Photosynthesis Parameters in Two Cultivars of Mulberry Differing in Salt Tolerance

Three-month-old mulberry (Morus alba L.) cultivars (salt tolerant cv. S1 and salt sensitive cv. ATP) were subjected to different concentrations of NaCl for 12 d. Leaf area, dry mass accumulation, total chlorophyll (Chl) content, net CO₂ assimilation rate (P _N), stomatal conductance (g _s), and transpiration rate (E) declined, and intercellular CO₂ concentration (C _i) increased. The changes in these parameters were dependent on stress severity and duration, and differed between the two cultivars. The tolerant cultivar showed a lesser reduction in P _N and g _s coupled with a better C _i and water use efficiency (WUE) than the sensitive cultivar. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effect of drought on ear and flag leaf photosynthesis of two wheat cultivars differing in drought resistance

We investigated net photosynthetic rate (P_N) of ear and two uppermost (flag and penultimate) leaves of wheat cultivars Hongmangmai (drought resistant) and Haruhikari (drought sensitive) during post-anthesis under irrigated and non-irrigated field conditions. The P_Nof ear and flag leaf were significantly higher and less affected by drought in Hongmangmai than in Haruhikari. The rate of reduction in stomatal conductance (g_s) was similar for the two cultivars, but intercellular CO₂concentration (C_i) in the flag leaf of Hongmangmai was lower than that of Haruhikari in non-irrigated treatment. No differences were observed in leaf water potential (₁) and osmotic adjustment of the flag leaf of the cultivars. These results imply that differences in photosynthetic inhibition on the flag leaf at low leaf ₁between the cultivars were primarily due to non-stomatal effects. Hence the main physiological factor associated with yield stability of Hongmangmai under drought stress may be attributed to the capacity for chloroplast activity in the flag leaf, which apparently allows sustained P_Nof flag leaf during grain filling under drought stress. The higher P_Nof ear in Hongmangmai under drought could also be related to its drought resistance.This revised version was published online in March 2005 with corrections to the page numbers.     

Effect of Foliar Application of Chitin and Chitosan Oligosaccharides on Photosynthesis of Maize and Soybean

On the first day after foliar application, chitosan pentamer (CH5) and chitin pentamer (CHIT5) decreased net photosynthetic rate (P _N) of soybean and maize, however, on subsequent days there was an increase in P _N in some treatments. CH5 caused an increase in maize P _N on day 3 at 10^–5 and 10^–7 M; the increases were 18 and 10 % over the control plants. This increase was correlated with increases in stomatal conductance (g _s) and transpiration rate (E), while the intercellular CO₂ concentration (C _i) was not different from the control plants. P _N of soybean plants did not differ from the control plants except for treatment CH5 (10^–7 M) which caused an 8 % increase on day 2, along with increased g _s, E, and C _i. On days 5 and 6 the CHIT5 treatment caused a 6–8 % increase in P _N of maize, which was accompanied by increases in g _s, E, and C _i. However, there was no such increase for soybean plants treated with CHIT5. In general, foliar application of high molecular mass chitin (CHH) resulted in decreased P _N, particularly for 0.010 % treated plants, both in maize and soybean. Foliar applications of chitosan and chitin oligomers did not affect (p > 0.05) maize or soybean height, root length, leaf area, shoot or root or total dry mass.     

Ionic and Osmotic Effects of Salinity on Single-Leaf Photosynthesis in Two Wheat Cultivars with Different Drought Tolerance

The effects of iso-osmotic salinity and drought stresses on leaf net photosynthetic rate (P _N) in two wheat (Triticum aestivum L.) cultivars BR 8 and Norin 61, differing in drought tolerance, were compared. In drought-sensitive Norin 61, the decline of P _N was larger than that in drought-tolerant BR 8. Under NaCl treatment, P _N decreased in two phases similarly in both cultivars. In the first phase, photosynthetic depression was gradual without any photochemical changes. In the second phase, photosynthetic depression was rapid and accompanied with a decline of the energy conversion efficiency in photosystem 2 (_PS2). Our observations suggest that the osmotic factor may induce a gradual depression of photosynthesis due to stomatal closure under both stress treatments. However, under NaCl treatment, a ionic factor (uptake and accumulation of excess Na⁺) may have direct effects on electron transport and cause more severe photosynthetic depression. The drought tolerance mechanism of BR 8 was insufficient to maintain single-leaf photosynthesis under salinity.     

