Photosynthesis of six barley genotypes as affected by water stress

The effect of water stress on plant water status and net photosynthetic gas exchange (PN) in six barley genotypes (Hordeum vulgare L.) differing in productivity and drought tolerance was studied in a controlled growth chamber. Osmotic adjustment (OA), PN, stomatal conductance (gs), and the ratio intercellular/ambient. CO2 concentration (Ci/Ca) were evaluated at four different levels of soil water availability, corresponding to 75, 35, 25 and 15 % of total available water. Variability in OA capacity was observed between genotypes: the drought tolerant genotypes Albacete and Alpha showed higher OA than drought susceptible genotypes Express and Mogador. The genotype Albacete exhibited also higher PN than the others at low water potential (Ψ). The ratios of PN/gs and Ci/Ca showed that differences in photosynthetic inhibition between genotypes at low Ψ were probably due to nonstomatal effects. In Tichedrett, a landrace genotype with a very extensive root development, OA was not observed, however, it exhibited a capacity to maintain its photosynthetic activity under water stress.     

Photosynthesis of six barley genotypes as affected by water stress

The effect of water stress on plant water status and net photosynthetic gas exchange (PN) in six barley genotypes (Hordeum vulgare L.) differing in productivity and drought tolerance was studied in a controlled growth chamber. Osmotic adjustment (OA), PN, stomatal conductance (gs), and the ratio intercellular/ambient. CO2 concentration (Ci/Ca) were evaluated at four different levels of soil water availability, corresponding to 75, 35, 25 and 15 % of total available water. Variability in OA capacity was observed between genotypes: the drought tolerant genotypes Albacete and Alpha showed higher OA than drought susceptible genotypes Express and Mogador. The genotype Albacete exhibited also higher PN than the others at low water potential (Ψ). The ratios of PN/gs and Ci/Ca showed that differences in photosynthetic inhibition between genotypes at low Ψ were probably due to nonstomatal effects. In Tichedrett, a landrace genotype with a very extensive root development, OA was not observed, however, it exhibited a capacity to maintain its photosynthetic activity under water stress. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effect of Water Stress on Photosynthesis in Two Mulberry Genotypes with Different drought Tolerance

Three-month-old mulberry (Morus alba L.) cultivars (drought tolerant S13 and drought sensitive S54) were subjected to water stress for 15 d. Water stress decreased the leaf water potential, net photosynthetic rate (PN), and stomatal conductance (gs) in both the cultivars. However, the magnitude of decline was comparatively greater in the sensitive cultivar (S54). Intercellular CO2 concentration (Ci) was unaltered during mild stress, but significantly increased at severe stress in both cultivars. The photosystem 2 activity significantly declined only at a severe stress in both cultivars. The Ci/gs ratio representing the mesophyll efficiency was greater in the tolerant cultivar S13. Involvement of stomatal and/or non-stomatal components in declining PN depended on the severity and duration of stress. However, the degree of non-stomatal limitations was relatively less in the drought tolerant cultivar.     

Seasonal Changes in Photosynthesis and Stomatal Conductance of Five Plant Species from a Semiarid Ecosystem

In order to determine whether stomatal closure alone regulates photosynthesis during drought under natural conditions, seasonal changes in leaf gas exchange were studied in plants of five species differing in life form and carbon fixation pathway growing in a thorn scrub in Venezuela. The species were: Ipomoea carnea, Jatropha gossypifolia, (C3 deciduous shrubs), Alternanthera crucis (C4 deciduous herb), and Prosopis juliflora and Capparis odoratissima (evergreen phreatophytic trees). Xylem water potential () of all species followed very roughly the precipitation pattern, being more closely governed by soil water content in I. carnea and A. crucis. Maximum rate of photosynthesis, Pmax, decreased with in I. carnea, J. gossypifolia, and A. crucis. In I. carnea and J. gossypifolia stomatal closure was responsible for a 90 % decline in net photosynthetic rate (PN) as decreased from -0.3 to -2.0 MPa, since stomatal conductance (gs) was sensitive to water stress, and stomatal limitation on PN increased with drought. In A. crucis, PN decreased by 90 % at a much lower (-9.3 MPa), and gs was relatively less sensitive to . In P. juliflora and C. odoratissima, Pmax, gs, and intercellular CO2 concentration (Ci) were independent of soil water content. In the C3 shrubs stomatal closure was apparently the main constraint on photosynthesis during drought, Ci declining with in I. carnea. In the C4 herb, Ci was constant along the range of values, which suggested a coordinated decrease in both gs and mesophyll capacity. In P. juliflora Ci showed a slow decrease with which may have been due to seasonal leaf developmental changes, rather than to soil water availability.     

