Acclimation of peanut (Arachis hypogaea L.) leaf photosynthesis to elevated growth CO2 and temperature

Peanut (Arachis hypogaea L. cv. Florunner) was grown from seed sowing to plant maturity under two daytime CO₂ concentrations ([CO₂]) of 360 μmol mol⁻¹ (ambient) and 720 μmol mol⁻¹ (elevated) and at two temperatures of 1.5 and 6.0 °C above ambient temperature. The objectives were to characterize peanut leaf photosynthesis responses to long-term elevated growth [CO₂] and temperature, and to assess whether elevated [CO₂] regulated peanut leaf photosynthetic capacity, in terms of activity and protein content of ribulose bisphosphate carboxylase-oxygenase (Rubisco), Rubisco photosynthetic efficiency, and carbohydrate metabolism. At both growth temperatures, leaves of plants grown under elevated [CO₂] had higher midday photosynthetic CO₂ exchange rate (CER), lower transpiration and stomatal conductance and higher water-use efficiency, compared to those of plants grown at ambient [CO₂]. Both activity and protein content of Rubisco, expressed on a leaf area basis, were reduced at elevated growth [CO₂]. Declines in Rubisco under elevated growth [CO₂] were 27–30% for initial activity, 5–12% for total activity, and 9–20% for protein content. Although Rubisco protein content and activity were down-regulated by elevated [CO₂], Rubisco photosynthetic efficiency, the ratio of midday light-saturated CER to Rubisco initial or total activity, of the elevated-[CO₂] plants was 1.3- to 1.9-fold greater than that of the ambient-[CO₂] plants at both growth temperatures. Leaf soluble sugars and starch of plants grown at elevated [CO₂] were 1.3- and 2-fold higher, respectively, than those of plants grown at ambient [CO₂]. Under elevated [CO₂], leaf soluble sugars and starch, however, were not affected by high growth temperature. In contrast, high temperature reduced leaf soluble sugars and starch of the ambient-[CO₂] plants. Activity of sucrose-P synthase, but not adenosine 5′-diphosphoglucose pyrophosphorylase, was up-regulated under elevated growth [CO₂]. Thus, in the absence of other environmental stresses, peanut leaf photosynthesis would perform well under rising atmospheric [CO₂] and temperature as predicted for this century.     

Root-zone temperature effects on photosynthesis, <Superscript>14</Superscript>C-photoassimilate partitioning and growth of temperate lettuce (<Emphasis Type="Italic">Lactuca sativa</Emphasis> cv. ‘Panama’) in the tropics

The effect of root growth temperature on maximal photosynthetic CO₂ assimilation (P _max), carbohydrate content, ¹⁴C-photoassimilate partitioning, growth, and root morphology of lettuce was studied after transfer of the root system from cool root-zone temperature (C-RZT) of 20 °C to hot ambient-RZT (A-RZT) and vice versa. Four days after RZT transfer, P _max and leaf total soluble sugar content were highest and lowest, respectively, in C-RZT and A-RZT plants. P _max and total leaf soluble sugar content were much lower in plants transferred from C-to A-RZT (C→A-RZT) than in C-RZT plants. However, these two parameters were much higher in plants transferred from A-to C-RZT (A→C-RZT) than in A-RZT plants. A-RZT and C→A-RZT plants had higher root total soluble sugar content than A→C-RZT and C-RZT plants. Leaf total insoluble sugar content was similar in leaves of all plants while it was the highest in the roots of C-RZT plants. Developing leaves of C-RZT plants had higher ¹⁴C-photoassimilate content than A-RZT plants. The A→C-RZT plants also had higher ¹⁴C-photoassimilate content in their developing leaves than A-RZT plants. However, more ¹⁴C-photoassimilates were translocated to the roots of A-RZT and C→A-RZT plants, but they were mainly used for root thickening than for its elongation. Increases in leaf area, shoot and root fresh mass were slower in C→A-RZT than in C-RZT plants. Conversely, A→C-RZT plants had higher increases in these parameters than A-RZT plants. Lower root/shoot ratio (R/S) in C-RZT than in A-RZT plants confirmed that more photoassimilates were channelled to the shoots than to the roots of C-RZT plants. Roots of C-RZT plants had greater total length with a greater number of tips and surface area, and smaller average diameter as compared to A-RZT plants. In C→A-RZT plants, there was root thickening but the increases in its length, tip number and surface area decreased. The reverse was observed for A→C-RZT plants. These results further supported the idea that newly fixed photoassimilates contributed more to root thickening than to root elongation in A-RZT and C→A-RZT plants.     

