Effects of elevated CO2 concentration and increased temperature on winter wheat: test of ARCWHEAT1 simulation model

Winter wheat (Triticum aestivum L., ev. Mercia) was grown in a controlled-environment facility at two CO₂ concentrations (targets 350 and 700 μmol mol^?1), and two temperature regimes (tracking ambient and ambient + 4°C). Observations of phenology, canopy growth, dry matter production and grain yield were used to test the ARCWHEAT1 simulation model. Dry-matter production and grain yield were increased at elevated CO₂ concentration (27 and 39%, respectively) and reduced at increased temperature (?16 and ?35%, respectively). ARCWHEAT1 substantially underestimated canopy growth for all treatments. However, differences in the facility environment from field conditions over the winter, indicated by the unusually rapid canopy growth observed in this period, meant that empirical model relationships were being used outside the conditions for which they were developed. The ARCWHEAT productivity submodel, given observed green area indices as inputs, overestimated the effect of CO₂ on productivity. An alternative, more mechanistic submodel of productivity, based on the SUCROS87 and Farquhar & von Caemmerer models, simulated observed crop biomass very closely. When these productivity simulations were inputed into the ARCWHEAT1 partitioning and grain-fill submodels, grain yield was predicted poorly, mainly as a result of the assumption that the number of grains is proportional to total growth during a short pre-anthesis phase. While yield was not correlated with growth in this phase, it was correlated with growth in longer pre-anthesis phases, indicating that ARCWHEAT1 could be improved by taking into account the contribution of earlier growth in determining yield.     

Acquisition and within-plant allocation of 13C and 15N in CO2-enriched Quercus robur plants

We assessed the effects of doubling atmospheric CO₂ concentration, [CO₂], on C and N allocation within pedunculate oak plants (Quercus robur L.) grown in containers under optimal water supply. A short-term dual ¹³CO₂ and ¹⁵NO₃^? labelling experiment was carried out when the plants had formed their third growing flush. The 22-week exposure to 700 μl l^?1 [CO₂] stimulated plant growth and biomass accumulation (+53% as compared with the 350 μl l^?1 [CO₂] treatment) but decreased the root/shoot biomass ratio (-23%) and specific leaf area (-18%). Moreover, there was an increase in net CO₂ assimilation rate (+37% on a leaf dry weight basis; +71% on a leaf area basis), and a decrease in both above- and below-ground CO₂ respiration rates (-32 and -26%, respectively, on a dry mass basis) under elevated [CO₂]. ¹³C acquisition, expressed on a plant mass basis or on a plant leaf area basis, was also markedly stimulated under elevated [CO₂] both after the 12-h ¹³CO₂ pulse phase and after the 60-h chase phase. Plant N content was increased under elevated CO₂ (+36%), but not enough to compensate for the increase in plant C content (+53%). Thus, the plant C/N ratio was increased (+13%) and plant N concentration was decreased (-11%). There was no effect of elevated [CO₂] on fine root-specific ¹⁵N uptake (amount of recently assimilated ¹⁵N per unit fine root dry mass), suggesting that modifications of plant N pools were merely linked to root size and not to root function. N concentration was decreased in the leaves of the first and second growing flushes and in the coarse roots, whereas it was unaffected by [CO₂] in the stem and in the actively growing organs (fine roots and leaves of the third growth flush). Furthermore, leaf N content per unit area was unaffected by [CO₂]. These results are consistent with the short-term optimization of N distribution within the plants with respect to growth and photosynthesis. Such an optimization might be achieved at the expense of the N pools in storage compartments (coarse roots, leaves of the first and second growth flushes). After the 60-h ¹³C chase phase, leaves of the first and second growth flushes were almost completely depleted in recent ¹³C under ambient [CO₂], whereas these leaves retained important amounts of recently assimilated ¹³C (carbohydrate reserves?) under elevated [CO₂].     

