Contrasting crop species responses to CO2 and temperature: rice,soybean and citrus

The continuing increase in atmospheric carbon dioxide concentration ([CO₂]) and projections of possible future increases in global air temperatures have stimulated interest in the effects of these climate variables on plants and, in particular, on agriculturally important food crops. Mounting evidence from many different experiments suggests that the magnitude and even direction of crop responses to [CO₂] and temperature is almost certain to be species dependent and very likely, within a species, to be cultivar dependent. Over the last decade, [CO₂] and temperature experiments have been conducted on several crop species in the outdoor, naturally-sunlit, environmentally controlled, plant growth chambers by USDA-ARS and the University of Florida, at Gainesville, Florida, USA. The objectives for this paper are to summarize some of the major findings of these experiments and further to compare and contrast species responses to [CO₂] and temperature for three diverse crop species: rice (Oryza sativa, L.), soybean (Glycine max, L.) and citrus (various species). Citrus had the lowest growth and photosynthetic rates but under [CO₂] enrichment displayed the greatest percentage increases over ambient [CO₂] control treatments. In all three species the direct effect of [CO₂] enrichment was always an increase in photosynthetic rate. In soybean, photosynthetic rate depended on current [CO₂] regardless of the long-term [CO₂] history of the crop. In rice, photosynthetic rate measured at a common [CO₂], decreased with increasing long-term [CO₂] growth treatment due to a corresponding decline in RuBP carboxylase content and activity. Rice specific respiration decreased from subambient to ambient and superambient [CO₂] due to a decrease in plant tissue nitrogen content and a decline in specific maintenance respiration rate. In all three species, crop water use decreased with [CO₂] enrichment but increased with increases in temperature. For both rice and soybean, [CO₂] enrichment increased growth and grain yield. Rice grain yields declined by roughly 10 % per each 1 °C rise in day/night temperature above 28/21 °C.     

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.     

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].     

Rice yield enhancement by elevated CO2 is reduced in cool weather

The projected increase of atmospheric CO₂ concentration ([CO₂]) is expected to increase rice yield, but little is known of the effects of [CO₂] at low temperature, which is the major constraint to growing rice in cool climates. We grew rice under two levels of [CO₂] (ambient and elevated by 200 μmol mol^?1) and two nitrogen (N) fertilization regimes in northern Japan in 2003 (cool weather) and 2004 (warm weather) in the field in a free‐air CO₂ enrichment (FACE) system. Elevated [CO₂] significantly increased grain yield in both years in both N regimes, but the magnitude of the increase differed between years: 6% in 2003 vs. 17% in 2004, with a significant interaction between [CO₂] and year. This difference resulted from responses of spikelet number and ripening percentage to elevated [CO₂]. Enhancement of dry matter production and N uptake at heading by elevated [CO₂] was smaller in 2003 than in 2004, although at maturity there was no difference between years. No significant interaction between N regime and [CO₂] was detected in yield and yield components. The results suggest that yield gain due to elevated [CO₂] can be reduced by low temperature.     

The growth of soybean under free air [CO<Subscript>2</Subscript>] enrichment (FACE) stimulates photosynthesis while decreasing in vivo Rubisco capacity

