Leaf photosynthesis and carbohydrates of CO2-enriched maize and grain sorghum exposed to a short period of soil water deficit during vegetative development.

Among C4 species, sorghum is known to be more drought tolerant than maize. The objective was to evaluate differences in leaf gas exchanges, carbohydrates, and two enzyme activities of these nicotinamide adenine dinucleotide phosphate-malic enzyme (NADP-ME) C4 subtype monocots in response to water deficit and CO2 concentration ([CO2]). Maize and sorghum were grown in pots in sunlit environmental-controlled chambers. Treatments included well watered (WW) and water stressed (WS) (water withheld at 26 days) and daytime [CO2] of 360 (ambient) and 720 (elevated) µmol mol⁻¹. Midday gas exchange rates, concentrations of nonstructural carbohydrates, and activities of sucrose-phosphate synthase (SPS) and adenosine 5'-diphosphoglucose pyrophosphorylase (ADGP) were determined for fully expanded leaf sections. There was no difference in leaf CO2 exchange rates (CER) between ambient and elevated [CO2] control plants for both maize and sorghum. After withholding water, leaf CER declined to zero after 8 days in maize and 10 days for sorghum. Sorghum had lower stomatal conductance and transpiration rates than maize, which resulted in a longer period of CER under drought. Nonstructural carbohydrates of both control maize and sorghum were hardly affected by elevated [CO2]. Under drought, however, increases in soluble sugars and decreases in starch were generally observed for maize and sorghum at both [CO2] levels. For stressed maize and sorghum, decreases in starch occurred earlier and were greater at ambient [CO2] than at elevated [CO2]. For maize, drought did not meaningfully affect SPS activity. However, a decline in SPS activity was observed for drought-stressed sorghum under both [CO2] treatments. There was an increase in ADGP activity in maize under drought for both [CO2] treatments. Such a response in ADGP to drought, however, did not occur for sorghum. The generally more rapid response of maize than sorghum to drought might be related to the more rapid growth of leaf area of maize.

Elevated CO2 increases water use efficiency by sustaining photosynthesis of water-limited maize and sorghum.

Maize and grain sorghum seeds were sown in pots and grown for 39 days in sunlit controlled-environment chambers at 360 (ambient) and 720 (double-ambient, elevated)µmol mol(-1) carbon dioxide concentrations [CO(2)]. Canopy net photosynthesis (PS) and evapotranspiration (TR) was measured throughout and summarized daily from 08:00 to 17:00h Eastern Standard Time. Irrigation was withheld from matched pairs of treatments starting on 26 days after sowing (DAS). By 35 DAS, cumulative PS of drought-stress maize, compared to well-watered plants, was 41% lower under ambient [CO(2)] but only 13% lower under elevated [CO(2)]. In contrast, by 35 DAS, cumulative PS of drought-stress grain sorghum, compared to well-watered plants, was only 9% lower under ambient [CO(2)] and 7% lower under elevated [CO(2)]. During the 27-35 DAS drought period, water use efficiency (WUE, mol CO(2)Kmol(-1)H(2)O), was 3.99, 3.88, 5.50, and 8.65 for maize and 3.75, 4.43, 5.26, and 9.94 for grain sorghum, for ambient-[CO(2)] well-watered, ambient-[CO(2)] stressed, elevated-[CO(2)] well-watered and elevated-[CO(2)] stressed plants, respectively. Young plants of maize and sorghum used water more efficiently at elevated [CO(2)] than at ambient [CO(2)], especially under drought. Reductions in biomass by drought for young maize and grain sorghum plants were 42 and 36% at ambient [CO(2)], compared to 18 and 14% at elevated [CO(2)], respectively. Results of our water stress experiment demonstrated that maintenance of relatively high canopy photosynthetic rates in the face of decreased transpiration rates enhanced WUE in plants grown at elevated [CO(2)]. This confirms experimental evidence and conceptual models that suggest that an increase of intercellular [CO(2)] (or a sustained intercellular [CO(2)]) in the face of decreased stomatal conductance results in relative increases of growth of C(4) plants. In short, drought stress in C(4) crop plants can be ameliorated at elevated [CO(2)] as a result of lower stomatal conductance and sustaining intercellular [CO(2)]. Furthermore, less water might be required for C(4) crops in future higher CO(2) atmospheres, assuming weather and climate similar to present conditions.

Stem juice production of the C4 sugarcane (Saccharum officinarum) is enhanced by growth at double-ambient CO2 and high temperature.