水分胁迫对冬小麦CO_2同化作用的影响

比较了两个小麦品种陕合6号和郑引1号经受不同程度的水分胁迫处理后,叶片多种光合参数:净光合速率(P_n)、气孔导度(G_s)、细胞间隙CO_2浓度(G_i)、表观量子需要量和羧化效率以及Rubis CO蛋白量与活性等的变化。在轻度水分胁迫下,叶片光合速率降低的根本原因在于气孔导度的下降;而在严重胁迫下,非气孔因素起主要作用。     

The effects of soil and atmospheric drought on photosynthesis and stomatal control of gas exchange in three coniferous species

The responses of steady-state CO₂ assimilation rate (A), transpiration rate (E), and stomatal conductance (g_s) to changes in leaf-to-air vapour pressure difference (δW) on one hand and to increasing soil drought on the other hand were examined in 2-year-old seedlings of Pseudotsuga menziesii, Pseudotsuga macrocarpa and Cedrus atlantica. Analysing the data through A vs intercellular CO₂ molar fraction (c_i) graphs, we could determine stomatal and mesophyll contributions to changes in A as δW or soil drought were increased. Increasing soil drought affected g_s and mesophyll photosynthesis independently, since clearly distinct predawn leaf water potential (ψ_p) regions appeared in which either stomatal or mesophyll effects prevailed for explaining the changes in A. The two Pseudotsuga species exhibited a large ψ_P range (between ca -0.8 and -1.5 to -1.9 MPa) in which only stomata were responsible for the decrease in A. A dramatic decline in mesophyll photosynthesis was noticed starting from values as high as -1.2 MPa ( C. atlantica ), -1.5 MPa ( P. macrocarpa ) and -1.9 MPa ( P. menziesii ). Increasing ΔW at high soil water content led to a sharp decline in A primarily due to an alteration of mesophyll photosynthesis. Stomatal conductance for CO₂ diffusion was affected in a lesser extent and in close correlation with the changes in mesophyll photosynthesis, which could suggest the existence of a functional linkage between mesophyll photosynthesis and stomata. Surprisingly, the drought resistant P. macrocarpa exhibited the least conservative water use efficiency in response to the two types of drought. In this species drought adaptation seems to be mainly due to its high root growth and soil prospection ability.     

Comparison of Isoprene Emission, Intercellular Isoprene Concentration and Photosynthetic Performance in Water-Limited Oak (Quercus pubescens Willd. and Quercus robur L.) Saplings

Abstract: The influence of prolonged water limitation on leaf gas exchange, isoprene emission, isoprene synthase activities and intercellular isoprene concentrations was investigated under standard conditions (30 °C leaf temperature and 1000 μmol photons m^-2 s^-1 PPFD) in greenhouse experiments with five-year-old pubescent oak ( Quercus pubescens Willd.) and four-year-old pedunculate oak ( Quercus robur L.) saplings. Net assimilation rates proved to be highly sensitive to moderate drought in both oak species, and were virtually zero at water potentials (Ψ_pd) below - 1.3 MPa in Q. robur and below - 2.5 MPa in Q. pubescens . The response of stomatal conductance to water stress was slightly less distinct. Isoprene emission was much more resistant to drought and declined significantly only at Ψ_pd below - 2 MPa in Q. robur and below - 3.5 MPa in Q. pubescens . Even during the most severe water stress, isoprene emission of drought-stressed saplings was still approximately one-third of the control in Q. robur and one-fifth in Q. pubescens . Isoprene synthase activities were virtually unaffected by drought stress. Re-watering led to partial recovery of leaf gas exchange and isoprene emission. Intercellular isoprene concentrations were remarkably enhanced in water-limited saplings of both oak species during the first half of the respective drought periods with maximum mean values up to ca. 16 μl l^-1 isoprene for Q. pubescens and ca. 11 μl l^-1 isoprene for pedunculate oak, supporting the hypothesis that isoprene serves as a short-term thermoprotective agent in isoprene-emitting plant species.     

