carbohydrate dilution regarding plant nutrient response to elevated CO₂.     

Effect of elevated CO<Subscript>2</Subscript> levels on leaf starch,nitrogen and photosynthesis of plants growing at three natural CO<Subscript>2</Subscript> springs in Japan

Plant communities around natural CO₂ springs have been exposed to elevated CO₂ levels over many generations and give us a unique opportunity to investigate the effects of long-term elevated CO₂ levels on wild plants. We searched for natural CO₂ springs in cool temperate climate regions in Japan and found three springs that were suitable for studying long-term responses of plants to elevated levels of CO₂: Ryuzin-numa, Yuno-kawa and Nyuu. At these CO₂ springs, the surrounding air was at high CO₂ concentration with no toxic gas emissions throughout the growth season, and there was natural vegetation around the springs. At each site, high-CO₂ (HC) and low-CO₂ (LC) plots were established, and three dominant species at the shrub layers were used for physiological analyses. Although the microenvironments were different among the springs, dicotyledonous species growing at the HC plots tended to have more starch and less nitrogen per unit dry mass in the leaves than those growing at the LC plots. In contrast, monocotyledonous species growing in the HC and LC plots had similar starch and nitrogen concentrations. Photosynthetic rates at the mean growth CO₂ concentration were higher in HC plants than LC plants, but photosynthetic rates at a common CO₂ concentration were lower in HC plants. Efficiency of water and nitrogen use of leaves at growth CO₂ concentration was greatly increased in HC plants. These results suggest that natural plants growing in elevated CO₂ levels under cool temperate climate conditions have down-regulated their photosynthetic capacity but that they increased photosynthetic rates and resource use efficiencies due to the direct effect of elevated CO₂ concentration.     

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.     

Effects of elevated CO2 and temperature on plant growth and herbivore defensive chemistry

Concentration of atmospheric CO₂ and temperature have both been rising for the last three decades. In this century, the temperature has been predicted to rise by 2–5 °C and the CO₂ concentration to double. These changes may affect the primary and secondary metabolism of plants and thus have implications for other trophic levels. However, the biotic interactions in changing climate conditions are poorly known. In this study, two questions were addressed: (i) How will climate change affect growth and the amounts of secondary compounds in flexible plant species? and (ii) How will this affect herbivores living on this species. Four clones of the dark‐leaved willow (Salix myrsinifolia (Salisb.)) seedlings were grown in closed‐top chambers with two controlled factors: concentration of atmospheric CO₂ and temperature (T). There were four combinations of these factors, each combination replicated four times (total of 16 chambers): (i) Control CO₂ (350 ppm) and control T, (ii) Elevated CO₂ (700 ppm) and control T, (iii) Control CO₂ and elevated T (2 °C), and (iv) Elevated CO₂ and elevated T. Stem growth and aerial biomass of the plants were determined; and the leaf phenolics, nitrogen and water concentrations were analysed. In addition the growth rate of larvae and feeding preference of adults of a specialist herbivore, the chrysomelid beetle Phratora vitellinae (L.), on the treated willow leaves were measured. Elevated temperature and CO₂ concentration increased the stem biomass and elevated CO₂ increased leaf biomass and total aerial biomass of the willows. Patterns of biomass allocation were different in different temperature treatments. At elevated temperature there was less branch and leaf material in relation to stems than at the control temperature. Moreover, patterns of biomass allocation differed among clones. CO₂ enhancement increased the specific leaf weight (SLW) and reduced both water and nitrogen content of the leaves, however, leaf area was unaffected by the treatments. Carbon dioxide (CO₂) and T enhancement reduced the concentrations of several phenolic compounds in the leaves. Phenolic compounds, nutrients, and water in the leaves might be diluted partly due to increased carbon allocation to different structures (e.g. thickening of cell wall and increase of trichomes, etc.). In some cases plant clones showed specific responses to treatments. The CO₂ enhancement reduced the relative growth rate (RGR) of the beetle larvae, and in contrast, temperature elevation increased it. Adult beetles did not clearly discriminate between willow leaves grown in different T and CO₂ environments, but tended to eat more leaf material from chambers with doubled CO₂ concentration. At elevated CO₂ adult beetles may need to eat more leaf material in order to reproduce, which may in turn prolong the life cycles, increasing the risk of being eaten and possibly affecting ability to overwinter successfully. Overall, climate change may significantly modify the dynamic interaction between willow and beetle populations.     

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.     