Down-regulation of light-saturated photosynthesis (A_sat) at elevated atmospheric CO₂ concentration, [CO₂], has been demonstrated for many C₃ species and is often associated with inability to utilize additional photosynthate and/or nitrogen limitation. In soybean, a nitrogen-fixing species, both limitations are less likely than in crops lacking an N-fixing symbiont. Prior studies have used controlled environment or field enclosures where the artificial environment can modify responses to [CO₂]. A soybean free air [CO₂] enrichment (FACE) facility has provided the first opportunity to analyze the effects of elevated [CO₂] on photosynthesis under fully open-air conditions. Potential ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) carboxylation (V_c,max) and electron transport through photosystem II (J_max) were determined from the responses of A_sat to intercellular [CO₂] (C_i) throughout two growing seasons. Mesophyll conductance to CO₂ (g_m) was determined from the responses of A_sat and whole chain electron transport (J) to light. Elevated [CO₂] increased A_sat by 15–20% even though there was a small, statistically significant, decrease in V_c,max. This differs from previous studies in that V_c,max/J_max decreased, inferring a shift in resource investment away from Rubisco. This raised the C_i at which the transition from Rubisco-limited to ribulose-1,5-bisphosphate regeneration-limited photosynthesis occurred. The decrease in V_c,max was not the result of a change in g_m, which was unchanged by elevated [CO₂]. This first analysis of limitations to soybean photosynthesis under fully open-air conditions reveals important differences to prior studies that have used enclosures to elevate [CO₂], most significantly a smaller response of A_sat and an apparent shift in resources away from Rubisco relative to capacity for electron transport.Abbreviations FACE Free air [CO₂] enrichment - Rubisco Ribulose-1,5-bisphosphate carboxylase/oxygenase - RuBP Ribulose-1,5-bisphosphate - SoyFACE Soybean free air [CO₂] enrichment - VPD Vapor pressure deficit     

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.     

Soybean nodulation and N2 fixation response to drought under carbon dioxide enrichment

The combined effects of carbon dioxide (CO₂) enrichment and water deficits on nodulation and N₂ fixation were analysed in soybean [Glycine max (L.) Merr.]. Two short-term experiments were conducted in greenhouses with plants subjected to soil drying, while exposed to CO₂ atmospheres of either 360 or 700 μmol CO₂ mol^–1. Under drought-stressed conditions, elevated [CO₂] resulted in a delay in the decrease in N₂ fixation rates associated with drying of the soil used in these experiments. The elevated [CO₂] also allowed the plants under drought to sustain significant increases in nodule number and mass relative to those under ambient [CO₂]. The total non-structural carbohydrate (TNC) concentration was lower in the shoots of the plants exposed to drought; however, plants under elevated CO₂ had much higher TNC levels than those under ambient CO₂. For both [CO₂] treatments, drought stress induced a substantial accumulation of TNC in the nodules that paralleled N₂ fixation decline, which indicates that nodule activity under drought may not be carbon limited. Under drought stress, ureide concentration increased in all plant tissues. However, exposure to elevated [CO₂] resulted in substantially less drought-induced ureide accumulation in leaf and petiole tissues. A strong negative correlation was found between ureide accumulation and TNC levels in the leaves. This relationship, together with the large effect of elevated [CO₂] on the decrease of ureide accumulation in the leaves, indicated the importance of ureide breakdown in the response of N₂ fixation to drought and of feedback inhibition by ureides on nodule activity. It is concluded that an important effect of CO₂ enrichment on soybean under drought conditions is an enhancement of photoassimilation, an increased partitioning of carbon to nodules and a decrease of leaf ureide levels, which is associated with sustained nodule growth and N₂ rates under soil water deficits. We suggest that future [CO₂] increases are likely to benefit soybean production by increasing the drought tolerance of N₂ fixation.     

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.     

Smaller than predicted increase in aboveground net primary production and yield of field-grown soybean under fully open-air [CO2] elevation

The Intergovernmental Panel on Climate Change projects that atmospheric [CO₂] will reach 550 ppm by 2050. Numerous assessments of plant response to elevated [CO₂] have been conducted in chambers and enclosures, with only a few studies reporting responses in fully open‐air, field conditions. Reported yields for the world's two major grain crops, wheat and rice, are substantially lower in free‐air CO₂ enrichment (FACE) than predicted from similar elevated [CO₂] experiments within chambers. This discrepancy has major implications for forecasting future global food supply. Globally, the leguminous‐crop soybean (Glycine max (L.) Merr.) is planted on more land than any other dicotyledonous crop. Previous studies have shown that total dry mass production increased on average 37% in response to increasing [CO₂] to approximately 700 ppm, but harvestable yield will increase only 24%. Is this representative of soybean responses under open‐air field conditions? The effects of elevation of [CO₂] to 550 ppm on total production, partitioning and yield of soybean over 3 years are reported. This is the first FACE study of soybean ( http://www.soyface.uiuc.edu ) and the first on crops in the Midwest of North America, one of the major food production regions of the globe. Although increases in both aboveground net primary production (17–18%) and yield (15%) were consistent across three growing seasons and two cultivars, the relative stimulation was less than projected from previous chamber experiments. As in previous studies, partitioning to seed dry mass decreased; however, net production during vegetative growth did not increase and crop maturation was delayed, not accelerated as previously reported. These results suggest that chamber studies may have over‐estimated the stimulatory effect of rising [CO₂], with important implications on global food supply forecasts.     