小麦和大豆叶片的气孔不均匀关闭现象

用14CO2放射自显影的方法研究了田间小麦和大豆叶片在水分胁迫下的气孔关闭状况。正常浇水的小麦和大豆叶片呈现出对14CO2的均匀吸收。在小麦与大豆"片水势分别降至-1．75和-1．32MPa的土壤干旱条件下,两种作物叶片都发生气孔不均匀关闭。离休叶片在空气中快速脱水易引起气孔不均匀关闭。正常供水小麦叶片在晴天中午明显的光合午休时,无CO2的不均匀吸收。某些晴天中午,在大豆光合午休低谷时段观察到较明显的气孔不均匀关闭,用气体交换资料计算出的细胞间隙CO2浓度并不随气孔年度的降低而下降,反而略有回升。     

The importance of xylem constraints in the distribution of conifer species

Vulnerability of stem xylem to cavitation was measured in 10 species of conifers using high pressure air to induce xylem embolism. Mean values of air pressure required to induce a 50% loss in hydraulic conductivity (φ₅₀) varied enormously between species, ranging from a maximum of 14.2±0.6 MPa (corresponding to a xylem water potential of −14.2 MPa) in the semi-arid species Actinostrobus acuminatus to a minimum of 2.3±0.2 MPa in the rainforest species Dacrycarpus dacrydioides . Mean φ₅₀ was significantly correlated with the mean rainfall of the driest quarter within the distribution of each species. The value of φ₅₀ was also compared with leaf drought tolerance data for these species in order to determine whether xylem dysfunction during drought dictated drought response at the leaf level. Previous data describing the maximum depletion of internal CO₂ concentration (c_i) in the leaves of these species during artificial drought was strongly correlated with φ₅₀ suggesting a primary role of xylem in effecting leaf drought response. The possibility of a trade-off between xylem conductivity and xylem vulnerability was tested in a sub-sample of four species, but no evidence of an inverse relationship between φ₅₀ and either stem-area specific (K_a) or leaf-area specific conductivity (K₁) was found.     

Stomatal and nonstomatal regulation of water use in cotton, corn, and sorghum

Stomata of corn (Zea mays L.) and sorghum (Sorghum bicolor L.) responded to changes in leaf water potential during the vegetative growth phase. During reproductive growth, leaf resistances were minimal and stomata were no longer sensitive to bulk leaf water status even when leaf water potentials approached −27 bars. Stomata of corn, cotton (Gossypium hirsutum L.), and sorghum appear to respond to changes in the humidity deficit between the leaf and air and in this manner, regulated transpirational flux to some degree. Distinct differences in water transport efficiency were observed in the three species. Under nonlimiting soil water conditions, sorghum exhibited the greatest efficiency of water transport while under limiting soil moisture conditions, cotton appeared most efficient. Corn was the least efficient with respect to nonstomatal regulation of water use. Differences in drought tolerance among the three species are partially dependent on stomatal regulation of water loss, but efficiency of the water transport system may be more related to drought adaptation. This is particularly important since stomata of all three species did not respond to bulk leaf water status during a large portion of the growing season.     

Spatial Distribution of Photosynthesis during Drought in Field-Grown and Acclimated and Nonacclimated Growth Chamber-Grown Cotton

Inhomogeneous photosynthetic activity has been reported to occur in drought-stressed leaves. In addition, it has been suggested that these water stress-induced nonuniformities in photosynthesis are caused by “patchy” stomatal closure and that the phenomenon may have created the illusion of a nonstomatal component to the inhibition of photosynthesis. Because these earlier studies were performed with nonacclimated growth chamber-grown plants, we sought to determine whether such “patches” existed in drought-treated, field-grown plants or in chamber-grown plants that had been acclimated to low leaf water potentials (ψ_leaf). Cotton (Gossypium hirsutum L.) was grown in the field and subjected to drought by withholding irrigation and rain from 24 d after planting. The distribution of photosynthesis, which may reflect the stomatal aperture distribution in a heterobaric species such as cotton, was assayed by autoradiography after briefly exposing attached leaves of field-grown plants to ¹⁴CO₂. A homogeneous distribution of radioactive photosynthate was evident even at the lowest ψ_leaf of −1.34 MPa. “Patchiness” could, however, be induced by uprooting the plant and allowing the shoot to air dry for 6 to 8 min. In parallel studies, growth chamber-grown plants were acclimated to drought by withholding irrigation for three 5-d drought cycles interspersed with irrigation. This drought acclimation lowered the ψ_leaf value at which control rates of photosynthesis could be sustained by approximately 0.7 MPa and was accompanied by a similar decline in the ψ_leaf at which patchiness first appeared. Photosynthetic inhomogeneities in chamber-grown plants that were visible during moderate water stress and ambient levels of CO₂ could be largely removed with elevated CO₂ levels (3000 μL L⁻¹), suggesting that they were stomatal in nature. However, advanced dehydration (less than approximately 2.0 MPa) resulted in “patches” that could not be so removed and were probably caused by nonstomatal factors. The demonstration that patches do not exist in drought-treated, field-grown cotton and that the presence of patches in chamber-grown plants can be altered by treatments that cause an acclimation of photosynthesis leads us to conclude that spatial heterogeneities in photosynthesis probably do not occur frequently under natural drought conditions.     

