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.     

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 [CO₂]. Canopy net photosynthesis (PS) and evapotranspiration (TR) was measured throughout and summarized daily from 08:00 to 17:00 h 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 [CO₂] but only 13% lower under elevated [CO₂]. In contrast, by 35 DAS, cumulative PS of drought-stress grain sorghum, compared to well-watered plants, was only 9% lower under ambient [CO₂] and 7% lower under elevated [CO₂]. During the 27-35 DAS drought period, water use efficiency (WUE, mol CO₂ Kmol⁻¹ H₂O), 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-[CO₂] well-watered, ambient-[CO₂] stressed, elevated-[CO₂] well-watered and elevated-[CO₂] stressed plants, respectively. Young plants of maize and sorghum used water more efficiently at elevated [CO₂] than at ambient [CO₂], especially under drought. Reductions in biomass by drought for young maize and grain sorghum plants were 42 and 36% at ambient [CO₂], compared to 18 and 14% at elevated [CO₂], 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 [CO₂]. This confirms experimental evidence and conceptual models that suggest that an increase of intercellular [CO₂] (or a sustained intercellular [CO₂]) in the face of decreased stomatal conductance results in relative increases of growth of C₄ plants. In short, drought stress in C₄ crop plants can be ameliorated at elevated [CO₂] as a result of lower stomatal conductance and sustaining intercellular [CO₂]. Furthermore, less water might be required for C₄ crops in future higher CO₂ atmospheres, assuming weather and climate similar to present conditions.     

Comparative field water relations of three Mediterranean shrub species co-occurring at a natural CO(2) vent

Annual variations in the water relations and stomatal response of Erica arborea, Myrtus communis and Juniperus communis occurring at a natural CO(2) vent were analysed under Mediterranean field conditions. A distinct gradient of CO(2)concentration ([CO(2)]) exists between two sites near a natural CO(2)-emitting vent, with higher [CO(2)] (700 micromol mol(-1)) in the proximity of the CO(2) spring. Plants at the CO(2) spring site have been growing for generations at elevated [CO(2)]. At both sites, maximum leaf conductance was related to predawn shoot water potential. The effects of water deficits during the summer drought were severe. Leaf conductance and water potential recovered after major rainfalls in September to predrought values. Strong relationships between leaf conductance, predawn water potential, and leaf-specific hydraulic resistance are consistent with the role of stomata in regulating plant water status. Considerable between-species variation in sensitivity of water potentials and stomatal characters to elevated [CO(2)] were observed. Common to all the shrubs were a reduction in leaf conductance and an increase in water potentials in response to elevated [CO(2)]. Elevated [CO(2)] decreased the sensitivity of leaf conductance to vapour pressure deficit. Morphological characters (including stomatal density and degree of sclerophylly) showed site-dependent variations, but degree and sign of such changes varied with the species and/or the season. Measurements of discrimination against (13)C provided evidence for long-term decreases of water use efficiency in CO(2) spring plants. Analysis of C isotope composition suggested that a downward adjustment of photosynthetic capacity may have occurred under elevated [CO(2)]. Elevated [CO(2)] effects on water relations and leaf morphology persisted in the long term, but the three shrubs growing in the same environment showed species-specific responses.     

Water relations and gas exchange in poplar and willow under water stress and elevated atmospheric CO2

Predictions of shifts in rainfall patterns as atmospheric [CO2] increases could impact the growth of fast growing trees such as Populus spp. and Salix spp. and the interaction between elevated CO2 and water stress in these species is unknown. The objectives of this study were to characterize the responses to elevated CO2 and water stress in these two species, and to determine if elevated CO2 mitigated drought stress effects. Gas exchange, water potential components, whole plant transpiration and growth response to soil drying and recovery were assessed in hybrid poplar (clone 53-246) and willow (Salix sagitta) rooted cuttings growing in either ambient (350 &mgr;mol mol-1) or elevated (700 &mgr;mol mol-1) atmospheric CO2 concentration ([CO2]). Predawn water potential decreased with increasing water stress while midday water potentials remained unchanged (isohydric response). Turgor potentials at both predawn and midday increased in elevated [CO2], indicative of osmotic adjustment. Gas exchange was reduced by water stress while elevated [CO2] increased photosynthetic rates, reduced leaf conductance and nearly doubled instantaneous transpiration efficiency in both species. Dark respiration decreased in elevated [CO2] and water stress reduced Rd in the trees growing in ambient [CO2]. Willow had 56% lower whole plant hydraulic conductivity than poplar, and showed a 14% increase in elevated [CO2] while poplar was unresponsive. The physiological responses exhibited by poplar and willow to elevated [CO2] and water stress, singly, suggest that these species respond like other tree species. The interaction of [CO2] and water stress suggests that elevated [CO2] did mitigate the effects of water stress in willow, but not in poplar.     