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].     

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.     

Nitrogen assimilation and transpiration: key processes conditioning responsiveness of wheat to elevated [CO2] and temperature

Although climate scenarios have predicted an increase in [CO₂] and temperature conditions, to date few experiments have focused on the interaction of [CO₂] and temperature effects in wheat development. Recent evidence suggests that photosynthetic acclimation is linked to the photorespiration and N assimilation inhibition of plants exposed to elevated CO₂. The main goal of this study was to analyze the effect of interacting [CO₂] and temperature on leaf photorespiration, C/N metabolism and N transport in wheat plants exposed to elevated [CO₂] and temperature conditions. For this purpose, wheat plants were exposed to elevated [CO₂] (400 vs 700 µmol mol^?1) and temperature (ambient vs ambient + 4°C) in CO₂ gradient greenhouses during the entire life cycle. Although at the agronomic level, elevated temperature had no effect on plant biomass, physiological analyses revealed that combined elevated [CO₂] and temperature negatively affected photosynthetic performance. The limited energy levels resulting from the reduced respiratory and photorespiration rates of such plants were apparently inadequate to sustain nitrate reductase activity. Inhibited N assimilation was associated with a strong reduction in amino acid content, conditioned leaf soluble protein content and constrained leaf N status. Therefore, the plant response to elevated [CO₂] and elevated temperature resulted in photosynthetic acclimation. The reduction in transpiration rates induced limitations in nutrient transport in leaves of plants exposed to elevated [CO₂] and temperature, led to mineral depletion and therefore contributed to the inhibition of photosynthetic activity.     

Influence of carbon dioxide enrichment, ozone and nitrogen fertilization on cotton (Gossypium hirsutum L.) leaf and root composition

The objective of this study was to test whether elevated [CO₂], [O₃] and nitrogen (N) fertility altered leaf mass per area (LMPA), non‐structural carbohydrate (TNC), N, lignin (LTGA) and proanthocyanidin (PA) concentrations in cotton (Gossypium hirsutum L.) leaves and roots. Cotton was grown in 14 dm³ pots with either sufficient (0·8 g N dm ^{? 3}) or deficient (0·4 and 0·2 g N dm ^{? 3}) N fertilization, and treated in open‐top chambers with either ambient or elevated ( + 175 and + 350 μ mol mol ^{? 1}) [CO₂] in combination with either charcoal‐filtered air (CF) or non‐filtered air plus 1·5 times ambient [O₃]. At about 50 d after planting, LMPA, starch and PA concentrations in canopy leaves were as much as 51–72% higher in plants treated with elevated [CO₂] compared with plants treated with ambient [CO₂], whereas leaf N concentration was 29% lower in elevated [CO₂]‐treated plants compared with controls. None of the treatments had a major effect on LTGA concentrations on a TNC‐free mass basis. LMPA and starch levels were up to 48% lower in plants treated with elevated [O₃] and ambient [CO₂] compared with CF controls, although the elevated [O₃] effect was diminished when plants were treated concurrently with elevated [CO₂]. On a total mass basis, leaf N and PA concentrations were higher in samples treated with elevated [O₃] in ambient [CO₂], but the difference was much reduced by elevated [CO₂]. On a TNC‐free basis, however, elevated [O₃] had little effect on tissue N and PA concentrations. Fertilization treatments resulted in higher PA and lower N concentrations in tissues from the deficient N fertility treatments. The experiment showed that suppression by elevated [O₃] of LMPA and starch was largely prevented by elevated [CO₂], and that interpretation of [CO₂] and [O₃] effects should include comparisons on a TNC‐free basis. Overall, the experiment indicated that allocation to starch and PA may be related to how environmental factors affect source–sink relationships in plants, although the effects of elevated [O₃] on secondary metabolites differed in this respect.     