Gas Exchange Characteristics and Water Relations in Some Elite Okra Cultivars Under Water Deficit

Thirty-days-old plants of two cultivars of okra (Hibiscus esculentus L.), Sabzpari and Chinese-red, were subjected for 30 d to two water regimes (100 and 60 % field capacity). Leaf water potential and osmotic potential of both lines decreased significantly with the imposition of drought. Both the leaf pressure potential and osmotic adjustment were much lower in Chinese-red than those in Sabzpari. Chlorophyll (Chl) b content increased, whereas Chl a content remained unchanged and thus Chl a/b ratios were reduced in both lines. Drought stress also caused a significant reduction in net photosynthetic rate (P _N), transpiration rate (E), stomatal conductance (g _s), and water use efficiency (WUE) especially in cv. Sabzpari. The lines did not differ in intrinsic WUE (P _Ng_s) or intercellular/ambient CO₂ ratio. Overall, the growth of two okra cultivars was positively correlated with P _N, but not with g _s or P _N/E, and negatively correlated with osmotic adjustment.     

Changes of leaf water potential and gas exchange during and after drought in triticale and maize genotypes differing in drought tolerance

Influence of drought (D) on changes of leaf water potential (Ψ) and parameters of gas exchange in D-resistant and D-sensitive genotypes of triticale and maize was compared. Soil D (from −0.01 to −2.45 MPa) was simulated by mannitol solutions. At −0.013 MPa significant differences in Ψ, net photosynthetic rate (P _N), transpiration rate (E), stomatal conductance (g _s), and internal CO₂ concentration (C _i) of D-resistant and D-sensitive triticale and maize genotypes were not found. Together with the increase in concentration of the mannitol solution the impact of D on E and g _s for D-sensitive genotypes (CHD-12, Ankora) became lower than for the D-resistant ones (CHD-247, Tina). Inversely, impact of D on Ψ was higher in D-sensitive than D-resistant genotypes. From 1 to 3 d of D, a higher decrease in P _N was observed in D-resistant genotypes than in the D-sensitive ones. Under prolonged D (5–14 d) and simultaneous more severe D the decrease in P _N was lower in D-resistant than in D-sensitive genotypes. Changes in Ψ, P _N, E, and g _s caused by D in genotypes differing in the drought susceptibility were similar for triticale and maize. Compared to control plants, increase of C _i was different for triticale and maize genotypes. Hence one of the physiological reasons of different susceptibility to D between sensitive and resistant genotypes is more efficient protection of tissue water status in resistant genotypes reflected in higher decrease in g _s and limiting E compared to the sensitive ones. Other reason, observed in D-resistant genotypes during the recovery from D-stress, was more efficient removal of detrimental effects of D.     

Peanut Photosynthesis Under Drought and Re-Watering

The photosynthetic response of three Arachis hypogaea L. cultivars (57-422, 73-30, and GC 8-35) grown for two months was measured under water available conditions, severe water stress, and 24, 72, and 93 h following re-watering. At the end of the drying cycle, all the cultivars reached dehydration, relative water content (RWC) ranging between 40 and 50 %. During dehydration, leaf stomatal conductance (g _s), transpiration rate (E), and net photosynthetic rate (P _N) decreased more in cvs. 57-422 and GC 8-35 than in 73-30. Instantaneous water use efficiency (WUE_i) and photosynthetic capacity (P _max) decreased mostly in cv. GC 8-35. Except in cv. GC 8-35, the activity of photosystem 1 (PS1) was only slightly affected. PS2 and ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPCO) were the main targets of water stress. After re-watering, cvs. 73-30 and GC 8-35 rapidly regained g _s, E, and P _N activities. Twenty-four hours after re-watering, the electron transport rates and RuBPCO activity strongly increased. P _N and P _max fully recovered later. Considering the different photosynthetic responses of the studied genotype, a general characterisation of the interaction between water stress and this metabolism is presented.     