Impairment of C(4) photosynthesis by drought is exacerbated by limiting nitrogen and ameliorated by elevated [CO(2)] in maize

Predictions of future ecosystem function and food supply from staple C(4) crops, such as maize, depend on elucidation of the mechanisms by which environmental change and growing conditions interact to determine future plant performance. To test the interactive effects of elevated [CO(2)], drought, and nitrogen (N) supply on net photosynthetic CO(2) uptake (A) in the world's most important C(4) crop, maize (Zea mays) was grown at ambient [CO(2)] (～385 ppm) and elevated [CO(2)] (550 ppm) with either high N supply (168 kg N ha(-1) fertilizer) or limiting N (no fertilizer) at a site in the US Corn Belt. A mid-season drought was not sufficiently severe to reduce yields, but caused significant physiological stress, with reductions in stomatal conductance (up to 57%), A (up to 44%), and the in vivo capacity of phosphoenolpyruvate carboxylase (up to 58%). There was no stimulation of A by elevated [CO(2)] when water availability was high, irrespective of N availability. Elevated [CO(2)] delayed and relieved both stomatal and non-stomatal limitations to A during the drought. Limiting N supply exacerbated stomatal and non-stomatal limitation to A during drought. However, the effects of limiting N and elevated [CO(2)] were additive, so amelioration of stress by elevated [CO(2)] did not differ in magnitude between high N and limiting N supply. These findings provide new understanding of the limitations to C(4) photosynthesis that will occur under future field conditions of the primary region of maize production in the world.     

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.     

Sugar accumulation in roots of two grape varieties with contrasting response to water stress

Root sugar accumulation was studied in two grapevine varieties contrasting in tolerance to water stress. During a 10‐day water withholding treatment, the drought‐tolerant variety, Grenache, sustained less negative predawn and midday leaf water potentials as well as root water potential compared with the sensitive variety, Semillon. Grenache vines also maintained lower stomatal conductance and transpiration than Semillon vines throughout the drying period. In both varieties there was accumulation of sucrose in the roots and concentrations were inversely correlated to leaf and root water status. In both Grenache and Semillon, elevated root osmolality was associated with decreased soil moisture indicating that sugar accumulation may play a role in osmotic protection. Petiole xylem sap abscisic acid (ABA) concentrations increased with water deficit in both varieties and were highest for vines with the most negative root and predawn leaf water potentials. Furthermore, root sucrose concentrations were positively correlated with leaf xylem sap ABA concentrations, indicative of integration between carbohydrate metabolism and the ABA signalling system. Similar root sugar accumulation patterns between the two varieties, however, demonstrate that other factors are likely influencing the ability of the drought‐tolerant variety to remain hydrated.     

Maintenance of C sinks sustains enhanced C assimilation during long-term exposure to elevated [CO2] in Mojave Desert shrubs