Infection with the parasitic angiosperm Striga hermonthica influences the response of the C3 cereal Oryza sativa to elevated CO2

Upland rice (Oryza sativa L.) was grown at both ambient (350 μmol mol^?1) and elevated (700 μmol mol^?1) CO₂ in either the presence or absence of the root hemi‐parasitic angiosperm Striga hermonthica (Del) Benth. Elevated CO₂ alleviated the impact of the parasite on host growth: biomass of infected rice grown at ambient CO₂ was 35% that of uninfected, control plants, while at elevated CO₂, biomass of infected plants was 73% that of controls. This amelioration occurred despite the fact that O. sativa grown at elevated CO₂ supported both greater numbers and a higher biomass of parasites per host than plants grown at ambient CO₂. The impact of infection on host leaf area, leaf mass, root mass and reproductive tissue mass was significantly lower in plants grown at elevated as compared with ambient CO₂. There were significant CO₂ and Striga effects on photosynthetic metabolism and instantaneous water‐use efficiency of O. sativa. The response of photosynthesis to internal [CO₂] (A/C_i curves) indicated that, at 45 days after sowing (DAS), prior to emergence of the parasites, uninfected plants grown at elevated CO₂ had significantly lower CO₂ saturated rates of photosynthesis, carboxylation efficiencies and ribulose‐1,5‐bisphosphate carboxylase/oxygenase (Rubisco; EC 4.1.1.39) contents than uninfected, ambient CO₂‐grown O. sativa. In contrast, infection with S. hermonthica prevented down‐regulation of photosynthesis in O. sativa grown at elevated CO₂, but had no impact on photosynthesis of hosts grown at ambient CO₂. At 76 DAS (after parasites had emerged), however, infected plants grown at both elevated and ambient CO₂ had lower carboxylation efficiencies and Rubisco contents than uninfected O. sativa grown at ambient CO₂. The reductions in carboxylation efficiency (and Rubisco content) were accompanied by similar reductions in nitrogen concentration of O. sativa leaves, both before and after parasite emergence. There were no significant CO₂ or infection effects on the concentrations of soluble sugars in leaves of O. sativa, but starch concentration was significantly lower in infected plants at both CO₂ concentrations. These results demonstrate that elevated CO₂ concentrations can alleviate the impact of infection with Striga on the growth of C₃ hosts such as rice and also that infection can delay the onset of photosynthetic down‐regulation in rice grown at elevated CO₂.     

Altered physiological and growth responses to elevated [CO2] in offspring from holm oak (Quercus ilex L.) mother trees with lifetime exposure to naturally elevated [CO2]

An important question with respect to plant performance in future climatic scenarios is whether the offspring of mature trees that have experienced lifelong exposure to elevated [CO₂] show altered physiological responses to elevated [CO₂] compared with those originating from current ambient CO₂ concentrations. To investigate this question, acorns were collected from two seed sources, denoted as ‘control’ and ‘spring’, from Quercus ilex mother trees grown at ambient (36 Pa) and at about twice ambient CO₂ concentrations, respectively, close to a natural CO₂ spring, Laiatico, central Italy. The seedlings were raised for 8 months under controlled conditions at ambient and elevated [CO₂] in a reciprocal experimental design and were used for the determination of biomass, photosynthesis and foliar carbohydrate concentrations, as well as the accumulation of structural biomass and lignin during leaf maturation. Under ambient [CO₂], biomass and foliar carbon acquisition in control progeny were not significantly different from spring progeny. However, under elevated [CO₂], spring seedlings showed less CO₂ acclimation than control seedlings but no significant differences in non‐structural carbohydrate concentrations and structural biomass per unit leaf dry mass. Developmental lignin accumulation in leaves was delayed under elevated [CO₂] compared with ambient [CO₂], but only in control progeny. Under elevated [CO₂], whole‐plant biomass, leaf area and stem diameter were significantly increased in Quercus ilex seedlings from both seed sources but with a higher stimulation of above‐ground biomass in spring than in control seedlings and a higher stimulation of below‐ground biomass in control seedlings. These results indicate that life history and/or progeny may determine the species‐specific CO₂ response and suggest that positive CO₂ acclimation is possible.     