Growth dynamics and genotypic variation in tropical, field-grown paddy rice (Oryza sativa L.) in response to increasing carbon dioxide and temperature

While previous studies have examined the growth and yield response of rice to continued increases in CO₂ concentration and potential increases in air temperature, little work has focused on the long-term response of tropical paddy rice (i.e. the bulk of world rice production) in situ, or genotypic differences among cultivars in response to increasing CO₂ and/or temperature. At the International Rice Research Institute, rice (cv IR72) was grown from germination until maturity for 4 field seasons, the 1994 and 1995 wet and the 1995 and 1996 dry seasons at three different CO₂ concentrations (ambient, ambient + 200 and ambient + 300 μL L^–1 CO₂) and two air temperatures (ambient and ambient + 4 °C) using open-top field chambers placed within a paddy site. Overall, enhanced levels of CO₂ alone resulted in significant increases in total biomass at maturity and increased seed yield with the relative degree of enhancement consistent over growing seasons across both temperatures. Enhanced levels of temperature alone resulted in decreases or no change in total biomass and decreased seed yield at maturity across both CO₂ levels. In general, simultaneous increases in air temperature as well as CO₂ concentration offset the stimulation of biomass and grain yield compared to the effect of CO₂ concentration alone. For either the 1995 wet and 1996 dry seasons, additional cultivars (N-22, NPT1 and NPT2) were grown in conjunction with IR72 at the same CO₂ and temperature treatments. Among the cultivars tested, N-22 showed the greatest relative response of both yield and biomass to increasing CO₂, while NPT2 showed no response and IR72 was intermediate. For all cultivars, however, the combination of increasing CO₂ concentration and air temperature resulted in reduced grain yield and declining harvest index compared to increased CO₂ alone. Data from these experiments indicate that (a) rice growth and yield can respond positively under tropical paddy conditions to elevated CO₂, but that simultaneous exposure to elevated temperature may negate the CO₂ response to grain yield; and, (b) sufficient intraspecific variation exists among cultivars for future selection of rice cultivars which may, potentially, convert greater amounts of CO₂ into harvestable yield.     

Effects of elevated CO2 and O3 on the rate and duration of grain growth and harvest index in spring wheat (Triticum aestivum L.)

Wheat (Triticum aestivum L.) cv. Minaret was grown in open-top chambers (OTCs) in 1995 and 1996 under three carbon dioxide (CO₂) and two ozone (O₃) levels. Plants were harvested regularly between anthesis and maturity to examine the rate of grain growth (dG/dt; mg d^–1) and the rate of increase in harvest index (dHI/dt;% d^–1). The duration of grain filling was not affected by elevated CO₂ or O₃, but was 12 days shorter in 1995, when the daily mean temperature was over 3 °C higher than in 1996. Season-long exposure to elevated CO₂ (680 μmol mol^–1) significantly increased the rate of grain growth in both years and mean grain weight at maturity (MGW) was up to 11% higher than in the chambered ambient air control (chAA; 383 μmol mol^–1). However, the increase in final yield obtained under elevated CO₂ relative to the chAA control in 1996 resulted primarily from a 27% increase in grain number per unit ground area. dG/dt was significantly reduced by elevated O₃ under ambient CO₂ conditions in 1995, but final grain yield was not affected because of a concurrent increase in grain number. Neither dG/dt nor dHI/dt were affected by the higher mean O₃ concentrations applied in 1996 (77 vs. 66 nmol mol^–1); the differing effects of O₃ on grain growth in 1995 and 1996 observed in both the ambient and elevated CO₂ treatments may reflect the contrasting temperature environments experienced. Grain yield was nevetheless reduced under elevated O₃ in 1996, primarily because of a substantial decrease in grain number. The data obtained show that, although exposure to elevated CO₂ and O₃ individually or in combination may affect both dG/dt and dHI/dt, the presence of elevated CO₂ does not protect against substantial O₃-induced yield losses resulting from its direct deleterious impact on reproductive processes. The implications of these results for food production under future climatic conditions are considered.     