During the first few years of elevated atmospheric [CO(2)] treatment at the Nevada Desert FACE Facility, photosynthetic downregulation was observed in desert shrubs grown under elevated [CO(2)], especially under relatively wet environmental conditions. Nonetheless, those plants maintained increased A (sat) (photosynthetic performance at saturating light and treatment [CO(2)]) under wet conditions, but to a much lesser extent under dry conditions. To determine if plants continued to downregulate during long-term exposure to elevated [CO(2)], responses of photosynthesis to elevated [CO(2)] were examined in two dominant Mojave Desert shrubs, the evergreen Larrea tridentata and the drought-deciduous Ambrosia dumosa, during the eighth full growing season of elevated [CO(2)] treatment at the NDFF. A comprehensive suite of physiological processes were collected. Furthermore, we used C labeling of air to assess carbon allocation and partitioning as measures of C sink activity. Results show that elevated [CO(2)] enhanced photosynthetic performance and plant water status in Larrea, especially during periods of environmental stress, but not in Ambrosia. δ(13)C analyses indicate that Larrea under elevated [CO(2)] allocated a greater proportion of newly assimilated C to C sinks than Ambrosia. Maintenance by Larrea of C sinks during the dry season partially explained the reduced [CO(2)] effect on leaf carbohydrate content during summer, which in turn lessened carbohydrate build-up and feedback inhibition of photosynthesis. δ(13)C results also showed that in a year when plant growth reached the highest rates in 5 years, 4% (Larrea) and 7% (Ambrosia) of C in newly emerging organs were remobilized from C that was assimilated and stored for at least 2 years prior to the current study. Thus, after 8 years of continuous exposure to elevated [CO(2)], both desert perennials maintained their photosynthetic capacities under elevated [CO(2)]. We conclude that C storage, remobilization, and partitioning influence the responsiveness of these desert shrubs during long-term exposure to elevated [CO(2)].     

Effect of drought and high solar radiation on 1-aminocyclopropane-1-carboxylic acid and abscisic acid concentrations in Rosmarinus officinalis plants

The endogenous concentrations of ACC and ABA were measured, at predawn and at maximum solar radiation, during a summer drought, and recovery after autumn rainfalls, in rosemary (Rosmarinus officinalis L.), a drought-tolerant species, growing under Mediterranean field conditions. During the summer, plants were subjected to both water deficit and high solar radiation. Plants showed severe reductions in shoot water potential to -3 MPa, which were associated with drastic stomatal closure (73%), a decrease in net photosynthesis, reaching almost zero, and a severe chlorophyll loss (74%). Despite the severity of the stress, plants recovered after the autumn rainfalls. The concentration of ACC was not enhanced by drought, and at predawn these concentrations remained constant at approximately 600 pmol ACC-1 DW throughout the experiment. Thus, ethylene did not regulate the response of rosemary to drought. However, a sharp increase in ACC levels between predawn and midday was observed. This increase was positively correlated to the intensity of the incident solar radiation. ACC levels recorded in June at midday reached 16 000 pmol g DW and in October values of 1000 pmol g-1 DW were observed. In contrast, in drought-stressed plants predawn concentrations of ABA were up to 130-fold those of recovered plants, and the levels of ABA scored at midday were double of those scored at predawn. In conclusion, although drought-stressed rosemary plants showed a relatively moderate ABA accumulation (approximately 500 pmol g-1 DW#, at predawn), it seems to be an essential factor for the regulation of the plant response to stress, thereby enabling a rapid recovery after stress release, although other mechanisms can not be excluded. As drought stress did not induce ACC accumulation, it was concluded that ethylene production was not a major factor in the drought stress resistance of rosemary plants. The increased ACC and ABA concentrations at midday were correlated with day length and light intensity and not with the water status of the plant.     

Heterophylly in the yellow waterlily, Nuphar variegata (Nymphaeaceae): effects of [CO2], natural sediment type, and water depth

We transplanted Nuphar variegata with submersed leaves only into natural lake sediments in pH-, [CO(2)]-, depth-, and temperature-controlled greenhouse tanks to test the hypotheses that more fertile sediment, lower free [CO(2)], and shallower depth would all stimulate the development of floating leaves. Sediment higher in porewater [NH(4)(+)] favored floating leaf development. Low CO(2)-grown plants initiated floating leaf development significantly earlier than high CO(2)-grown plants, which produced significantly more submersed leaves and fewer floating leaves. Mean floating leaf biomass was significantly greater than mean submersed leaf biomass but was not influenced by CO(2) enrichment, whereas mean submersed leaf biomass increased 88% at high [CO(2)]. At the shallower depth (35 cm), floating leaves required 50% less biomass investment per leaf than at 70 cm, and a significantly greater proportion of plants had floating leaves (70 vs. 23-43% at 35 vs. 70 cm, respectively) for the last three of the eight leaf censuses. Sediment type, water depth, and especially free [CO(2)] all can influence leaf morphogenesis in Nuphar variegata, and the development of more and larger submersed leaves with CO(2) enrichment favors the exploitation of high [CO(2)] when it is present in the water column.     