Changes in Leaf Morphology and Composition with Future Increases in CO<Subscript>2</Subscript> and Temperature Revisited: Wheat in Field Chambers

Whether leaf morphology is altered by future increases in atmospheric CO₂ and temperature has been reexamined over 3 years in wheat grown in field chambers at two levels of nitrogen supply. Flag leaf fresh and dry mass, area, volume, and ratios of these parameters, as well as the contents of water, chlorophyll, nonstructural carbohydrates, and nitrogen compounds have been determined at anthesis and 14 days later. High CO₂ decreased rather than increased, as reported in the literature, leaf mass per area and leaf density, and increased water content per area and per volume and water percentage. Warmer temperatures also decreased leaf mass per area, but did not affect density or water per area or per volume, whereas they increased water percentage. Nitrogen supply did not change CO₂ and temperature effects on leaf morphology. Nonstructural carbohydrates increased and nitrogen compounds decreased in elevated CO₂, and the sum of these compounds decreased with warmer temperatures. These changes in composition did not account for modifications of leaf morphology. We conclude that increases in atmospheric CO₂ and temperature after leaf initiation can decrease leaf mass per area, and elevated CO₂ can also decrease leaf density, due to decreases in leaf structural compounds. The functional significance of these changes is probably a decrease in photosynthetic capacity per unit leaf area.     

Profiles of isoprene emission and photosynthetic parameters in hybrid poplars exposed to free‐air CO2 enrichment†

Poplar (Populus × euroamericana) saplings were grown in the field to study the changes of photosynthesis and isoprene emission with leaf ontogeny in response to free air carbon dioxide enrichment (FACE) and soil nutrient availability. Plants growing in elevated [CO₂] produced more leaves than those in ambient [CO₂]. The rate of leaf expansion was measured by comparing leaves along the plant profile. Leaf expansion and nitrogen concentration per unit of leaf area was similar between nutrient treatment, and this led to similar source–sink functional balance. Consequently, soil nutrient availability did not cause downward acclimation of photosynthetic capacity in elevated [CO₂] and did not affect isoprene synthesis. Photosynthesis assessed in growth [CO₂] was higher in plants growing in elevated than in ambient [CO₂]. After normalizing for the different number of leaves over the profile, maximal photosynthesis was reached and started to decline earlier in elevated than in ambient [CO₂]. This may indicate a [CO₂]‐driven acceleration of leaf maturity and senescence. Isoprene emission was adversely affected by elevated [CO₂]. When measured on the different leaves of the profile, isoprene peak emission was higher and was reached earlier in ambient than in elevated [CO₂]. However, a larger number of leaves was emitting isoprene in plant growing in elevated [CO₂]. When integrating over the plant profile, emissions in the two [CO₂] levels were not different. Normalization as for photosynthesis showed that profiles of isoprene emission were remarkably similar in the two [CO₂] levels, with peak emissions at the centre of the profile. Only the rate of increase of the emission of young leaves may have been faster in elevated than in ambient [CO₂]. Our results indicate that elevated [CO₂] may overall have a limited effect on isoprene emission from young seedlings and that plants generally regulate the emission to reach the maximum at the centre of the leaf profile, irrespective of the total leaf number. In comparison with leaf expansion and photosynthesis, isoprene showed marked and repeatable differences among leaves of the profile and may therefore be a useful trait to accurately monitor changes of leaf ontogeny as a consequence of elevated [CO₂].     

Mesophyll conductance in leaves of Japanese white birch (Betula platyphylla var. japonica) seedlings grown under elevated CO2 concentration and low N availability

To test the hypothesis that mesophyll conductance (g_m) would be reduced by leaf starch accumulation in plants grown under elevated CO₂ concentration [CO₂], we investigated g_m in seedlings of Japanese white birch grown under ambient and elevated [CO₂] with an adequate and limited nitrogen supply using simultaneous gas exchange and chlorophyll fluorescence measurements. Both elevated [CO₂] and limited nitrogen supply decreased area‐based leaf N accompanied with a decrease in the maximum rate of Rubisco carboxylation (V_c,max) on a CO₂ concentration at chloroplast stroma (C_c) basis. Conversely, only seedlings grown at elevated [CO₂] under limited nitrogen supply had significantly higher leaf starch content with significantly lower g_m among the treatment combinations. Based on a leaf anatomical analysis using microscopic photographs, however, there were no significant difference in the area of chloroplast surfaces facing intercellular space per unit leaf area among treatment combinations. Thicker cell walls were suggested in plants grown under limited N by increases in leaf mass per area subtracting non‐structural carbohydrates. These results suggest that starch accumulation and/or thicker cell walls in the leaves grown at elevated [CO₂] under limited N supply might hinder CO₂ diffusion in chloroplasts and cell walls, which would be an additional cause of photosynthetic downregulation as well as a reduction in Rubisco activity related to the reduced leaf N under elevated [CO₂].     