Effects of shading in different developmental phases on biomass and grain yield of winter wheat at ambient and elevated CO2

Winter wheat (Triticum aestivuin cv. Mercia) was grown in a controlled-environment facility under simulated Held conditions at ambient (360μmol mol^?1) and elevated (690 μmol mol^?1) CO₂ concentrations. Some of the plants were shaded to mimic cloudy conditions during three periods of about 20d duration between terminal spikelet and start of grain-fill, giving 16 treatments in all. Elevated CO₂, increased grain yield by about 20%, while shading in any period decreased yield, with the greatest effect in the last period, encompassing anthesis. No interactions between these effects were significant for grain yield, but there were complex interactions for mean grain size. Observed effects of shading and elevated CO₂ on biomass production were well predicted by a simulation model. Observed effects of treatments on yield could be related to effects on biomass using a simple model which assumes that yield is proportional to biomass production, with coefficients of 0.42 (g grain yield g^?1 biomass) for the first two periods and 0.74 for the last period. Wheat models should therefore include developmental changes in sensitivity of yield to biomass production, but biomass changes induced by different CO₂ concentrations or light environments can be treated as having equivalent effects on grain yield.     

Seasonal changes in the response of winter wheat to elevated atmospheric CO2 concentration grown in Open-Top Chambers and field tracking enclosures

Winter wheat was grown at ambient and elevated (ambient plus 350 μL L^–1) CO₂ concentrations in open top chambers and in field-tracking sun-lit climatized enclosures (elevated is 718 μL L^–1). There was no significant effect of CO₂ concentration on sheath, leaf and root biomass and leaf area in the early spring (January to April). 24-h canopy CO₂ exchange rate (CCER) was not significantly affected either. However, elevated CO₂ concentration increased CCER at midday, decreased evapotranspiration rate and increased instantaneous water-use-efficiency during early spring. Leaf, sheath and root nitrogen concentration per unit dry weight decreased and nonstructural carbohydrate concentration increased under elevated CO₂, and N-uptake per unit ground area decreased significantly (– 22%) towards the end of this period. These results contrast with results from the final harvest, when grain yield and biomass were increased by 19% under elevated CO₂. N concentration per dry weight was reduced by 5%, but N-uptake per unit ground area was significantly higher (+ 11%) for the elevated CO₂ treatment. 24-h and midday-CCER increased significantly more in late spring (period of 21 April to 30 May) (respectively by + 40% and 53%) than in the early spring (respectively 5% and 19%) in response to elevated CO₂. Midday evapotranspiration rate was reduced less by elevated CO₂ in the late spring (– 13%) than in early spring (– 21%). The CO₂ response of midday and 24-h CCER decreased again (+ 27% and + 23% resp.) towards the end of the growing season. We conclude that the low response to CO₂ concentration during the early spring was associated with a growth-restriction, caused by low temperature and irradiance levels. The reduction of nitrogen concentration, the increase of nonstructural carbohydrate, and the lower evapotranspiration indicated that CO₂ did have an effect towards the end of early spring, but not on biomass accumulation. Regression analysis showed that both irradiance and temperature affected the response to CO₂.     

Ozone and density affect the response of biomass and seed yield to elevated CO2 in rice

Tropospheric O₃ reduces growth and yield of many crop species, whereas CO₂ ameliorates the negative effects of O₃. Thus, in a combined elevated CO₂ and O₃ atmosphere, seed yield is at least restored to that of charcoal‐filtered (CF) air at ambient CO₂. The CO₂‐induced yield increase in CF air is highly variable, suggesting other potential resource limitations. To understand such variability in response, we tested that (1) competition for resources precludes some of the CO₂ enhancement on biomass and yield; and (2) O₃ reduces competition in elevated CO₂. We grew rice (Oryza sativa L.) at five densities in CF and O₃‐fumigated (+O3) air at ambient (A) and elevated [CO₂] (+CO2) in 1997 and 1998. O₃ reduced biomass by 25% and seed yield by 13–20% in A, but had little effect in +CO2. A competition model of biomass and yield response to density based on resource availability without competition showed that fewer resources were used for biomass in +O3 than in CF (average 53% vs. 70%) in A, while in +CO2 85% of resources were used for biomass regardless of O₃ suggesting greater depletion of resources. The enhanced biomass response to CO₂ with O₃ is consistent with a 22% greater CO₂ enhancement ratio [mass in +CO2 air/mass in A air; enhancement ratio (ER)] in +O3 than in CF air. For seed yield, few resources were used (average 17% and 25% for CF in 1997 and 1998, respectively), and ER was 13% greater in +O3. With competition the rate of change of individual plant biomass to density was not affected by +CO2 in CF air in 1997 but was increased 19% with more nutrients in 1998, indicating resource limitations with +CO2. The rate of change of individual plant yield to density was reduced with CO₂ in 1997 and unchanged in 1998 showing a different response to resource limitation for reproductive biomass. The resource use in +O3‐A suggested that increased density and soil fertility might compensate for pollutant damage. Although ambient [O₃] can modulate the response to elevated CO₂, resource limitation precludes the CO₂ fertilization impact and both factors need consideration for better management and forecasts of future productivity.     