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].     

Acclimation of photosynthesis to elevated CO2 and temperature in five British native species of contrasting functional type

Acclimation of photosynthesis to growth at elevated CO₂ concentration varies markedly between species. Species functionally classified as stress-tolerators (S) and ruderals (R), are thought to be incapable, or the least capable, of responding positively in terms of growth to elevated [CO₂]. Is this pattern of response also apparent in leaf photosynthesis of wild S- and R-strategists? Acclimatory loss of a photosynthetic and growth response to elevated [CO₂] is assumed to reflect limitation on capacity to utilize additional photosynthate. The doubling of pre-industrial global [CO₂] is expected to coincide with a 3 °C increase in mean temperature which could stimulate growth; will photosynthetic capacity at elevated [CO₂] be greater when the concurrent temperature increase is simulated? Five species from natural grassland of NW Europe and of contrasting ecological strategy were grown in hemispherical greenhouses, environmentally controlled to track the external microclimate. Within a replicated design, plants were grown at (i) current ambient [CO₂] and temperature, (ii) elevated [CO₂] (ambient + 340 μmol mol^–1) and ambient temperature, (iii) ambient [CO₂] and elevated temperature (ambient + 3 °C), or (iv) elevated [CO₂] and elevated temperature. After 75–104 days, the CO₂ response of light-saturated rates of photosynthesis (A_sat) was analysed in controlled-environment cuvettes in a field laboratory. There was no acclimatory loss of photosynthetic capacity with growth in elevated [CO₂] or elevated temperature over this period in Poa alpina (S), Bellis perennis (R) or Plantago lanceolata (mixed C-S-R strategist), and a significant (P ? ? bl 0.05) increase in capacity in Helianthemum nummularium (S) and Poa annua (R). Photosynthetic rates of leaves grown and measured in elevated [CO₂] were therefore significantly higher than rates for leaves grown and measured in ambient [CO₂], for all species. With the exception of Poa alpina, stomatal conductance and stomatal limitation on A_sat showed no acclimatory response to growth in elevated [CO₂]. Carboxylation efficiency, determined from the initial slope of the response of A_sat to intercellular CO₂ concentration was significantly increased by elevated [CO₂] and elevated temperature in H.nummularium, implying a possible increase in in vivo RubisCO activity. Increased carboxylation efficiency of this species was also reflected by an increase in the CO₂- and light-saturated rates of photosynthesis, indicating an increased capacity for regeneration of the primary CO₂ acceptor in photosynthesis. The results show that R-strategists and slow-growing S-strategists, are inherently capable of large increases in leaf photosynthetic capacity with growth in elevated [CO₂] in contrast to expectations from growth studies. With the exception of P.annua, where there was a significant negative interaction between CO₂ and temperature, concurrent increase in growth temperature had little effect on this pattern of response.     

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].     

Responses of Senna reticulata,a legume tree from the Amazonian floodplains,to elevated atmospheric CO2 concentration and waterlogging

Key message

The Amazonian tree Senna reticulata showed an increase in photosynthesis and starch content under elevated [CO ₂ ] that led an increment in biomass after 90 days. Elevated [CO ₂ ] was also capable of reducing the negative effect of waterlogging.