Interannual climatic variation mediates elevated CO2 and O3 effects on forest growth

We analyzed growth data from model aspen (Populus tremuloides Michx.) forest ecosystems grown in elevated atmospheric carbon dioxide ([CO₂]; 518 μL L^?1) and ozone concentrations ([O₃]; 1.5 × background of 30–40 nL L^?1 during daylight hours) for 7 years using free‐air CO₂ enrichment technology to determine how interannual variability in present‐day climate might affect growth responses to either gas. We also tested whether growth effects of those gasses were sustained over time. Elevated [CO₂] increased tree heights, diameters, and main stem volumes by 11%, 16%, and 20%, respectively, whereas elevated ozone [O₃] decreased them by 11%, 8%, and 29%, respectively. Responses similar to these were found for stand volume and basal area. There were no growth responses to the combination of elevated [CO₂+O₃]. The elevated [CO₂] growth stimulation was found to be decreasing, but relative growth rates varied considerably from year to year. Neither the variation in annual relative growth rates nor the apparent decline in CO₂ growth response could be explained in terms of nitrogen or water limitations. Instead, growth responses to elevated [CO₂] and [O₃] interacted strongly with present‐day interannual variability in climatic conditions. The amount of photosynthetically active radiation and temperature during specific times of the year coinciding with growth phenology explained 20–63% of the annual variation in growth response to elevated [CO₂] and [O₃]. Years with higher photosynthetic photon flux (PPF) during the month of July resulted in more positive growth responses to elevated [CO₂] and more negative growth responses to elevated [O₃]. Mean daily temperatures during the month of October affected growth in a similar fashion the following year. These results indicate that a several‐year trend of increasingly cloudy summers and cool autumns were responsible for the decrease in CO₂ growth response.     

The temporal and species dynamics of photosynthetic acclimation in flag leaves of rice (Oryza sativa) and wheat (Triticum aestivum) under elevated carbon dioxide

In this study, we tested for the temporal occurrence of photosynthetic acclimation to elevated [CO₂] in the flag leaf of two important cereal crops, rice and wheat. In order to characterize the temporal onset of acclimation and the basis for any observed decline in photosynthetic rate, we characterized net photosynthesis, g_s, g_m, C_i/C_a, C_i/C_c, V_cmax, J_max, cell wall thickness, content of Rubisco, cytochrome (Cyt) f, N, chlorophyll and carbohydrate, mRNA expression for rbcL and petA, activity for Rubisco, sucrose phosphate synthase (SPS) and sucrose synthase (SS) at full flag expansion, mid‐anthesis and the late grain‐filling stage. No acclimation was observed for either crop at full flag leaf expansion. However, at the mid‐anthesis stage, photosynthetic acclimation in rice was associated with RuBP carboxylation and regeneration limitations, while wheat only had the carboxylation limitation. By grain maturation, the decline of Rubisco content and activity had contributed to RuBP carboxylation limitation of photosynthesis in both crops at elevated [CO₂]; however, the sharp decrease of Rubisco enzyme activity played a more important role in wheat. Although an increase in non‐structural carbohydrates did occur during these later stages, it was not consistently associated with changes in SPS and SS or photosynthetic acclimation. Rather, over time elevated [CO₂] appeared to enhance the rate of N degradation and senescence so that by late‐grain fill, photosynthetic acclimation to elevated [CO₂] in the flag leaf of either species was complete. These data suggest that the basis for photosynthetic acclimation with elevated [CO₂] may be more closely associated with enhanced rates of senescence, and, as a consequence, may be temporally dynamic, with significant species variation.     