Evidence for involvement of photosynthetic processes in the stomatal response to CO2

Stomatal conductance (gs) typically declines in response to increasing intercellular CO2 concentration (ci). However, the mechanisms underlying this response are not fully understood. Recent work suggests that stomatal responses to ci and red light (RL) are linked to photosynthetic electron transport. We investigated the role of photosynthetic electron transport in the stomatal response to ci in intact leaves of cocklebur (Xanthium strumarium) plants by examining the responses of gs and net CO2 assimilation rate to ci in light and darkness, in the presence and absence of the photosystem II inhibitor 3-(3,4-dichlorophenyl)-1,1-dimethylurea (DCMU), and at 2% and 21% ambient oxygen. Our results indicate that (1) gs and assimilation rate decline concurrently and with similar spatial patterns in response to DCMU; (2) the response of gs to ci changes slope in concert with the transition from Rubisco- to electron transport-limited photosynthesis at various irradiances and oxygen concentrations; (3) the response of gs to ci is similar in darkness and in DCMU-treated leaves, whereas the response in light in non-DCMU-treated leaves is much larger and has a different shape; (4) the response of gs to ci is insensitive to oxygen in DCMU-treated leaves or in darkness; and (5) stomata respond normally to RL when ci is held constant, indicating the RL response does not require a reduction in ci by mesophyll photosynthesis. Together, these results suggest that part of the stomatal response to ci involves the balance between photosynthetic electron transport and carbon reduction either in the mesophyll or in guard cell chloroplasts.     

Influence of germination date on Dioon edule (Zamiaceae) seedling tolerance to water stress

Dioon edule seedling mortality is mostly attributed to dehydration by prolonged drought, even when they present xeromorphic characteristics like the adult plants. The effect of germination date (GD) and soil water deficit on seedling tolerance to water stress was assessed. The seedlings germinated and grown from mature seeds every month from December to April GD were selected to evaluate the leaf area, photosynthetic pigment content, crassulacean acid metabolism (CAM) activity, stomatal conductance (gs) and leaflet anatomy at soil water potential (Ψs) of 0.0 MPa (day 1), ?0.1 MPa (day 40), ?1.0 MPa (day 90), ?1.5 MPa (day 130), and a control (0.0 MPa at day 130) to recognize differences due to leaf development. The seedlings shifted from C₃ to CAM cycling when exposed to water stress at Ψs of ?1.0 MPa, like adult plants. The March–April GD seedlings with undeveloped sclerified hypodermis and stomata, presented reduced leaf area, lower Chlorophyll a/b ratio, higher CAM activity and midday partial stomatal closure when reached Ψs of ?1.0 MPa. These have higher probability of dehydration during severe drought (February–April) than those of the December–February GD with similar Ψs. Plants used for restoration purposes must have full leaf development to increase the survival.     

Soil and plant water relations determine photosynthetic responses of C3 and C4 grasses in a semi-arid ecosystem under elevated CO2