Abstract

Tree species from the Amazonian floodplains have to cope with low oxygen availability due to annual pulses of inundation that can last up to 7 months. Species capable of adapting to flooding and/or waterlogged conditions usually partition their storage to favor starch and allocate it to roots, where carbohydrates are used to maintain respiration rates during waterlogging. In spite of climate change, virtually nothing is known about how elevated atmospheric CO₂ concentration ([CO₂]) will affect plants when combined with waterlogging. In this work, we used open top chambers to evaluate the effect of elevated [CO₂] during a period of terrestrial phase and in subsequent combination with waterlogged conditions to determine if the surplus carbon provided by elevated [CO₂] may improve the waterlogging tolerance of the fast-growing Amazonian legume tree Senna reticulata. During the terrestrial phase, photosynthesis was ca. 28 % higher after 30, 45 and 120 days of elevated [CO₂], and starch content in the leaves was, on average, 49 % higher than with ambient [CO₂]. Total biomass was inversely correlated to the starch content of leaves, indicating that starch might be the main carbohydrate source for biomass production during the terrestrial phase. This response was more pronounced under elevated [CO₂], resulting in 30 % more biomass in comparison to ambient [CO₂] plants. After 135 days at elevated [CO₂] an inversion has been observed in total biomass accumulation, in which ambient [CO₂] presented a greater increment in total biomass in comparison to elevated [CO₂], indicating negative effects on growth after long-term CO₂ exposure. However, plants with elevated [CO₂]/waterlogged displayed a greater increment in biomass in comparison with ambient [CO₂]/waterlogged that, unlike during the terrestrial phase, was unrelated to starch reserves. We conclude that S. reticulata displays mechanisms that make this species capable of responding positively to elevated [CO₂] during the first pulse of growth. This response capacity is also associated with a “buffering effect” that prevents the plants from decreasing their biomass under waterlogged conditions.     

Elevated [CO2] and increased N supply reduce leaf disease and related photosynthetic impacts on Solidago rigida

To evaluate whether leaf spot disease and related effects on photosynthesis are influenced by increased nitrogen (N) input and elevated atmospheric CO₂ concentration ([CO₂]), we examined disease incidence and photosynthetic rate of Solidago rigida grown in monoculture under ambient or elevated (560 μmol mol⁻¹) [CO₂] and ambient or elevated (+4 g N m⁻² year⁻¹) N conditions in a field experiment in Minnesota, USA. Disease incidence was lower in plots with either elevated [CO₂] or enriched N (−57 and −37%, respectively) than in plots with ambient conditions. Elevated [CO₂] had no significant effect on total plant biomass, or on photosynthetic rate, but reduced tissue%N by 13%. In contrast, N fertilization increased both biomass and total plant N by 70%, and as a consequence tissue%N was unaffected and photosynthetic rate was lower on N fertilized plants than on unfertilized plants. Regardless of treatment, photosynthetic rate was reduced on leaves with disease symptoms. On average across all treatments, asymptomatic leaf tissue on diseased leaves had 53% lower photosynthetic rate than non-diseased leaves, indicating that the negative effect from the disease extended beyond the visual lesion area. Our results show that, in this instance, indirect effects from elevated [CO₂], i.e., lower disease incidence, had a stronger effect on realized photosynthetic rate than the direct effect of higher [CO₂].     

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.     

Effects of elevated CO2 and temperature on photosynthesis and Rubisco in rice and soybean

Rice (Oryza sativa L. cv. IR-72) and soybean (Glycine max L. Merr. cv. Bragg), which have been reported to differ in acclimation to elevated CO₂, were grown for a season in sunlight at ambient and twice-ambient [CO₂], and under daytime temperature regimes ranging from 28 to 40°C. The objectives of the study were to test whether CO₂ enrichment could compensate for adverse effects of high growth temperatures on photosynthesis, and whether these two C₃ species differed in this regard. Leaf photosynthetic assimilation rates (A) of both species, when measured at the growth [CO₂], were increased by CO₂ enrichment, but decreased by supraoptimal temperatures. However, CO₂ enrichment more than compensated for the temperature-induced decline in A. For soybean, this CO₂ enhancement of A increased in a linear manner by 32–95% with increasing growth temperatures from 28 to 40°C, whereas with rice the degree of enhancement was relatively constant at about 60%, from 32 to 38°C. Both elevated CO₂ and temperature exerted coarse control on the Rubisco protein content, but the two species differed in the degree of responsiveness. CO₂ enrichment and high growth temperatures reduced the Rubisco content of rice by 22 and 23%, respectively, but only by 8 and 17% for soybean. The maximum degree of Rubisco down-regulation appeared to be limited, as in rice the substantial individual effects of these two variables, when combined, were less than additive. Fine control of Rubisco activation was also influenced by both elevated [CO₂] and temperature. In rice, total activity and activation were reduced, but in soybean only activation was lowered. The apparent catalytic turnover rate (K_cat) of rice Rubisco was unaffected by these variables, but in soybean elevated [CO₂] and temperature increased the apparent K_cat by 8 and 22%, respectively. Post-sunset declines in Rubisco activities were accelerated by elevated [CO₂] in rice, but by high temperature in soybean, suggesting that [CO₂] and growth temperature influenced the metabolism of 2-carboxyarabinitol-1-phosphate, and that the effects might be species-specific. The greater capacity of soybean for CO₂ enhancement of A at supraoptimal temperatures was probably not due to changes in stomatal conductance, but may be partially attributed to less down-regulation of Rubisco by elevated [CO₂] in soybean than in rice. However, unidentified species differences in the temperature optimum for photosynthesis also appeared to be important. The responses of photosynthesis and Rubisco in rice and soybean suggest that among C₃ plants species-specific differences will be encountered as a result of future increases in global [CO₂] and air temperatures.     