Growth, light interception and yield responses of spring wheat (Triticum aestivum L.) grown under elevated CO2 and O3 in open-top chambers

Spring wheat cv. Minaret was grown to maturity under three carbon dioxide (CO₂) and two ozone (O₃) concentrations in open-top chambers (OTC). Green leaf area index (LAI) was increased by elevated CO₂ under ambient O₃ conditions as a direct result of increases in tillering, rather than individual leaf areas. Yellow LAI was also greater in the 550 and 680 μmol mol^–1 CO₂ treatments than in the chambered ambient control; individual leaves on the main shoot senesced more rapidly under 550 μmol mol^–1 CO₂, but senescence was delayed at 680 μmol mol^–1 CO₂. Fractional light interception (f) during the vegetative period was up to 26% greater under 680 μmol mol^–1 CO₂ than in the control treatment, but seasonal accumulated intercepted radiation was only increased by 8%. As a result of greater carbon assimilation during canopy development, plants grown under elevated CO₂ were taller at anthesis and stem and ear biomass were 27 and 16% greater than in control plants. At maturity, yield was 30% greater in the 680 μmol mol^–1 CO₂ treatment, due to a combination of increases in the number of ears per m^–2, grain number per ear and individual grain weight (IGW). Exposure to a seasonal mean (7 h d^–1) of 84 nmol mol^–1 O₃ under ambient CO₂ decreased green LAI and increased yellow LAI, thereby reducing both f and accumulated intercepted radiation by ≈ 16%. Individual leaves senesced completely 7–28 days earlier than in control plants. At anthesis, the plants were shorter than controls and exhibited reductions in stem and ear biomass of 15 and 23%. Grain yield at maturity was decreased by 30% due to a combination of reductions in ear number m^–2, the numbers of grains per spikelet and per ear and IGW. The presence of elevated CO₂ reduced the rate of O₃-induced leaf senescence and resulted in the maintenance of a higher green LAI during vegetative growth under ambient CO₂ conditions. Grain yields at maturity were nevertheless lower than those obtained in the corresponding elevated CO₂ treatments in the absence of elevated O₃. Thus, although the presence of elevated CO₂ reduced the damaging impact of ozone on radiation interception and vegetative growth, substantial yield losses were nevertheless induced. These data suggest that spring wheat may be susceptible to O₃-induced injury during anthesis irrespective of the atmospheric CO₂ concentration. Possible deleterious mechanisms operating through effects on pollen viability, seed set and the duration of grain filling are discussed.     

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.     

Seasonal CO2 exchange patterns of developing peach (Prunus persica) fruits in response to temperature, light and CO2 concentration