Hourly and seasonal variation in photosynthesis and stomatal conductance of soybean grown at future CO(2) and ozone concentrations for 3 years under fully open-air field conditions

It is anticipated that enrichment of the atmosphere with CO(2) will increase photosynthetic carbon assimilation in C3 plants. Analysis of controlled environment studies conducted to date indicates that plant growth at concentrations of carbon dioxide ([CO(2)]) anticipated for 2050 ( approximately 550 micromol mol(-1)) will stimulate leaf photosynthetic carbon assimilation (A) by 20 to 40%. Simultaneously, concentrations of tropospheric ozone ([O(3)]) are expected to increase by 2050, and growth in controlled environments at elevated [O(3)] significantly reduces A. However, the simultaneous effects of both increases on a major crop under open-air conditions have never been tested. Over three consecutive growing seasons > 4700 individual measurements of A, photosynthetic electron transport (J(PSII)) and stomatal conductance (g(s)) were measured on Glycine max (L.) Merr. (soybean). Experimental treatments used free-air gas concentration enrichment (FACE) technology in a fully replicated, factorial complete block design. The mean A in the control plots was 14.5 micromol m(-2) s(-1). At elevated [CO(2)], mean A was 24% higher and the treatment effect was statistically significant on 80% of days. There was a strong positive correlation between daytime maximum temperatures and mean daily integrated A at elevated [CO(2)], which accounted for much of the variation in CO(2) effect among days. The effect of elevated [CO(2)] on photosynthesis also tended to be greater under water stress conditions. The elevated [O(3)] treatment had no statistically significant effect on mean A, g(s) or J(PSII) on newly expanded leaves. Combined elevation of [CO(2)] and [O(3)] resulted in a slightly smaller increase in average A than when [CO(2)] alone was elevated, and was significantly greater than the control on 67% of days. Thus, the change in atmospheric composition predicted for the middle of this century will, based on the results of a 3 year open-air field experiment, have smaller effects on photosynthesis, g(s) and whole chain electron transport through photosystem II than predicted by the substantial literature on relevant controlled environment studies on soybean and likely most other C3 plants.     

Response of respiration of soybean leaves grown at ambient and elevated carbon dioxide concentrations to day-to-day variation in light and temperature under field conditions

BACKGROUND AND AIMS: Respiration is an important component of plant carbon balance, but it remains uncertain how respiration will respond to increases in atmospheric carbon dioxide concentration, and there are few measurements of respiration for crop plants grown at elevated [CO(2)] under field conditions. The hypothesis that respiration of leaves of soybeans grown at elevated [CO(2)] is increased is tested; and the effects of photosynthesis and acclimation to temperature examined. METHODS: Net rates of carbon dioxide exchange were recorded every 10 min, 24 h per day for mature upper canopy leaves of soybeans grown in field plots at the current ambient [CO(2)] and at ambient plus 350 micromol mol(-1) [CO(2)] in open top chambers. Measurements were made on pairs of leaves from both [CO(2)] treatments on a total of 16 d during the middle of the growing seasons of two years. KEY RESULTS: Elevated [CO(2)] increased daytime net carbon dioxide fixation rates per unit of leaf area by an average of 48 %, but had no effect on night-time respiration expressed per unit of area, which averaged 53 mmol m(-2) d(-1) (1.4 micromol m(-2) s(-1)) for both the ambient and elevated [CO(2)] treatments. Leaf dry mass per unit of area was increased on average by 23 % by elevated [CO(2)], and respiration per unit of mass was significantly lower at elevated [CO(2)]. Respiration increased by a factor of 2.5 between 18 and 26 degrees C average night temperature, for both [CO(2)] treatments. CONCLUSIONS: These results do not support predictions that elevated [CO(2)] would increase respiration per unit of area by increasing photosynthesis or by increasing leaf mass per unit of area, nor the idea that acclimation of respiration to temperature would be rapid enough to make dark respiration insensitive to variation in temperature between nights.     