Seasonal patterns of photosynthetic response and acclimation to elevated carbon dioxide in field-grown strawberry

Strawberry (Fragaria × ananassa) plants were grown in field plots at the current ambient [CO₂], and at ambient + 300 and ambient + 600 μmol mol⁻¹ [CO₂]. Approximately weekly measurements were made of single leaf gas exchange of upper canopy leaves from early spring through fall of two years, in order to determine the temperature dependence of the stimulation of photosynthesis by elevated [CO₂], whether growth at elevated [CO₂] resulted in acclimation of photosynthesis, and whether any photosynthetic acclimation was reduced when fruiting created additional demand for the products of photosynthesis. Stimulation of photosynthetic CO₂ assimilation by short-term increases in [CO₂] increased strongly with measurement temperature. The stimulation exceeded that predicted from the kinetic characteristics of ribulose-1,5-bisphosphate carboxylase at all temperatures. Acclimation of photosynthesis to growth at elevated [CO₂] was evident from early spring through summer, including the fruiting period in early summer, with lower rates under standard measurement conditions in plants grown at elevated [CO₂]. The degree of acclimation increased with growth [CO₂]. However, there were no significant differences between [CO₂] treatments in total nitrogen per leaf area, and photosynthetic acclimation was reversed one day after switching the [CO₂] treatments. Tests showed that acclimation did not result from a limitation of photosynthesis by triose phosphate utilization rate at elevated [CO₂]. Photosynthetic acclimation was not evident during dry periods in midsummer, when the elevated [CO₂] treatments conserved soil water and photosynthesis declined more at ambient than at elevated [CO₂]. Acclimation was also not evident during the fall, when plants were vegetative, despite wet conditions and continued higher leaf starch content at elevated [CO₂]. Stomatal conductance responded little to short-term changes in [CO₂] except during drought, and changed in parallel with photosynthetic acclimation through the seasons in response to the long-term [CO₂] treatments. The data do not support the hypothesis that source-sink balance controls the seasonal occurrence of photosynthetic acclimation to elevated [CO₂] in this species. This revised version was published online in June 2006 with corrections to the Cover Date.     

Elevated CO2 and water deficit effects on photosynthesis, ribulose bisphosphate carboxylase‐oxygenase, and carbohydrate metabolism in rice

Rice (Oryza sativa[L.] cv. IR-72) was grown for a season in sunlit, controlled-environment chambers at 350 or 700 µmol CO₂ mol^?1 under continuously flooded (unstressed) or drought-imposed periods at panicle initiation (stressed). The midday canopy photosynthetic rates (P_n), measured at the CO₂ concentration ([CO₂]) used for growth, were enhanced by high [CO₂] but reduced by drought. High [CO₂] increased P_n by 18 to 34% for the unstressed plants, and 6 to 12% for the stressed plants. In the unstressed plants, CO₂ enrichment increased water-use efficiency (WUE) by 26%, and reduced evapotranspiration (ET) by 8 to 14%. Both high [CO₂] and severe drought decreased the activity and content of ribulose bisphosphate carboxylase-oxygenase (Rubisco). High-CO₂-unstressed plants had 6 to 22% smaller content and 5 to 25%, lower activity of Rubisco than ambient-CO₂-unstressed plants. Under severe drought, reductions of Rubisco were 53 and 27% in activity and 40 and 12% in content, respectively, for ambient- and high-CO₂ treatments. The apparent catalytic turnover rate (K_cat) of midday fully activated Rubisco was not altered by high [CO₂], but severe drought reduced K_cat by 17 to 23%. Chloroplasts of the high-CO₂ leaves contained more, and larger starch grains than those of the ambient CO₂ leaves. High [CO₂] did not affect the leaf sucrose content of unstressed plants. In contrast, severe drought reduced the leaf starch and increased the sucrose content in both CO₂ treatments. The activity of leaf sucrose phosphate synthase of unstressed plants was not affected by high [CO₂], whereas that of ambient-CO₂-grown plants was reduced 45% by severe drought. Reduction in ET and enhancements in both P_n and WUE for rice grown under high [CO₂] helped to delay the adverse effects of severe drought and allowed the stressed plants to assimilate CO₂ for an extra day. Thus, rice grown in the next century may utilize less water, use water more efficiently, and be able to tolerate drought better under some situations.     