CO₂ exchange rates per unit dry weight, measured in the field on attached fruits of the late-maturing Cal Red peach cultivar, at 1200 μmol photons m^?2S^?1 and in dark, and photosynthetic rates, calculated by the difference between the rates of CO₂ evolution in light and dark, declined over the growing season. Calculated photosynthetic rates per fruit increased over the season with increasing fruit dry matter, but declined in maturing fruits apparently coinciding with the loss of chlorophyll. Slight net fruit photosynthetic rates ranging from 0. 087 ± 0. 06 to 0. 003 ± 0. 05 nmol CO₂ (g dry weight)^?1 S^?1 were measured in midseason under optimal temperature (15 and 20°C) and light (1200 μmol photons m^?2 S^?1) conditions. Calculated fruit photosynthetic rates per unit dry weight increased with increasing temperatures and photon flux densities during fruit development. Dark respiration rates per unit dry weight doubled within a temperature interval of 10°C; the mean seasonal O₁₀ value was 2. 03 between 20 and 30°C. The highest photosynthetic rates were measured at 35°C throughout the growing season. Since dark respiration rates increased at high temperatures to a greater extent than CO₂ exchange rates in light, fruit photosynthesis was apparently stimulated by high internal CO₂ concentrations via CO₂ refixation. At 15°C, fruit photosynthetic rates tended to be saturated at about 600 μmol photons m^?2 S^?1. Young peach fruits responded to increasing ambient CO₂ concentrations with decreasing net CO₂ exchange rates in light, but more mature fruits did not respond to increases in ambient CO₂. Fruit CO₂ exchange rates in the dark remained fairly constant, apparently uninfluenced by ambient CO₂ concentrations during the entire growing season. Calculated fruit photosynthetic rates clearly revealed the difference in CO₂ response of young and mature peach fruits. Photosynthetic rates of younger peach fruits apparently approached saturation at 370 μl CO₂1^?2. In CO₂ free air, fruit photosynthesis was dependent on CO₂ refixation since CO₂ uptake by the fruits from the external atmosphere was not possible. The difference in photosynthetic rates between fruits in CO₂-free air and 370 μl CO₂ 1^?1 indicated that young peach fruits were apparently able to take up CO₂ from the external atmosphere. CO₂ uptake by peach fruits contributed between 28 and 16% to the fruit photosynthetic rate early in the season, whereas photosynthesis in maturing fruits was supplied entirely by CO₂ refixation.     

Responses of wheat and rice to factorial combinations of ambient and elevated CO2 and temperature in FACE experiments

Elevated CO₂ and temperature strongly affect crop production, but understanding of the crop response to combined CO₂ and temperature increases under field conditions is still limited while data are scarce. We grew wheat (Triticum aestivum L.) and rice (Oryza sativa L.) under two levels of CO₂ (ambient and enriched up to 500 μmol mol^?1) and two levels of canopy temperature (ambient and increased by 1.5–2.0 °C) in free‐air CO₂ enrichment (FACE) systems and carried out a detailed growth and yield component analysis during two growing seasons for both crops. An increase in CO₂ resulted in higher grain yield, whereas an increase in temperature reduced grain yield, in both crops. An increase in CO₂ was unable to compensate for the negative impact of an increase in temperature on biomass and yield of wheat and rice. Yields of wheat and rice were decreased by 10–12% and 17–35%, respectively, under the combination of elevated CO₂ and temperature. The number of filled grains per unit area was the most important yield component accounting for the effects of elevated CO₂ and temperature in wheat and rice. Our data showed complex treatment effects on the interplay between preheading duration, nitrogen uptake, tillering, leaf area index, and radiation‐use efficiency, and thus on yield components and yield. Nitrogen uptake before heading was crucial in minimizing yield loss due to climate change in both crops. For rice, however, a breeding strategy to increase grain number per m² and % filled grains (or to reduce spikelet sterility) at high temperature is also required to prevent yield reduction under conditions of global change.     

Free Air CO2 Enrichment of potato (Solanum tuberosum L.): development, growth and yield

A FACE (Free Air CO₂ Enrichment) experiment was carried out on Potato (Solanum tuberosum L., cv. Primura) in 1995 in Italy. Three FACE rings were used to fumigate circular field plots of 8 m diameter while two rings were used as controls at ambient CO₂ concentrations. Four CO₂ exposure levels were used in the rings (ambient, 460, 560 and 660 μmol mol^–1). Phenology and crop development, canopy surface temperature, above- and below-ground biomass were monitored during the growing season. Crop phenology was affected by elevated CO₂, as the date of flowering was progressively anticipated in the 660, 560, 460 μmol mol^–1 treatments. Crop development was not affected significantly as plant height, leaf area and the number of leaves per plant were the same in the four treatments. Elevated atmospheric CO₂ levels had, instead, a significant effect on the accumulation of total nonstructural carbohydrates (TNC = soluble sugars + starch) in the leaves during a sunny day. Specific leaf area was decreased under elevated CO₂ with a response that paralleled that of TNC concentrations. This reflected the occurrence of a progressive increase of photosynthetic rates and carbon assimilation in plants exposed to increasingly higher levels of atmospheric CO₂. Tuber growth and final tuber yield were also stimulated by rising CO₂ levels. When calculated by regression of tuber yield vs. the imposed levels of CO₂concentration, yield stimulation was as large as 10% every 100 μmol mol^–1 increase, which translated into over 40% enhancement in yield under 660 μmol mol^–1. This was related to a higher number of tubers rather than greater mean tuber mass or size. Leaf senescence was accelerated under elevated CO₂ and a linear relationship was found between atmospheric CO₂ levels and leaf reflectance measured at 0.55 μm wavelength. We conclude that significant CO₂ stimulation of yield has to be expected for potato under future climate scenarios, and that crop phenology will be affected as well.     