To model the effect of increasing atmospheric CO2 on semi-arid grasslands, the gas exchange responses of leaves to seasonal changes in soil water, and how they are modified by CO2, must be understood for C3 and C4 species that grow in the same area. In this study, open-top chambers were used to investigate the photosynthetic and stomatal responses of Pascopyrum smithii (C3) and Bouteloua gracilis (C4) grown at 360 (ambient CO2) and 720 micro mol mol-1 CO2 (elevated CO2) in a semi-arid shortgrass steppe. Assimilation rate (A) and stomatal conductance (gs) at the treatment CO2 concentrations and at a range of intercellular CO2 concentrations and leaf water potentials (psileaf) were measured over 4 years with variable soil water content caused by season and CO2 treatment. Carboxylation efficiency of ribulose bisphosphate carboxylase/oxygenase (Vc,max), and ribulose bisphosphate regeneration capacity (Jmax) were reduced in P. smithii grown in elevated CO2, to the degree that A was similar in elevated and ambient CO2 (when soil moisture was adequate). Photosynthetic capacity was not reduced in B. gracilis under elevated CO2, but A was nearly saturated at ambient CO2. There were no stomatal adaptations independent of photosynthetic acclimation. Although photosynthetic capacity was reduced in P. smithii growing in elevated CO2, reduced gs and transpiration improved soil water content and psileaf in the elevated CO2 chambers, thereby improving A of both species during dry periods. These results suggest that photosynthetic responses of C3 and C4 grasses in this semi-arid ecosystem will be driven primarily by the effect of elevated CO2 on plant and soil water relations.     

Diurnal and Seasonal Changes in Prunus Amygdalus Gas Exchanges

Diurnal courses in net photosynthetic rate (PN), stomatal conductance (gs), leaf water potential (), internal CO2 concentration (ci), and water use efficiency (WUE) were studied as season progressed, in relation to environmental factors in field grown Prunus amygdalus. In sun leaves PN reached maximum between 09:00 to 11:00 h and subsequently declined when high temperature and low humidity occurred. An increase was observed late in the afternoon. A decrease in gs and was found as season progressed in both years of measurements. In periods of high evaporative demand, was very low, however, it did not explain the reductions of PN in all the three periods (spring, early and late summer). Midday depression of PN and gs seemed to be related with leaf temperature (Tl) and high irradiance. Increase in ci and F0 and decrease in Fv/Fm found between 12:00 and 14:00 h corresponded to the decrease in PN. Therefore, a transient modification of photosynthetic machinery might be considered. WUE was negatively correlated with vapour pressure difference of leaf to air, that decreased during the day. The September values, higher than in the previous months, were due to the lower seasonal decreases in PN than in gs.     

Drought response of two Mexican oak species, Quercus laceyi and Q. sideroxyla (Fagaceae), in relation to elevational position

To investigate the relationship between the altitudinal distribution of Quercus laceyi and Q. sideroxyla and their physiological responses to drought, we measured relative water content (RWC), water potentials (Ψ(predawn) and Ψ(midday)), photosynthesis (A(max)), stomatal conductance (g), chlorophyll fluorescence (F(v)/F(m)), and spectral reflectance (400-1100 nm) five times during a 7 wk acute drought. Quercus laceyi was drought tolerant, while Q. sideroxyla was a drought avoider; Q. laceyi tolerated lower RWC (Q. sideroxyla = 54%, Q. laceyi = 44%), Ψ(pd) (Q. sideroxyla = -2.6 MPa, Q. laceyi = -3.3 MPa), and Ψ(md) (Q. sideroxyla = -4.5 MPa, Q. laceyi = -6.6 MPa). The F(v)/F(m) also declined first in Q. sideroxyla in wk 6, whereas F(v)/F(m) did not decline in Q. laceyi until wk 7. A(max) and g fell in wk 4, 6, and 7 in drought seedlings of both species, suggesting a decline in CO(2) assimilation during the drought. Leaf spectral reflectance increased with time in response to decreases in leaf photosynthetic pigment concentrations in latter weeks of the drought. The results suggest a close association between the altitudinal distributions of these species and their adaptation to water stress.     