Drought stress, plant water status, and floral trait expression in fireweed, Epilobium angustifolium (Onagraceae)

In a controlled environment, we artificially induced drought during flowering of Epilobium angustifolium, an animal-pollinated plant. Leaf water potential (ψ(l)) and floral traits were monitored over a 12-d period of soil moisture depletion. Soil moisture depletion induced drought stress over time, as revealed by significant treatment × day interactions for predawn and midday ψ(l). Nectar volume and flower size showed significant negative responses to drought stress, but nectar sugar concentration did not vary between treatments. Floral traits were more buffered from drought than leaf water potentials. We used path analysis to examine direct and indirect effects of ψ(l) on floral traits for plants in well-watered (control) vs. drought treatments. According to the best-fit path models, midday ψ(l) has significant positive effects on flower size and nectar volume in both environments. However, for controls midday ψ(l) also had a significant negative effect on nectar sugar concentration. Results indicate that traits influencing floral attractiveness to pollinators in E. angustifolium vary with plant water status, such that pollinator-mediated selection could indirectly target physiological or biochemical controls on ψ(l). Moreover, under mesic conditions selection for greater nectar sugar reward may be constrained by the antagonistic effects of plant water status on nectar volume and sugar concentration.     

Elevated CO(2) induces biochemical and ultrastructural changes in leaves of the C(4) cereal sorghum

We analyzed the impact of growth at either 350 (ambient) or 700 (elevated) microL L(-1) CO(2) on key elements of the C(4) pathway (photosynthesis, carbon isotope discrimination, and leaf anatomy) using the C(4) cereal sorghum (Sorghum bicolor L. Moench.). Gas-exchange analysis of the CO(2) response of photosynthesis indicated that both carboxylation efficiency and the CO(2) saturated rate of photosynthesis were lower in plants grown at elevated relative to ambient CO(2). This was accompanied by a 49% reduction in the phosphoenolpyruvate carboxylase content of leaves (area basis) in the elevated CO(2)-grown plants, but no change in Rubisco content. Despite the lower phosphoenolpyruvate carboxylase content, there was a 3-fold increase in C isotope discrimination in leaves of plants grown at elevated CO(2) and bundle sheath leakiness was estimated to be 24% and 33%, respectively, for the ambient and elevated CO(2)-grown plants. However, we could detect no difference in quantum yield. The ratio of quantum yield of CO(2) fixation to PSII efficiency was lower in plants grown at elevated CO(2), but only when leaf internal was below 50 microL L(-1). This suggests a reduction in the efficiency of the C(4) cycle when [CO(2)] is low, and also implies increased electron transport to acceptors other than CO(2). Analysis of leaf sections using a transmission electron microscope indicated a 2-fold decrease in the thickness of the bundle sheath cell walls in plants grown at elevated relative to ambient CO(2). These results suggest that significant acclimation to increased CO(2) concentrations occurs in sorghum.     

Growth under elevated atmospheric CO(2) concentration accelerates leaf senescence in sunflower (Helianthus annuus L.) plants

Some morphogenetic and metabolic processes were sensitive to a high atmospheric CO(2) concentration during sunflower primary leaf ontogeny. Young leaves of sunflower plants growing under elevated CO(2) concentration exhibited increased growth, as reflected by the high specific leaf mass referred to as dry weight in young leaves (16days). The content of photosynthetic pigments decreased with leaf development, especially in plants grown under elevated CO(2) concentrations, suggesting that high CO(2) accelerates chlorophyll degradation, and also possibly leaf senescence. Elevated CO(2) concentration increased the oxidative stress in sunflower plants by increasing H(2)O(2) levels and decreasing activity of antioxidant enzymes such as catalase and ascorbate peroxidase. The loss of plant defenses probably increases the concentration of reactive oxygen species in the chloroplast, decreasing the photosynthetic pigment content as a result. Elevated CO(2) concentration was found to boost photosynthetic CO(2) fixation, especially in young leaves. High CO(2) also increased the starch and soluble sugar contents (glucose and fructose) and the C/N ratio during sunflower primary leaf development. At the beginning of senescence, we observed a strong increase in the hexoses to sucrose ratio that was especially marked at high CO(2) concentration. These results indicate that elevated CO(2) concentration could promote leaf senescence in sunflower plants by affecting the soluble sugar levels, the C/N ratio and the oxidative status during leaf ontogeny. It is likely that systemic signals produced in plants grown with elevated CO(2), lead to early senescence and a higher oxidation state of the cells of these plant leaves.     