Water availability moderates N2 fixation benefit from elevated [CO2]: A 2‐year free‐air CO2 enrichment study on lentil (Lens culinaris MEDIK.) in a water limited agroecosystem

Increased biomass and yield of plants grown under elevated [CO₂] often corresponds to decreased grain N concentration ([N]), diminishing nutritional quality of crops. Legumes through their symbiotic N₂ fixation may be better able to maintain biomass [N] and grain [N] under elevated [CO₂], provided N₂ fixation is stimulated by elevated [CO₂] in line with growth and yield. In Mediterranean‐type agroecosystems, N₂ fixation may be impaired by drought, and it is unclear whether elevated [CO₂] stimulation of N₂ fixation can overcome this impact in dry years. To address this question, we grew lentil under two [CO₂] (ambient ~400 ppm and elevated ~550 ppm) levels in a free‐air CO₂ enrichment facility over two growing seasons sharply contrasting in rainfall. Elevated [CO₂] stimulated N₂ fixation through greater nodule number (+27%), mass (+18%), and specific fixation activity (+17%), and this stimulation was greater in the high than in the low rainfall/dry season. Elevated [CO₂] depressed grain [N] (?4%) in the dry season. In contrast, grain [N] increased (+3%) in the high rainfall season under elevated [CO₂], as a consequence of greater post‐flowering N₂ fixation. Our results suggest that the benefit for N₂ fixation from elevated [CO₂] is high as long as there is enough soil water to continue N₂ fixation during grain filling.     

Effects of atmospheric CO2 enrichment and root restriction on leaf gas exchange and growth of banana (Musa)

The effects of atmospheric CO₂ enrichment and root restriction on photosynthetic characteristics and growth of banana (Musa sp. AAA cv. Gros Michel) plants were investigated. Plants were grown aeroponically in root chambers in controlled environment glasshouse rooms at CO₂ concentrations of 350 or 1 000 μmol CO₂ mol^-1. At each CO₂ concentration, plants were grown in large (2001) root chambers that did not restrict root growth or in small (20 1) root chambers that restricted root growth. Plants grown at 350 μmol CO₂ mol^-1 generally had a higher carboxylation efficiency than plants grown at 1 000 μmol CO₂ mol^-1 although actual net CO₂ assimilation (A) was higher at the higher ambient CO₂ concentration due to increased intercellular CO₂ concentrations (C_i resulting from CO₂ enrichment. Thus, plants grown at 1 000 μmol CO₂ mol^-1 accumulated more leaf area and dry weight than plants grown at 350 μmol CO₂ mol^-1. Plants grown in the large root chambers were more photosynthetically efficient than plants grown in the small root chambers. At 350 μmol CO₂ mol^-1, leaf area and dry weights of plant organs were generally greater for plants in the large root chambers compared to those in the small root chambers. Atmospheric CO₂ enrichment may have compensated for the effects of root restriction on plant growth since at 1 000 μmol CO₂ mol^-1 there was generally no effect of root chamber size on plant dry weight.     