Physiological and Biochemical Characteristics of Sugar Beet Plants Grown at an Increased Carbon Dioxide Concentration and at Various Nitrate Doses

Three-week-old sugar beet (Beta vulgaris L.) seedlings were grown for an additional four weeks under controlled conditions: in river sand watered with a modified Knop mixture containing one half-fold (0.5N), standard (1N), and or threefold (3N) nitrate amount, at the irradiance of 90 W/m² PAR, and at the carbon dioxide concentrations of 0.035% (1C treatment) or 0.07% (2C treatment). The increase in the carbon dioxide concentration and in the nitrogen dose resulted in an increase in the leaf area and the leaf and root dry weight per plant. With the increase in the nitrogen dose, morphological indices characterizing leaf growth increased more noticeably in 1C plants than in 2C plants. And vice versa, the effects of increased CO₂ concentration were reduced with the increase in the nitrogen dose. Roots responded to the changes in the CO₂ and nitrate concentrations otherwise than leaves. At a standard nitrate dose (1N), the contents of proteins and nonstructural carbohydrates (sucrose and starch) in leaves depended little on the CO₂ concentration. At a double CO₂ concentration, the content of chlorophyll somewhat decreased, and the net photosynthesis rate (P _n) calculated per leaf area unit increased. An increase in the nitrogen dose did not affect the leaf carbohydrate content of the 1C and 2C plants except the leaves of the 2C-3N plants, where the carbohydrate content decreased. In 1C and 2C plants, an increase in the nitrogen dose caused an increase in the protein and chlorophyll content. Specific P _n values somewhat decreased in 1C-0.5N plants and had hardly any dependence on the nitrate dose in the 2C plants. The carbohydrate content in roots did not depend on the CO₂ concentration, and the content was the highest at 0.5N. Characteristic nitrogen dose-independent acclimation of photosynthesis to an increased carbon dioxide concentration, which was postulated previously [1], was not observed in our experiments with sugar beet grown at doubled carbon dioxide concentration.     

Involvement of nitrogen and cytokinins in photosynthetic acclimation to elevated CO2 of spring wheat

Acclimation of photosynthetic capacity to elevated CO₂ involves a decrease of the leaf Rubisco content. In the present study, it was hypothesized that nitrogen uptake and partitioning within the leaf and among different aboveground organs affects the down-regulation of Rubisco. Given the interdependence of nitrogen and cytokinin signals at the whole plant level, it is also proposed that cytokinins affect the nitrogen economy of plants under elevated CO₂, and therefore the acclimatory responses. Spring wheat received varying levels of nitrogen and cytokinin in field chambers with ambient (370 μmol mol⁻¹) or elevated (700 μmol mol⁻¹) atmospheric CO₂. Gas exchange, Rubisco, soluble protein and nitrogen contents were determined in the top three leaves in the canopy, together with total nitrogen contents per shoot. Growth in elevated CO₂ induced decreases in photosynthetic capacity only when nitrogen supply was low. However, the leaf contents of Rubisco, soluble protein and total nitrogen on an area basis declined in elevated CO₂ regardless of nitrogen supply. Total nitrogen in the shoot was no lower in elevated than ambient CO₂, but the fraction of this nitrogen located in flag and penultimate leaves was lower in elevated CO₂. Decreased Rubisco: chlorophyll ratios accompanied losses of leaf Rubisco with CO₂ enrichment. Cytokinin applications increased nitrogen content in all leaves and nitrogen allocation to senescing leaves, but decreased Rubisco contents in flag leaves at anthesis and in all leaves 20 days later, together with the amount of Rubisco relative to soluble protein in all leaves at both growth stages. The results suggest that down regulation of Rubisco in leaves at elevated CO₂ is linked with decreased allocation of nitrogen to the younger leaves and that cytokinins cause a fractional decrease of Rubisco and therefore do not alleviate acclimation to elevated CO₂.     