Effects of Pre-Treatments with Abscisic Acid and/or Benzyladenine on Gas Exchange of French Bean, Sugar Beet, and Maize Leaves During Water Stress and After Rehydration

Net photosynthetic rate (P_N), transpiration rate (E), and stomatal conductance (g_s) during water stress and after rehydration were measured in Phaseolus vulgaris, Beta vulgaris, and Zea mays. Immediately before imposition of water stress by cessation of watering, plants were irrigated with water (control), 100 M abscisic acid (ABA), and/or 10 M N⁶-benzyladenine (BA). In all three species, application of ABA decreased g_s, E, and P_N already 1 h after application. However, during water stress g_s, E, and P_N in plants pre-treated with ABA remained higher than in plants pre-treated with water. Positive effects of ABA application were observed also after rehydration. In contrast, the effects of pre-treatment with BA were species-specific. While in bean plants BA application ameliorated negative effect of water stress, only very slight effects were observed in maize, and in sugar beet BA even aggravated the effects of water stress.     

Diurnal Gas Exchange and Superior Resources Use Efficiency of Typical C4 Species in Hunshandak Sandland,China

Net photosynthetic rate (P _N), transpiration rate (E), stomatal conductance (g _s), leaf water potential (ψ_leaf), leaf nitrogen content, and photosynthetic nitrogen use efficiency (PNUE) were compared between a typical C₄ plant, Agriophyllum squarrosum and a C₃ plant, Leymus chinensis, in Hunshandak Sandland, China. The plant species showed different diurnal gas exchange patterns on June 12–14 when photosynthetic photon flux density (PPFD), air temperature (T _air), and water potential were moderate. P _N, E, and g _s of A. squarrosum showed distinct single peak while those of L. chinensis were depressed at noon and had two peaks in their diurnal courses. Gas exchange traits of both species showed midday depression under higher photosynthetic photon flux density (PPFD) and T _air when Ψ_leaf was significantly low down on August 6–8. However, those of A. squarrosum were depressed less seriously. Moreover, A. squarrosum had higher P _N, Ψ_leaf, water use efficiency (WUE), and PNUE than L. chinensis. Thus A. squarrosum was much more tolerant to heat and high irradiance and could utilise the resources on sand area more efficiently than L. chinensis. Hence species like A. squarrosum may be introduced and protected to reconstruct the degraded sand dunes because of their higher tolerance to stress and higher resource use efficiency. This revised version was published online in August 2006 with corrections to the Cover Date.     

Photosynthetic photon flux density,carbon dioxide concentration,and vapor pressure deficit effects on photosynthesis in cacao seedlings

Independent short-term effects of photosynthetic photon flux density (PPFD) of 50–400 μmol m⁻² s⁻¹, external CO₂ concentration (C _a) of 85–850 cm³ m⁻³, and vapor pressure deficit (VPD) of 0.9–2.2 kPa on net photosynthetic rate (P _N), stomatal conductance (g _s), leaf internal CO₂ concentration (C _i), and transpiration rates (E) were investigated in three cacao genotypes. In all these genotypes, increasing PPFD from 50 to 400 μmol m⁻² s⁻¹ increased P _N by about 50 %, but further increases in PPFD up to 1 500 μmol m⁻² s⁻¹ had no effect on P _N. Increasing C _a significantly increased P _N and C _i while g _s and E decreased more strongly than in most trees that have been studied. In all genotypes, increasing VPD reduced P _N, but the slight decrease in g _s and the slight increase in C _i with increasing VPD were non-significant. Increasing VPD significantly increased E and this may have caused the reduction in P _N. The unusually small response of g _s to VPD could limit the ability of cacao to grow where VPD is high. There were no significant differences in gas exchange characteristics (g _s, C _i, E) among the three cacao genotypes under any measurement conditions.     

Effect of Cd on growth, photosynthetic gas exchange, and chlorophyll fluorescence of wild and Cd-sensitive mutant rice

Growth, photosynthetic gas exchange, and chlorophyll fluorescence characteristics were investigated in wild type (WT) and Cd-sensitive mutant rice (Oryza sativa L.) plants using 50 μM Cd treatment for 12 d followed by a 3-d recovery. Under Cd stress, net dry mass and pigment contents were significantly lower in the mutant plants than in the WT. The mutant had lower net photosynthetic rate (P _N), transpiration rate (E), and stomatal conductance (g _s) than WT rice, however, it had higher intercellular CO₂ concentration (C _i), indicating that non-stomatal factors accounted for the inhibition of P _N. Maximal photochemical efficiency of photosystem 2 (F_v/F_m), effective quantum yield of PS2 (Φ_PS2), and photochemical quenching (q_P) decreased much in the mutant under Cd stress. Cd content in roots and leaves of the mutant was significantly higher than those in the WT. Hence Cd toxicity was associated with the marked increases in Cd contents of plant tissue. After the recovery for 3 d, the WT rice had higher capacity to recover from Cd injury than the mutant.     