Allocation responses to CO2 enrichment and defoliation by a native annual plant Heterotheca subaxillaris

Among plants grown under enriched atmospheric CO₂, root:shoot balance (RSB) theory predicts a proportionately greater allocation of assimilate to roots than among ambient‐grown plants. Conversely, defoliation, which decreases the plant's capacity to assimilate carbon, is predicted to increase allocation to shoot. We tested these RSB predictions, and whether responses to CO₂ enrichment were modified by defoliation, using Heterotheca subaxillaris, an annual plant native to south‐eastern USA. Plants were grown under near‐ambient (400 μmol mol^?1) and enriched (700 μmol mol^?1) levels of atmospheric CO₂. Defoliation consisted of the weekly removal of 25% of each new fully expanded, but not previously defoliated, leaf from either rosette or bolted plants. In addition to dry mass measurements of leaves, stems, and roots, Kjeldahl N, protein, starch and soluble sugars were analysed in these plant components to test the hypothesis that changes in C:N uptake ratio drive shifts in root:shoot ratio. Young, rapidly growing CO₂‐enriched plants conformed to the predictions of RSB, with higher root:shoot ratio than ambient‐grown plants (P < 0.02), whereas older, slower growing plants did not show a CO₂ effect on root:shoot ratio. Defoliation resulted in smaller plants, among which both root and shoot biomass were reduced, irrespective of CO₂ treatment (P < 0.03). However, H. subaxillaris plants were able to compensate for leaf area removal through flexible shoot allocation to more leaves vs. stem (P < 0.01). Increased carbon availability through CO₂ enrichment did not enhance the response to defoliation, apparently because of complete growth compensation for defoliation, even under ambient conditions. CO₂‐enriched plants had higher rates of photosynthesis (P < 0.0001), but this did not translate into increased final biomass accumulation. On the other hand, earlier and more abundant yield of flower biomass was an important consequence of growth under CO₂ enrichment.     

Effects of elevated CO2 on plant growth and nutrient partitioning of rice (Oryza sativa L.) at rapid tillering and physiological maturity

Abstract

This work investigates the relationship between plant growth, grain yield, nutrient acquisition and partitioning in rice (Oryza sativa L.) under elevated CO₂. Plants were grown hydroponically in growth chambers with a 12-h photoperiod at either 370 or 700 µmol CO₂ mol^?1 concentration. Plant dry mass (DM), grain yield and macro- and micronutrient concentrations of vegetative organs and grains were determined. Elevated CO₂ increased biomass at tillering, and this was largely due to an increase in root mass by 160%. Elevated CO₂ had no effect on total nutrient uptake (N, P, K, Mg and Ca). However, nutrient partitioning among organs was significantly altered. N partitioning to leaf blades was significantly decreased, whereas the N partitioning into the leaf sheaths and roots was increased. Nutrient use efficiency of N, P, K, and Mg in all organs was significantly increased at elevated CO₂. At harvest maturity, grain yield was increased by 27% at elevated CO₂ while grain (protein) concentration was decreased by a similar magnitude (28%), suggesting that critical nutrient requirements for rice might need to be reassessed with global climate change.     

Productivity and water use of wheat under free-air CO2 enrichment

A free-air CO₂ enrichment (FACE) experiment was conducted at Maricopa, Arizona, on wheat from December 1992 through May 1993. The FACE apparatus maintained the CO₂ concentration, [CO₂], at 550 μmol mol^?1 across four replicate 25-m-diameter circular plots under natural conditions in an open field. Four matching Control plots at ambient [CO₂] (about 370 μmol mol^?1) were also installed in the field. In addition to the two levels of [CO₂], there were ample (Wet) and limiting (Dry) levels of water supplied through a subsurface drip irrigation system in a strip, split-plot design. Measurements were made of net radiation, R_n; soil heat flux, G_o; soil temperature; foliage or surface temperature; air dry and wet bulb temperatures; and wind speed. Sensible heat flux, H, was calculated from the wind and temperature measurements. Latent heat flux, λET, and evapotranspiration, ET, were determined as the residual in the energy balance. The FACE treatment reduced daily total R_n by an average 4%. Daily FACE sensible heat flux, H, was higher in the FACE plots. Daily latent heat flux, λET, and evapotranspiration, ET, were consistently lower in the FACE plots than in the Control plots for most of the growing season, about 8% on the average. Net canopy photosynthesis was stimulated by an average 19 and 44% in the Wet and Dry plots, respectively, by elevated [CO₂] for most of the growing season. No significant acclimation or down regulation was observed. There was little above-ground growth response to elevated [CO₂] early in the season when temperatures were cool. Then, as temperatures warmed into spring, the FACE plants grew about 20% more than the Control plants at ambient [CO₂], as shown by above-ground biomass accumulation. Root biomass accumulation was also stimulated about 20%. In May the FACE plants matured and senesced about a week earlier than the Controls in the Wet plots. The FACE plants averaged 0.6 °C warmer than the Controls from February through April in the well-watered plots, and we speculate that this temperature rise contributed to the earlier maturity. Because of the acceleration of senescence, there was a shortening of the duration of grain filling, and consequently, there was a narrowing of the final biomass and yield differences. The 20% mid-season growth advantage of FACE shrunk to about an 8% yield advantage in the Wet plots, while the yield differences between FACE and Control remained at about 20% in the Dry plots.     