Canopy warming caused photosynthetic acclimation and reduced seed yield in maize grown at ambient and elevated [CO2]

Rising atmospheric CO₂ concentration ([CO₂]) and attendant increases in growing season temperature are expected to be the most important global change factors impacting production agriculture. Although maize is the most highly produced crop worldwide, few studies have evaluated the interactive effects of elevated [CO₂] and temperature on its photosynthetic physiology, agronomic traits or biomass, and seed yield under open field conditions. This study investigates the effects of rising [CO₂] and warmer temperature, independently and in combination, on maize grown in the field throughout a full growing season. Free‐air CO₂ enrichment (FACE) technology was used to target atmospheric [CO₂] to 200 μmol mol^?1 above ambient [CO₂] and infrared heaters to target a plant canopy increase of 3.5 °C, with actual season mean heating of ~2.7 °C, mimicking conditions predicted by the second half of this century. Photosynthetic gas‐exchange parameters, leaf nitrogen and carbon content, leaf water potential components, and developmental measurements were collected throughout the season, and biomass and yield were measured at the end of the growing season. As predicted for a C₄ plant, elevated [CO₂] did not stimulate photosynthesis, biomass, or yield. Canopy warming caused a large shift in aboveground allocation by stimulating season‐long vegetative biomass and decreasing reproductive biomass accumulation at both CO₂ concentrations, resulting in decreased harvest index. Warming caused a reduction in photosynthesis due to down‐regulation of photosynthetic biochemical parameters and the decrease in the electron transport rate. The reduction in seed yield with warming was driven by reduced photosynthetic capacity and by a shift in aboveground carbon allocation away from reproduction. This field study portends that future warming will reduce yield in maize, and this will not be mitigated by higher atmospheric [CO₂] unless appropriate adaptation traits can be introduced into future cultivars.     

How is the relationship between the C4 cereal Sorghum bicolor and the C3 root hemi-parasites Striga hermonthica and Striga asiatica affected by elevated CO2?

The C₄ cereal Sorghum bicolor was grown under either ambient (350 μmol mol^?1) or elevated (700 μmol mol^?1) [CO₂] in either the presence or absence of the C₃ obligate root hemi-parasites Striga hermonthica or S. asiatica. Both uninfected and infected sorghum plants were taller and had greater biomass, photosynthetic rates, water-use efficiencies and leaf areas under elevated compared with ambient [CO₂]. There was no evidence of any downregula-tion of photosynthesis in sorghum grown at elevated [CO₂]. Biomass of infected sorghum was lower under both ambient and elevated [CO₂], and although infected plants were larger under elevated [CO₂] the relative impact of infection on host biomass was either the same (S. asiatica) or only slightly less (S. hermonthica) than under ambient [CO₂]. In contrast, biomass of S. hermonthica and S. asiatica per host was lower under elevated than ambient [CO₂], although rates of photosynthesis were higher at elevated [CO₂] and parasite stomatal conductance was not responsive to [CO₂]. Parasites emerged above-ground and flowered earlier under ambient compared with elevated [CO₂]. It appears that the mechanism(s) by which the parasites affect host growth is (are) relatively insensitive to increased atmospheric [CO₂], although the parasites themselves were adversely affected by growth at elevated [CO₂].