Gas Exchange,Leaf Structure,and Hydraulic Features in Relation to Sex,Shoot Form,and Leaf Form in An Evergreen Shrub Sabina vulgaris in the Semi-Arid Mu Us Sandland in China

We examined differences in net photosynthetic rate (P _N), transpiration rate (E), water use efficiency (WUE), ratio of substomatal to atmospheric CO₂ concentration (C _i/C _a), cuticle thickness (CT), epidermis cell size (ECS), mesophyll cell size (MCS), vascular bundle size (VBS), tissue density (TD), and coefficient of water loss (k) in Sabina vulgaris as related to sex, shoot form, and leaf form. P _N, E, WUE, C _i/C _a, MCS, VBS, and k varied with sex, whereas CT, ECS, and TD did not. These differences in physiology and anatomy between the female and male plants may be closely related with their reproduction behaviour. P _N, E, C _i/C _a, CT, ECS, MCS, and VBS were significantly smaller in the erect shoots than in the prostrate shoots, WUE was just opposite; TD and k did not vary with shoot form. These changes in physiology with shoot form indicate that erect shoots may be more tolerant of water stress than prostrate shoots. P _N, E, C _i/C _a, TD, and k were significantly greater in the spine leaves than in the scale leaves, whereas WUE, CT, ECS, MCS, and VBS followed the opposite trends. The changes in physiology and anatomy with leaf form suggest that scale leaves have higher drought-resistant and water-holding capacities than spine leaves. Measurements of field gas exchange showed that three-year-old seedlings had lower drought-resistance and higher water loss than five-year-old seedlings, which provides some evidence that seedling survival decreases with decreasing plant age.     

Tolerance to Water Stress in Tomato Cultivars

The effects of plant water stress imposed at vegetative, flowering, and fruiting stages of four cultivars of tomato (Lycopersicon esculentum Mill.) on net photosynthetic rate (P _N), stomatal conductance (g _s), transpiration rate (E), osmotic adjustment, and crop water stress index (CWSI) were investigated. Osmotic adjustment was the highest in cv. Arka Meghali, followed by cv. RFS-1. CWSI was lowest in cv. Arka Meghali and highest in cv. Pusa Ruby. Significant reduction in g _s, E, and P _N was observed in all the cultivars. The maximum reduction in E was observed in cv. Arka Saurabh during the fruiting stage (62.4 %) and maximum reduction in P _N at the flowering stage in Pusa Ruby (53.1 %). Maximum P _N was observed in Arka Meghali under water stress. The values of internal CO₂ concentration (C _i) did not follow the decrease in g _s which might be taken as an indication of mesophyll (non-stomatal) limitation to P _N. Magnitude of P _N decrease accompanying g _s reductions varied in the four cultivars. Arka Meghali which had highest rate of gas exchange efficiency (P _N/g _s) under water deficits can be recommended for rainfed cultivation. This revised version was published online in August 2006 with corrections to the Cover Date.     

Photoprotective Function of Photorespiration in Several Grapevine Cultivars Under Drought Stress