Combined effects of elevated CO2 and soil drought on carbon and nitrogen allocation of the desert shrub Caragana intermedia

Impacts of either elevated CO₂ or drought stress on plant growth have been studied extensively, but interactive effects of these on plant carbon and nitrogen allocation is inadequately understood yet. In this study the response of the dominant desert shrub, Caragana intermedia Kuanget H.c.Fu, to the interaction of elevated CO₂ (700 ± 20 μmol mol⁻¹) and soil drought were determined in two large environmental growth chambers (18 m²). Elevated CO₂ increased the allocation of biomass and carbon into roots and the ratio of carbon to nitrogen (C:N) as well as the leaf soluble sugar content, but decreased the allocation of biomass and carbon into leaves, leaf nitrogen and leaf soluble protein concentrations. Elevated CO₂ significantly decreased the partitioning of nitrogen into leaves, but increased that into roots, especially under soil drought. Elevated CO₂ significantly decreased the carbon isotope discrimination (Δ) in leaves, but increased them in roots, and the ratio of Δ values between root and leaf, indicating an increased allocation into below-ground parts. It is concluded that stimulation of plant growth by CO₂ enrichment may be negated under soil drought, and under the future environment, elevated CO₂ may partially offset the negative effects of enhanced drought by regulating the partitioning of carbon and nitrogen.     

Altered physiological function,not structure,drives increased radiation-use efficiency of soybean grown at elevated CO<Subscript>2</Subscript>

Previous studies of elevated carbon dioxide concentration ([CO₂]) on crop canopies have found that radiation-use efficiency is increased more than radiation-interception efficiency. It is assumed that increased radiation-use efficiency is due to changes in leaf-level physiology; however, canopy structure can affect radiation-use efficiency if leaves are displayed in a manner that optimizes their physiological capacity, even though the canopy intercepts the same amount of light. In order to determine the contributions of physiology and canopy structure to radiation-use and radiation-interception efficiency, this study relates leaf-level physiology and leaf display to photosynthetic rate of the outer canopy. We used a new imaging approach that delivers three-dimensional maps of the outer canopy during the growing season. The 3D data were used to model leaf orientation and mean photosynthetic electron transport of the outer canopy to show that leaf orientation changes did not contribute to increased radiation-use; i.e. leaves of the outer canopy showed similar diurnal leaf movements and leaf orientation in both treatments. Elevated [CO₂] resulted in an increased maximum electron transport rate (ETR_max) of light reactions of photosynthesis. Modeling of canopy light interception showed that stimulated leaf-level electron transport at elevated [CO₂], and not alterations in leaf orientation, was associated with stimulated radiation-use efficiency and biomass production in elevated [CO₂]. This study provides proof of concept of methodology to quantify structure–function relationships in combination, allowing a quantitative estimate of the contribution of both effects to canopy energy conversion under elevated [CO₂].     

Interacting effects of elevated CO2, nutrient availability and plant species on a generalist invertebrate herbivore

By affecting plant growth and phytochemistry elevated CO₂ may have indirect effects on the performance of herbivores. These effects show considerable variability across studies and may depend on nutrient availability, the carbon/nutrient‐balance in plant tissues and the secondary metabolism of plants. We studied the responses to elevated CO₂ and different nutrient availability of 12 herbaceous plant species differing in their investment into secondary compounds. Caterpillars of the generalist herbivore Spodoptera littoralis were reared on the leaves produced and their consumption and growth rates analysed. Elevated CO₂ resulted in a similar increase of biomass in all plant species, whereas the positive effect of fertilization varied among plant species. Specific leaf weight was influenced by elevated CO₂, but the effect depended on nutrient level and identity of plant species. Elevated CO₂ increased the C/N ratio of the leaves of most species. Caterpillars consumed more leaf material when plants were grown under elevated CO₂ and low nutrients. This indicates compensatory feeding due to lower tissue quality. However, the effects of elevated CO₂, nutrient availability and plant species identity on leaf consumption interacted. Both the effects of CO₂ and nutrient availability on the relative growth rate of the herbivore depended on the plant species. The feeding rate of S. littoralis on plant species that do not produce nitrogen‐containing secondary compounds (NCSC) was higher under low nutrient availability. In contrast, in plants producing NCSC nutrient availability had no effect on the feeding rate. This suggests that compensatory feeding in response to low nutrient contents may not be possible if plants produce NCSC. We conclude that elevated CO₂ causes species‐specific changes in the quality of plant tissues and consequently in changes in the preferences of herbivores for plant species. This could result in changes in plant community composition.