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

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

Increased night temperature reduces the stimulatory effect of elevated carbon dioxide concentration on methane emission from rice paddy soil

To determine how elevated night temperature interacts with carbon dioxide concentration ([CO₂]) to affect methane (CH₄) emission from rice paddy soil, we conducted a pot experiment using four controlled‐environment chambers and imposed a combination of two [CO₂] levels (ambient: 380 ppm; elevated: 680 ppm) and two night temperatures (22 and 32 °C). The day temperature was maintained at 32 °C. Rice (cv. IR72) plants were grown outside until the early‐reproductive growth stage and then transferred to the chambers. After onset of the treatment, day and night CH₄ fluxes were measured every week. The CH₄ fluxes changed significantly with the growth stage, with the largest fluxes occurring around the heading stage in all treatments. The total CH₄ emission during the treatment period was significantly increased by both elevated [CO₂] (P=0.03) and elevated night temperature (P<0.01). Elevated [CO₂] increased CH₄ emission by 3.5% and 32.2% under high and low night temperature conditions, respectively. Elevated [CO₂] increased the net dry weight of rice plants by 12.7% and 38.4% under high and low night temperature conditions, respectively. These results imply that increasing night temperature reduces the stimulatory effect of elevated [CO₂] on both CH₄ emission and rice growth. The CH₄ emission during the day was larger than at night even under the high‐night‐temperature treatment (i.e. a constant temperature all day). This difference became larger after the heading stage. We observed significant correlations between the night respiration and daily CH₄ flux (P<0.01). These results suggest that net plant photosynthesis contributes greatly to CH₄ emission and that increasing night temperature reduces the stimulatory effect of elevated [CO₂] on CH₄ emission from rice paddy soil.     

Yield response of field‐grown soybean exposed to heat waves under current and elevated [CO2]

Elevated atmospheric CO₂ concentration ([CO₂]) generally enhances C₃ plant productivity, whereas acute heat stress, which occurs during heat waves, generally elicits the opposite response. However, little is known about the interaction of these two variables, especially during key reproductive phases in important temperate food crops, such as soybean (Glycine max). Here, we grew soybean under elevated [CO₂] and imposed high‐ (+9°C) and low‐ (+5°C) intensity heat waves during key temperature‐sensitive reproductive stages (R1, flowering; R5, pod‐filling) to determine how elevated [CO₂] will interact with heat waves to influence soybean yield. High‐intensity heat waves, which resulted in canopy temperatures that exceeded optimal growth temperatures for soybean, reduced yield compared to ambient conditions even under elevated [CO₂]. This was largely due to heat stress on reproductive processes, especially during R5. Low‐intensity heat waves did not affect yields when applied during R1 but increased yields when applied during R5 likely due to relatively lower canopy temperatures and higher soil moisture, which uncoupled the negative effects of heating on cellular‐ and leaf‐level processes from plant‐level carbon assimilation. Modeling soybean yields based on carbon assimilation alone underestimated yield loss with high‐intensity heat waves and overestimated yield loss with low‐intensity heat waves, thus supporting the influence of direct heat stress on reproductive processes in determining yield. These results have implications for rain‐fed cropping systems and point toward a climatic tipping point for soybean yield when future heat waves exceed optimum temperature.     

Seasonal changes in the effects of free-air CO2 enrichment (FACE) on growth, morphology and physiology of rice root at three levels of nitrogen fertilization