Four grapevine cultivars, i.e. Cabernet Sauvignon (a member of the Western Europe cultivar group), Rizamat (a member of the East cultivar group), Red Double Taste (a hybridized cultivar from Vitis vinifera L. and V. labrusca L.), and 1103Paulsen (a hybridized rootstock), were treated by three severity orders of drought stress for 25 d. Then net photosynthetic rate (P _N), maximal photochemical efficiency (F_v/F_m), actual photochemical efficiency (_PS2) of photosystem 2, total electron transport rate (J_T), and electron transport flows used in carboxylation (J_C) and in oxygenation (J_O) reactions catalysed by ribulose-1,5-bisphosphate carboxylase/oxygenase were determined. P _N was determined again after re-watering for 2 d by gas exchange measurement. Along with the increase in severity of drought stress, P _N, F_v/F_m, _PS2, J_T, and J_C in all four cultivars decreased. The range of decrease differed among cultivars. J_O expressed various trends from cultivar to cultivar. In Rizamat that received slight and moderate drought stress, P _N evidently decreased, but J_O markedly increased, thus maintaining high values of J_T and _PS2. Prior to the moderate drought stress, the F_v/F_m was high in Rizamat, indicating that the photodamage had not happened ahead of the moderate drought stress given. Under the severe drought stress, the photorespiration rate in Rizamat decreased by 70 %, and J_T, _PS2, and F_v/F_m also dropped to very low values, i.e. the photodamage of photosynthetic apparatus has taken place. This suggested that the photorespiration has consumed the excessive assimilatory power and the photo-protective function of photorespiration is very important for Rizamat. When Cabernet Sauvignon grew under drought stress, its J_O decreased in a small range, thus maintaining higher values of J_C, J_T, _PS2, and F_v/F_m; hence no serious photodamage occurred. Despite of the fact that P _N of cv. Red Double Taste decreased markedly under the slight drought stress, J_O still increased under the severe drought stress. This suggests that photorespiration is important in photoprotection under drought stress. J_O in cv. 1103Paulsen markedly decreased under slight stress. Accordingly, P _N, F_v/F_m, _PS2, J_T, and J_C decreased to extremely low values. Thus photorespiration effectively protects the photosynthetic apparatus from photo-damage under drought, assists in maintaining a relatively high _PS2, and helps P _N to be rapidly recovered after re-watering.     

Responses of Tobacco Plantlets to Change of Irradiance During Transfer from in vitro to ex vitro Conditions

Chlorophyll a fluorescence kinetics, net photosynthetic rate (P _N), water relations, and photosynthetic pigment contents were studied during acclimation of in vitro grown tobacco to higher irradiance (HL; 700 mol m^–2 s^–1). Plantlets were grown on medium containing sucrose in glass vessels (G-plants) or in Magenta boxes (M-plants) with better CO₂ supply in the latter ones. The effect of HL was studied either (1) in plantlets grown under original in vitro conditions (closed vessels), (2) in in vitro plantlets exposed to ambient CO₂ concentration (covers removed), or (3) in plantlets transplanted to ex vitro into pots with sand and nutrient solution. Higher P _N, and fraction of closed photosystem 2 (PS2) centres (1 – q_P), and lower content of xanthophyll cycle pigments were found in M-plants compared to G-plants. HL treatment caused photoinhibition particularly in plants kept in closed vessels. This was indicated by the decrease in the ratio of F_v/F_m and by the increase in non-photochemical quenching, 1 – qp, and content of xanthophyll cycle pigments. Better CO₂ supply ensured by the removal of closure lead to the moderate reduction of symptoms of photoinhibition, although stomatal conductance (g _s), transpiration rate (E), and P _N were negatively affected. The main reason was the decrease in relative air humidity, which caused similar reduction of P _N, E, and g _s after the transfer of plantlets to ex vitro. Nevertheless, plant response to HL seemed not to be affected by any possible root injury caused by transfer to ex vitro. The differences in contents of xanthophyll cycle pigments, degree of de-epoxidation, P _N, and quenching parameters between M- and G-plantlets were still significant 7 d after ex vitro transfer and HL acclimation.     

Diurnal Changes of Gas Exchange,Chlorophyll Fluorescence,and Stomatal Aperture of Hybrid Poplar Clones Subjected to Midday Light Stress

Diurnal changes in net photosynthetic rate (P _N), chlorophyll (Chl) fluorescence, and stomatal aperture of several hybrid poplar clones subjected to midday light stress were measured in July and August of 1996. Midday depression of P _N, photosystem 2 (PS2) efficiency, stomatal conductance (g _s), and stomatal aperture was observed in all clones, though at differing rates among them. Non-uniform stomatal closure occurred at noon and at other times, requiring a modification of intercellular CO₂ concentration (C ₁). A linear relationship was found between g _s and stomatal aperture. More than half of the photons absorbed by PS2 centre dissipated thermally when subjected to light stress at noon. There was a linear relationship between the rate of PS2 photochemical electron transport (PxPFD) and P _N. There was a consensus for two fluorescence indicators (1 – q_P/q_N and (F_m' – F)/F_m') in assessment of susceptibility of photoinhibition in the clones. According to P _N, Chl fluorescence, and stomatal aperture, we conclude that midday depression of photosynthesis can be attributed to both stomatal and non-stomatal limitations.     