Response of eucalypt species to drought

The response of three eucalypt species (Eucalyptus pulchella, Eucalyptus coccifera and Eucalyptus delegatensis) to a severe drought in the summer of 1982/83 was examined at Snug Plains, south-eastern Tasmania. Few large differences in leaf water potential (Ψ_l) or stomatal conductance (gs) were apparent even at the height of the drought when both Ψ_l and soil water potentials (Ψ_s) reached ca. — 4.5 MPa. However, E. pulchella maintained a higher relative water content (RWC) in its leaves than E. coccifera and E. delegatensis, and showed less severe crown damage. After the first light rains substantial interspecific differences in Ψ_l and gs occurred. Eucalyptus pulchella restored normal Ψ_l, gs and RWC more rapidly than the other two species and, even for severely droughted trees, crown growth commenced via epicormic buds near the ends of its branches while for E. delegatensis and E. coccifera crown regeneration was via epicormic buds arising from stems and larger branches. This resulted in a change in dominance in certain stands and showed that E. pulchella was more drought-resistant than E. coccifera, which was in turn more resistant than E. delegatensis. This conclusion was confirmed during competition experiments using potted seedlings. However, potted seedlings differed from mature field trees by maintaining moderate gs at high vapour pressure deficits and closing stomata at Ψ_l below ca. — 2.0 MPa. Substantial variation in the severity of drought symptoms was observed over short distances. This variation appeared to be determined by the moisture-holding capacity of the soil and the biomass of the stand. Although differences in the rooting patterns of seedlings were evident, field measurements of Ψ_l and Ψ_s suggested that all three species were exploiting the same water resource. In contrast to previous studies, the results suggest that large interspecific differences in tissue hydration and crown damage may be present, even though differences in Ψ_l, gs and characteristics of the root system may appear small.     

Limitations to photosynthesis of lettuce grown under tropical conditions: alleviation by root-zone cooling.

Aerial parts of lettuce plants were grown under natural tropical fluctuating ambient temperatures, but with their roots exposed to two different root-zone temperatures (RZTs): a constant 20 degrees C-RZT and a fluctuating ambient (A-) RZT from 23-40 degrees C. Plants grown at A-RZT showed lower photosynthetic CO2 assimilation (A), stomatal conductance (gs), midday leaf relative water content (RWC), and chlorophyll fluorescence ratio Fv/Fm than 20 degrees C-RZT plants on both sunny and cloudy days. Substantial midday depression of A and g(s) occurred on both sunny and cloudy days in both RZT treatments, although Fv/Fm did not vary diurnally on cloudy days. Reciprocal temperature transfer experiments investigated the occurrence and possible causes of stomatal and non-stomatal limitations of photosynthesis. For both temperature transfers, light-saturated stomatal conductance (gs sat) and photosynthetic CO2 assimilation (A(sat)) were highly correlated with each other and with midday RWC, suggesting that A was limited by water stress-mediated stomatal closure. However, prolonged growth at A-RZT reduced light- and CO2-saturated photosynthetic O2 evolution (Pmax), indicating non-stomatal limitation of photosynthesis. Tight temporal coupling of leaf nitrogen content and P(max) during both temperature transfers suggested that decreased nutrient status caused this non-stomatal limitation of photosynthesis.     

水分胁迫下烯效唑对百日草幼苗光合特性及叶解剖结构的影响

以百日草‘芳菲1号’为试材,研究不同水分胁迫下烯效唑(S3307)对其幼苗生长、光合特性及叶解剖结构的影响,以明确S3307对百日草的抗旱作用及其机理。结果显示:(1)在水分胁迫下,百日草的生长均受到不同程度的抑制,叶绿素含量显著降低,光合作用受到抑制,叶解剖结构有所变化。(2)S3307处理后,均能够显著降低所对应的不同程度水分胁迫下百日草的株高,显著增加茎粗、叶面积、叶片厚度、栅栏组织厚度和根冠比,显著增加叶绿素含量,提高百日草的净光合速率(Pn)、气孔导度(Gs)、蒸腾速率(Tr)和水分利用效率(WUE)。研究表明,S3307能够提高百日草的抗旱性,而且在轻度和中度水分胁迫下Pn的下降主要是由气孔因素引起,而在重度水分胁迫下光合速率的下降是由非气孔因素引起的。     

Photosynthetic Characteristics in Mulberry during Water Stress and Rewatering

Three-month-old plants of mulberry (Morus alba L. cv. Kanva-2) were subjected to a drought stress by withholding water supply. As the leaf water potential (W) was dropping progressively with the severity of treatment and increasing stress duration, the values of leaf area, dry mass accumulation, total chlorophyll (Chl) content, net photosynthetic rate (PN), stomatal conductance (gs), and transpiration rate (E) were declined. The photosystem 2 (PS2) photochemical efficiency significantly decreased only at a severe stress treatment. The intercellular CO2 concentration (Ci) remained unaltered during a mild stress, yet it increased under moderate and severe stresses. The Ci/gs ratio reflected the mesophyll efficiency during water stress. Rewatering of the plants led to an almost complete recovery of PN, E, and gs, indicating that a short-term stress brings about reversible effects only.