Over time, the relative effects of elevated [CO₂] on the aboveground photosynthesis, growth and development of rice (Oryza sativa L.) are likely to be changed with increasing duration of CO₂ exposure, but the resultant effects on rice belowground responses remain to be evaluated. To investigate the impacts of elevated [CO₂] on seasonal changes in root growth, morphology and physiology of rice, a free‐air CO₂ enrichment (FACE) experiment was performed at Wuxi, Jiangsu, China, in 2002–2003. A japonica cultivar with large panicle was exposed to two [CO₂] (ambient [CO₂], 370 μmol mol⁻¹; elevated [CO₂], 570 μmol mol⁻¹) at three levels of nitrogen (N): low (LN, 15 g N m⁻²), medium (MN, 25 g N m⁻²) and high N (HN, 35 g N m⁻²). Elevated [CO₂] increased cumulative root volume, root dry weight, adventitious root length and adventitious root number at all developmental stages by 25–71%, which was mainly associated with increased root growth rate during early growth period (EGP) and lower rate of root senescence during late growth period (LGP), while a slight inhibition of root growth rate occurred during middle growth period (MGP). For individual adventitious roots, elevated [CO₂] increased average length, volume, diameter and dry weight early in the season, but the effects gradually disappeared in subsequent stages. Total surface area and active adsorption area per unit root dry weight reached their maxima 10 days earlier in FACE vs. ambient plants, but both of them together with root oxidation ability per unit root dry weight declined with elevated [CO₂] during MGP and LGP, the decline being larger during MGP than LGP. The CO₂‐induced decreases in specific root activities during MGP and LGP were associated with a larger amount of root accumulation during EGP and lower N concentration and higher C/N ratio in roots during MGP and LGP in FACE vs. ambient plants. The results suggest that most of the CO₂‐induced increases in shoot growth of rice are similarly associated with increased root growth.     

Root-zone CO2 and root-zone temperature effects on photosynthesis and nitrogen metabolism of aeroponically grown lettuce (Lactuca sativa L.) in the tropics

Effects of elevated root-zone (RZ) CO₂ concentration (RZ [CO₂]) and RZ temperature (RZT) on photosynthesis, productivity, nitrate (NO₃ ^?), total reduced nitrogen (TRN), total leaf soluble and Rubisco proteins were studied in aeroponically grown lettuce plants in a tropical greenhouse. Three weeks after transplanting, four different RZ [CO₂] concentrations (ambient, 360 ppm, and elevated concentrations of 2,000; 10,000; and 50,000 ppm) were imposed on plants at 20°C-RZT or ambient(A)-RZT (24–38°C). Elevated RZ [CO₂] resulted in significantly higher light-saturated net photosynthetic rate, but lower light-saturated stomatal conductance. Higher elevated RZ [CO₂] also protected plants from both chronic and dynamic photoinhibition (measured by chlorophyll fluorescence Fv/Fm ratio) and reduced leaf water loss. Under each RZ [CO₂], all these variables were significantly higher in 20°C-RZT plants than in A-RZT plants. All plants accumulated more biomass at elevated RZ [CO₂] than at ambient RZ [CO₂]. Greater increases of biomass in roots than in shoots were manifested by lower shoot/root ratios at elevated RZ [CO₂]. Although the total biomass was higher at 20°C-RZT, the increase in biomass under elevated RZ [CO₂] was greater at A-RZT. Shoot NO₃ ^? and TRN concentrations, total leaf soluble and Rubisco protein concentrations were higher in all elevated RZ [CO₂] plants than in plants under ambient RZ [CO₂] at both RZTs. Under each RZ [CO₂], total leaf soluble and Rubisco protein concentrations were significantly higher at 20°C-RZT than at A-RZT. Our results demonstrated that increased P _Nmax and productivity under elevated [CO₂] was partially due to the alleviation of midday water loss, both dynamic and chronic photoinhibition as well as higher turnover of Calvin cycle with higher Rubisco proteins.     

Response of the floating aquatic fern <Emphasis Type="Italic">Azolla filiculoides</Emphasis> to elevated CO<Subscript>2</Subscript>, temperature,and phosphorus levels