Gas Exchange Characteristics of the Flue-Cured Tobacco (Nicotiana Tabacum L.) Cultivars Grown under Two Production Systems

The net photosynthetic rate (P _N), intercellular CO₂ concentration (C _i), stomatal conductance (g _s), transpiration rate (E), water use efficiency (WUE), and leaf biomass production of four American flue-cured tobacco (Nicotiana tabacum L.) cultivars K 326, K 358, and Speight G 28 were compared with three local Indian cultivars 16/103, Special FCV, and PCT-7, during 1994 and 1995 crop seasons under irrigated and rainfed production systems (Northern light soils, NLS, and Karnataka light soils, KLS) in India. By comparison, the American tobacco cv. K 326 showed the highest P _N and g _s. A positive correlation was found between P _N and biomass production in all the varieties tested (r = 0.55 in NLS and 0.73 in KLS). The American cultivars were superior than the local cultivars in their biomass production and P _N under Indian farming conditions.     

Influence of water stress on leaf photosynthetic characteristics in wheat cultivars differing in their susceptibility to drought

A gradual reduction in leaf water potential (Ψ_leaf), net photosynthetic rate (P _N), stomatal conductance, and transpiration rate was observed in two drought tolerant (C 306 and K 8027) and two susceptible (RW 893 and 899) genotypes subjected to water stress. The extent of reduction was lower in K 8027 and C 306 and higher in RW 893 and RW 899. Rewatering the plants after 5 d of stress restored P _N and other gas exchange traits in all four cultivars. Water stress had no significant effect on variable to maximum fluorescence ratio (F_v/F_m) indicating that water stress had no effect on primary photochemistry of photosystem 2 (PS2). However, water stress reduced the efficiency of excitation energy transfer (F′_v/F′_m) and the quantum yield of electron transport (Φ_PS2). The reduction was more pronounced in susceptible cultivars. Water stress had no significant effect on photochemical quenching, however, the non-photochemical quenching increased by water stress.     

Chain Correlation between Variables of Gas Exchange and Yield Potential in Different Winter Wheat Cultivars

Variables of gas exchange of flag leaves and grain yield potentials of five representative winter wheat (Triticum aestivum L.) cultivars varied greatly across different development stages under the same management and irrigation. The cultivars with high yield potential had higher net photosynthetic rate (P _N), PPFD (photosynthetic photon flux density) saturated photosynthetic rate (P _sat), stomatal conductance (g _s), and maximum apparent quantum yield of CO₂ fixation (_m,app) than those with low grain yield, but their dark respiration rate (R _D) and compensation irradiance (I _c) were remarkably lower. Compared with overall increase of yield potential of 71 % from low yield cultivars to high yield ones, P _N, P _sat, _m,app, and g _s were 13, 19, 57, and 32 % higher, respectively; but R _D and I _c decreased by 19 and 76 %, respectively. Such difference was evidently large during anthesis stage (e.g., P _N by 33 %), which indicated that this period could be the best for assisting further selection for better cultivars. However, transpiration rate (E) and water use efficiency (WUE) differed only little. At different development stages, especially at anthesis, P _N and P _sat were positively correlated with _m,app, g _s, and yield potential, and negatively correlated with R _D and I _c. Thus the high-yield-potential winter wheat cultivars possess many better characters in photosynthesis and associated parameters than the low-yield cultivars.     

Photosynthesis Parameters in Two Cultivars of Mulberry Differing in Salt Tolerance

Three-month-old mulberry (Morus alba L.) cultivars (salt tolerant cv. S1 and salt sensitive cv. ATP) were subjected to different concentrations of NaCl for 12 d. Leaf area, dry mass accumulation, total chlorophyll (Chl) content, net CO₂ assimilation rate (P _N), stomatal conductance (g _s), and transpiration rate (E) declined, and intercellular CO₂ concentration (C _i) increased. The changes in these parameters were dependent on stress severity and duration, and differed between the two cultivars. The tolerant cultivar showed a lesser reduction in P _N and g _s coupled with a better C _i and water use efficiency (WUE) than the sensitive cultivar.