Two cultivars of sugarcane (Saccharum officinarum cv. CP73-1547 and CP88-1508) were grown for 3 months in paired-companion, temperature-gradient, sunlit greenhouses under daytime [CO2] of 360 (ambient) and 720 (double ambient) micromol mol(-1) and at temperatures of 1.5 degrees C (near ambient) and 6.0 degrees C higher than outside ambient temperature. Leaf area and biomass, stem biomass and juice and CO2 exchange rate (CER) and activities of ribulose bisphosphate carboxylase-oxygenase (Rubisco) and phosphoenolpyruvate carboxylase (PEPC) of fully developed leaves were measured at harvest. On a main stem basis, leaf area, leaf dry weight, stem dry weight and stem juice volume were increased by growth at doubled [CO2] or high temperature. Such increases were even greater under combination of doubled [CO2]/high temperature. Plants grown at doubled [CO2]/high temperature combination averaged 50%, 26%, 84% and 124% greater in leaf area, leaf dry weight, stem dry weight and stem juice volume, respectively, compared with plants grown at ambient [CO2]/near-ambient temperature combination. In addition, plants grown at doubled [CO2]/high temperature combination were 2-3-fold higher in stem soluble solids than those at ambient [CO2]/near-ambient temperature combination. Although midday CER of fully developed leaves was not affected by doubled [CO2] or high temperature, plants grown at doubled [CO2] were 41-43% less in leaf stomatal conductance and 69-79% greater in leaf water-use efficiency, compared with plants grown at ambient [CO2]. Activity of PEPC was down-regulated 23-32% at doubled [CO2], while high temperature did not have a significant impact on this enzyme. Activity of Rubisco was not affected by growth at doubled [CO2], but was reduced 15-28% at high temperature. The increases in stem juice production and stem juice soluble solids concentration for sugarcane grown at doubled [CO2] or high temperature, or at doubled [CO2]/high temperature combination, were partially the outcome of an increase in whole plant leaf area. Such increase would enhance the ongoing and cumulative photosynthetic capability of the whole plant. The results indicate that a doubling of [CO2] would benefit sugarcane production more than the anticipated 10-15% increase for a C4 species.

Growth at elevated CO(2) delays the adverse effects of drought stress on leaf photosynthesis of the C(4) sugarcane.

Sugarcane (Saccharum officinarum L. cv. CP72-2086) was grown in sunlit greenhouses at daytime [CO(2)] of 360 (ambient) and 720 (elevated)mumolmol(-1). Drought stress was imposed for 13d when plants were 4 months old, and various photosynthetic parameters and levels of nonstructural carbohydrates were determined for uppermost fully expanded leaves of well-watered (control) and drought stress plants. Control plants at elevated [CO(2)] were 34% and 25% lower in leaf stomatal conductance (g(s)) and transpiration rate (E) and 35% greater in leaf water-use efficiency (WUE) than their counterparts at ambient [CO(2)]. Leaf CO(2) exchange rate (CER) and activities of Rubisco, NADP-malate dehydrogenase, NADP-malic enzyme and pyruvate P(i) dikinase were marginally affected by elevated [CO(2)], but were reduced by drought, whereas activity of PEP carboxylase was reduced by elevated [CO(2)], but not by drought. At severe drought developed at day 12, leaf g(s) and WUE of ambient-[CO(2)] stress plants declined to 5% and 7%, while elevated-[CO(2)] stress plants still maintained g(s) and WUE at 20% and 74% of their controls. In control plants, elevated [CO(2)] did not enhance the midday levels of starch, sucrose, or reducing sugars. For both ambient- and elevated-[CO(2)] stress plants, severe drought did not affect the midday level of sucrose but substantially reduced that of starch. Nighttime starch decomposition in control plants was 55% for ambient [CO(2)] and 59% for elevated [CO(2)], but was negligible for stress plants of both [CO(2)] treatments. For both ambient-[CO(2)] control and stress plants, midday sucrose level at day 12 was similar to the predawn value at day 13. In contrast, sucrose levels of elevated-[CO(2)] control and stress plants at predawn of day 13 were 61-65% of the midday values of day 12. Levels of reducing sugars were much greater for both ambient- and elevated-[CO(2)] stress plants, implying an adaptation to drought stress. Sugarcane grown at elevated [CO(2)] had lower leaf g(s) and E and greater leaf WUE, which helped to delay the adverse effects of drought and, thus, allowed the stress plants to continue photosynthesis for at least an extra day during episodic drought cycles.

Enhancement in leaf photosynthesis and upregulation of Rubisco in the C4 sorghum plant at elevated growth carbon dioxide and temperature occur at early stages of leaf ontogeny.

Rising atmospheric carbon dioxide (CO2) concentration and temperature will influence photosynthesis, growth and yield of agronomic crops. To investigate effects of elevated CO2 and high temperature on leaf gas exchanges, activities of Rubisco and phosphoenolpyruvate carboxylase (PEPC) and growth of grain sorghum (Sorghum bicolor L. Moench), plants were grown in controlled environments at day-time maximum/night-time minimum temperatures of 30/20°C or 36/26°C at ambient (350 µmol mol-1) or elevated (700 µmol mol-1) CO2. Gas-exchange rates, activities of Rubisco and PEPC and growth parameters (leaf, stem and total dry weights) were determined at different stages of leaf development. Between 6 and 25 days after leaf tip emergence, leaf carbon exchange rate (CER) of elevated CO2 plants was greater at 30/20°C and 36/26°C than that of ambient CO2 plants at the same temperatures. The positive response of CER to elevated CO2 was greater in young leaves than in old leaves. In young leaves, elevated CO2 enhanced Rubisco activity at 30/20°C and 36/26°C, whereas PEPC activity was not affected by elevated CO2 at 30/20°C but was marginally enhanced at 36/26°C. At 30/20°C, growth parameters were not affected by elevated CO2 until 50 days after sowing (DAS); at 36/26°C, they were progressively enhanced by elevated CO2 to as high as 49 to 62% by 50 DAS. Leaf CER and Rubisco activity were enhanced by elevated CO2 at early stages of leaf ontogeny for the C4 grain sorghum. Such enhancement should have a significant role in dry matter production under elevated CO2.