Protective mechanism of desiccation tolerance in Reaumuria soongorica: Leaf abscission and sucrose accumulation in the stem

Reaumuria soongorica (Pall.) Maxim., a perennial semi-shrub, is widely found in semi-arid areas in northwestern China and can survive severe desiccation of its vegetative organs. In order to study the protective mechanism of desiccation tolerance in R. soongorica, diurnal patterns of net photosynthetic rate (Pn), water use efficiency (WUE) and chlorophyll fluorescence parameters of Photosystem II (PSII), and sugar content in the source leaf and stem were investigated in 6-year-old plants during progressive soil drought imposed by the cessation of watering. The results showed that R. soongorica was characterized by very low leaf water potential, high WUE, photosynthesis and high accumulation of sucrose in the stem and leaf abscission under desiccation. The maximum Pn increased at first and then declined during drought, but intrinsic WUE increased remarkably in the morning with increasing drought stress. The maximal photochemical efficiency of PSII (Fv/Fm) and the quantum efficiency of noncyclic electric transport of PSII(Φ_PSII) decreased significantly under water stress and exhibited an obvious phenomenon of photoinhibition at noon. Drought stressed plants maintained a higher capacity of dissipation of the excitation energy (measured as NPQ) with the increasing intensity of stress. Conditions of progressive drought promoted sucrose and starch accumulation in the stems but not in the leaves. However, when leaf water potential was less than −21.3 MPa, the plant leaves died and then abscised. But the stem photosynthesis remained and, afterward the plants entered the dormant state. Upon rewatering, the shoots reactivated and the plants developed new leaves. Therefore, R. soongorica has the ability to reduce water loss through leaf abscission and maintain the vigor of the stem cells to survive desiccation. Supported by the Program of the Research of Vegetation Restoration in Arid Areas of Lanzhou (Grant No. 03-2-27) and the National Natural Science Foundation of China (Grant No. 30270243)     

Long-term growth of soybean at elevated [CO2] does not cause acclimation of stomatal conductance under fully open-air conditions

Accurately predicting plant function and global biogeochemical cycles later in this century will be complicated if stomatal conductance (g(s)) acclimates to growth at elevated [CO(2)], in the sense of a long-term alteration of the response of g(s) to [CO(2)], humidity (h) and/or photosynthetic rate (A). If so, photosynthetic and stomatal models will require parameterization at each growth [CO(2)] of interest. Photosynthetic acclimation to long-term growth at elevated [CO(2)] occurs frequently. Acclimation of g(s) has rarely been examined, even though stomatal density commonly changes with growth [CO(2)]. Soybean was grown under field conditions at ambient [CO(2)] (378 micromol mol(-1)) and elevated [CO(2)] (552 micromol mol(-1)) using free-air [CO(2)] enrichment (FACE). This study tested for stomatal acclimation by parameterizing and validating the widely used Ball et al. model (1987, Progress in Photosynthesis Research, vol IV, 221-224) with measurements of leaf gas exchange. The dependence of g(s) on A, h and [CO(2)] at the leaf surface was unaltered by long-term growth at elevated [CO(2)]. This suggests that the commonly observed decrease in g(s) under elevated [CO(2)] is due entirely to the direct instantaneous effect of [CO(2)] on g(s) and that there is no longer-term acclimation of g(s) independent of photosynthetic acclimation. The model accurately predicted g(s) for soybean growing under ambient and elevated [CO(2)] in the field. Model parameters under ambient and elevated [CO(2)] were indistinguishable, demonstrating that stomatal function under ambient and elevated [CO(2)] could be modelled without the need for parameterization at each growth [CO(2)].     