Combined effects of elevated CO2 and air temperature on carbon assimilation of Pinus taeda trees

Branches of 22-year-old loblolly pine (Pinus taeda, L.) trees growing in a plantation were exposed to ambient CO₂, ambient + 165 μmol mol^?1 CO₂ or ambient + 330 μmol mol^?1 CO₂ concentrations in combination with ambient or ambient + 2°C air temperatures for 3 years. Field measurements in the third year indicated that net carbon assimilation was enhanced in the elevated CO₂ treatments in all seasons. On the basis of A/C_i, curves, there was no indication of photosynthetic down-regulation. Branch growth and leaf area also increased significantly in the elevated CO₂ treatments. The imposed 2°C increase in air temperature only had slight effects on net assimilation and growth. Compared with the ambient CO₂ treatment, rates of net assimilation were ～1·6 times greater in the ambient + 165 μmol mol^?1 CO₂ treatment and 2·2 times greater in the ambient + 330 μmol mol^?1 CO₂ treatment. These ratios did not change appreciably in measurements made in all four seasons even though mean ambient air temperatures during the measurement periods ranged from 12·6 to 28·2°C. This indicated that the effect of elevated CO₂ concentrations on net assimilation under field conditions was primarily additive. The results also indicated that the effect of elevated CO₂ (+ 165 or + 330 μmol mol^?1) was much greater than the effect of a 2°C increase in air temperature on net assimilation and growth in this species.     

Seasonal changes in the effects of elevated CO2 on rice at three levels of nitrogen supply: a free air CO2 enrichment (FACE) experiment

Over time, the stimulative effect of elevated CO₂ on the photosynthesis of rice crops is likely to be reduced with increasing duration of CO₂ exposure, but the resultant effects on crop productivity remain unclear. To investigate seasonal changes in the effect of elevated CO₂ on the growth of rice (Oryza sativa L.) crops, a free air CO₂ enrichment (FACE) experiment was conducted at Shizukuishi, Iwate, Japan in 1998–2000. The target CO₂ concentration of the FACE plots was 200 µmol mol^?1 above that of ambient. Three levels of nitrogen (N) were supplied: low (LN, 4 g N m^?2), medium [MN, 8 (1998) and 9 (1999, 2000) g N m^?2] and high N (HN, 12 and 15 g N m^?2). For MN and HN but not for LN, elevated CO₂ increased tiller number at panicle initiation (PI) but this positive response decreased with crop development. As a result, the response of green leaf area index (GLAI) to elevated CO₂ greatly varied with development, showing positive responses during vegetative stages and negative responses after PI. Elevated CO₂ decreased leaf N concentration over the season, except during early stage of development. For MN crops, total biomass increased with elevated CO₂, but the response declined linearly with development, with average increases of 32, 28, 21, 15 and 12% at tillering, PI, anthesis, mid‐ripening and grain maturity, respectively. This decline is likely to be due to decreases in the positive effects of elevated CO₂ on canopy photosynthesis because of reductions in both GLAI and leaf N. Up to PI, LN‐crops tended to have a lower response to elevated CO₂ than MN‐ and HN‐crops, though by final harvest the total biomass response was similar for all N levels. For MN‐ and HN‐crops, the positive response of grain yield (ca. 15%) to elevated CO₂ was slightly greater than the response of final total biomass while for LN‐crops it was less. We conclude that most of the seasonal changes in crop response to elevated CO₂ are directly or indirectly associated with N uptake.     

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.