Azolla filiculoides is a floating aquatic fern growing in tropical and temperate freshwater ecosystems. As A. filiculoides has symbiotic nitrogen-fixing cyanobacteria (Anabaena azollae) within its leaf cavities, it is cultivated in rice paddies to improve N availability and suppress other wetland weeds. To understand how C assimilation and N accumulation in A. filiculoides respond to elevated atmospheric carbon dioxide concentration (CO₂) in combination with P addition and higher temperatures, we conducted pot experiments during the summer of 2007 and 2008. In 2007, we grew A. filiculoides in pots at two treatment levels of added P fertilizer and at two levels of [CO₂] (380 ppm for ambient and 680 ppm for elevated [CO₂]) in controlled-environment chambers. In 2008, we grew A. filiculoides in four controlled-environment chambers at two [CO₂] levels and two temperature levels (34/26°C (day/night) and 29/21°C). We found that biomass and C assimilation by A. filiculoides were significantly increased by elevated [CO₂], temperature, and P level (all P < 0.01), with a significant interaction between elevated [CO₂] and added P (P < 0.01). Tissue N content was decreased by elevated [CO₂] and increased by higher temperature and P level (all P < 0.01). The acetylene reduction assay showed that the N-fixation activity of A. filiculoides was not significantly different under ambient and elevated [CO₂] but was significantly stimulated by P addition. N-fixation activity decreased at higher temperatures (34/26°C), indicating that 29/21°C was more suitable for A. azollae growth. Therefore, we conclude that the N accumulation potential of A. filiculoides under future climate warming depends primarily on the temperature change and P availability, and C assimilation should be increased by elevated [CO₂].     

Down-regulation of Rubisco activity under combined increases of CO<Subscript>2</Subscript> and temperature minimized by changes in Rubisco k<Subscript>cat</Subscript> in wheat

Increases in growth temperature have been observed to affect photosynthesis differently under long-term exposure to ambient- and twice ambient-air CO₂ concentrations. This study investigates the causes of this interaction in wheat (Triticum aestivum L.) grown in the field over two consecutive years under temperature gradient chambers in ambient (370 μmol mol⁻¹) or elevated (700 μmol mol⁻¹) atmospheric CO₂ concentrations and at ambient or ambient +4°C temperatures, with either a low or a high nitrogen supply. The photosynthesis-internal CO₂ response curves and the activity, activation state, k_cat and amount of Ribulose-1, 5-bisphosphate carboxylase/oxygenase (Rubisco) were measured, as well as the soluble protein concentration in flag leaves at ear emergence and 8–15 days after anthesis. A high nitrogen supply increased V_cmax, the Rubisco amount and activity and soluble protein contents, but did not significantly change the Rubisco k_cat. Both elevated CO₂ and above ambient temperatures had negative effects on V_cmax and Rubisco activity, but at elevated CO₂, an increase in temperature did not decrease V_cmax or Rubisco activity in relation to ambient temperature. The amounts of Rubisco and soluble protein decreased with elevated CO₂ and temperature. The negative impact of elevated CO₂ on Rubisco properties was somewhat counteracted at elevated temperatures by an increase in k_cat. This effect can diminish the detrimental effects on photosynthesis of combined increases of CO₂ and temperature.     

Effects of lifelong [CO2] enrichment on carboxylation and light utilization of Quercus pubescens Willd. examined with gas exchange,biochemistry and optical techniques

Lifelong exposure to elevated concentrations of atmospheric CO₂ may enhance carbon assimilation of trees with unlimited rooting volume and consequently may reduce requirements for photoprotective pigments. In early summer the effects of elevated [CO₂] on carboxylation and light utilization of mature Quercus pubescens trees growing under chronic [CO₂] enrichment at two CO₂ springs and control sites in Italy were examined. Net photosynthesis was enhanced by 36 to 77%. There was no evidence of photosynthetic downregulation early in the growing season when sink demand presumably was greatest. Specifically, maximum assimilation at saturating [CO₂], electron transport capacity, and Rubisco content, activity and carboxylation capacity were not significantly different in trees growing at the CO₂ springs and their respective control sites. Foliar biochemical content, leaf reflectance index of chlorophyll pigments (NDVI), and photochemical efficiency of PSII (ΔF/F_m′) also were not significantly affected by [CO₂] enrichment except that starch content and ΔF/F_m′ tended to be higher at one spring (42 and 15%, respectively). Contrary to expectation, prolonged elevation of [CO₂] did not reduce xanthophyll cycle pigment pools or alter mid‐day values of leaf reflectance index of xanthophyll cycle pigments (PRI), despite the enhancement of carbon assimilation. However, both these pigments and PRI were well correlated with electron transport capacity.