Effects of water stress and rewatering on leaf water relations of lemon plants

Potted two-year-old lemon plants (Citrus limon (L.) Burm. fil.) cv. Fino, growing under field conditions were subjected to drought by withholding irrigation for 13 d. After that, plants were re-irrigated and the recovery was studied for 5 d. Control plants were daily irrigated maintaining the soil matric potential at about -30 kPa. Young leaves of control plants presented higher leaf conductance (g1) and lower midday leaf water potential (Ψmd) than mature ones. Young leaves also showed higher leaf water potential at the turgor loss point (Ψtlp) than mature leaves. In both leaf types g1 decreased with increased vapour pressure deficit of the atmosphere. From day 1 of the withholding water, predawn and midday leaf water potentials (Ψpd and Ψmd) decreased, reaching in both cases minimum values of -5.5 MPa, with no significant differences between mature and young leaves. Water stress induced stomatal closure, leaf rolling and partial defoliation. No osmotic adjustment was found in response to water stress in either leaf type, but both were able to enhance the cell wall elasticity (elastic adjustment). After rewatering, leaf water potential recovered quickly (within 2 d) but g1 did not. This revised version was published online in July 2006 with corrections to the Cover Date.     

Elucidating genomic regions determining enhanced leaf growth and delayed senescence in elevated CO2

Limited information is available on the genetic variation and control for plant growth response to elevated CO(2) (e[CO(2)]). Such information is necessary to understand plant adaptation and evolution in future rising CO(2). Here, quantitative trait loci (QTL) for leaf growth, development, quality and leaf senescence were determined in a tree pedigree - an F(2) hybrid of Populus trichocarpa T. & G and Populus deltoides Marsh, following season-long exposure to either current day ambient carbon dioxide (a[CO(2)]) or e[CO(2)] at 600 microL L(-1). Leaf growth and development differed between the grandparents such that P. trichocarpa showed greater response to e[CO(2)]. In the F(2) generation, leaf development and quality traits including leaf area, leaf shape, epidermal cell area, and stomatal number, specific leaf area (SLA), and the phenology trait, canopy senescence index, were sensitive to e[CO(2)]. Sixty-nine QTL were mapped for the 19 traits of plants in a[CO(2)] while 60 QTL were mapped for plants in e[CO(2)]. The results suggest that although many QTL mapped to common positions in a[CO(2)] and e[CO(2)], confirming their importance in determining growth, there was also differential genetic control for a number of traits including leaf senescence. Candidate genes were shown to collocate to regions where response QTL mapped. This study is the first to identify candidate genes that may be important in determining plant adaptation to future high-CO(2) world.     

Differential stress responses of antioxidative systems to drought in pendunculate oak (Quercus robur) and maritime pine (Pinus pinaster) grown under high CO(2) concentrations

Maritime pine (Pinus pinaster), a drought-avoiding species, contained 2--4-fold lower activities of superoxide dismutase, ascorbate peroxidase, catalase, dehydroascorbate reductase, and glutathione reductase than pendunculate oak (Quercus robur), a drought-tolerant species. The levels of ascorbate, monodehydroascorbate radical reductase activity, and glutathione in pine needles were similar to those in oak leaves. In both species the development of drought stress, characterized by decreasing predawn water potentials, caused gradual reductions in antioxidant protection, increased lipid peroxidation, increased oxidation of ascorbate and glutathione and in pine also significant loss in soluble proteins and carotenoids. These results support the idea that increased drought-tolerance in oak as compared with pine is related to increased biochemical protection at the tissue level. To test the hypothesis that elevated CO(2) ameliorated drought-induced injury, young oak and pine trees acclimated to high CO(2) were subjected to drought stress. Analysis of plots of enzymatic activities and metabolites against predawn water potentials revealed that the drought stress-induced decreases in antioxidant protection and increases in lipid peroxidation were dampened at high CO(2). In pine, protein and pigment degradation were also slowed down. At high CO(2), superoxide dismutase activities increased transiently in drought-stressed trees, but collapsed in pine faster than in oak. These observations suggest that the alleviation of drought-induced injury under elevated CO(2) is related to a higher stability of antioxidative enzymes and an increased responsiveness of SOD to stressful conditions. This ameliorating mechanism existed independently from the effects of elevated CO(2) on plant water relations and is limited within a species-specific metabolic window.