Photosynthetic limitations and volatile and non‐volatile isoprenoids in the poikilochlorophyllous resurrection plant Xerophyta humilis during dehydration and rehydration

We investigated the photosynthetic limitations occurring during dehydration and rehydration of Xerophyta humilis, a poikilochlorophyllous resurrection plant, and whether volatile and non‐volatile isoprenoids might be involved in desiccation tolerance. Photosynthesis declined rapidly after dehydration below 85% relative water content (RWC). Raising intercellular CO₂ concentrations during desiccation suggest that the main photosynthetic limitation was photochemical, affecting energy‐dependent RuBP regeneration. Imaging fluorescence confirmed that both the number of photosystem II (PSII) functional reaction centres and their efficiency were impaired under progressive dehydration, and revealed the occurrence of heterogeneous photosynthesis during desiccation, being the basal leaf area more resistant to the stress. Full recovery in photosynthetic parameters occurred on rehydration, confirming that photosynthetic limitations were fully reversible and that no permanent damage occurred. During desiccation, zeaxanthin and lutein increased only when photosynthesis had ceased, implying that these isoprenoids do not directly scavenge reactive oxygen species, but rather protect photosynthetic membranes from damage and consequent denaturation. X. humilis was found to emit isoprene, a volatile isoprenoid that acts as a membrane strengthener in plants. Isoprene emission was stimulated by drought and peaked at 80% RWC. We surmise that isoprene and non‐volatile isoprenoids cooperate in reducing membrane damage in X. humilis, isoprene being effective when desiccation is moderate while non‐volatile isoprenoids operate when water deficit is more extreme.     

Flavescence dorée phytoplasma deregulates stomatal control of photosynthesis in Vitis vinifera

Flavescence dorée (FD) is among the major grapevine diseases causing high management costs; curative methods against FD are unavailable. In FD‐infected plants, decrease in photosynthesis is usually recorded, but deregulation in stomatal control of leaf gas exchange during FD infection and recovery is unknown. We measured the seasonal time course of gas exchange rates in two cultivars (‘Barbera’ and ‘Nebbiolo’) during the term of 1 year when grapevines experienced a water stress and another with no drought, with difference in gas exchange rates in response to FD infection and recovery as assessed by symptom observation and phytoplasma detection through PCR analysis. Chlorophyll fluorescence was also evaluated at the time of maximum symptom severity in ‘Barbera’, the cultivar showing the most severe stress response to FD infection, causing the highest damage in vineyards of north‐western Italy. In FD‐infected plants, net photosynthesis and transpiration gradually decreased during the season, more during the no drought year than during drought. During recovery, healthy (PCR negative) plants infected 2 years before, but not those infected an year before, regained the gas exchange performances to the level as measured before infection. The relationships between stomatal conductance and the residual leaf intercellular CO₂ concentration (c_i) discriminated healthy versus FD‐infected and recovered plants; at the same c_i, FD‐infected leaves had higher non‐photochemical quenching than healthy ones. We conclude that metabolic, not stomatal, leaf gas exchange limitation in FD‐infected and recovered grapevines is the basis of plant response to FD disease. In addition, we also suggest that such response is dependent upon water stress, by showing that water stress superimposes on FD infection in terms of stomatal and metabolic non‐stomatal limitations to carbon assimilation.     

Photosynthetic limitations in olive cultivars with different sensitivity to salt stress

Olive (Olea europea L) is one of the most valuable and widespread fruit trees in the Mediterranean area. To breed olive for resistance to salinity, an environmental constraint typical of the Mediterranean, is an important goal. The photosynthetic limitations associated with salt stress caused by irrigation with saline (200 mm ) water were assessed with simultaneous gas‐exchange and fluorescence field measurements in six olive cultivars. Cultivars were found to possess inherently different photosynthesis when non‐stressed. When exposed to salt stress, cultivars with inherently high photosynthesis showed the highest photosynthetic reductions. There was no relationship between salt accumulation and photosynthesis reduction in either young or old leaves. Thus photosynthetic sensitivity to salt did not depend on salt exclusion or compartmentalization in the old leaves of the olive cultivars investigated. Salt reduced the photochemical efficiency, but this reduction was also not associated with photosynthesis reduction. Salt caused a reduction of stomatal and mesophyll conductance, especially in cultivars with inherently high photosynthesis. Mesophyll conductance was generally strongly associated with photosynthesis, but not in salt‐stressed leaves with a mesophyll conductance higher than 50 mmol m^?2 s^?1. The combined reduction of stomatal and mesophyll conductances in salt‐stressed leaves increased the CO₂ draw‐down between ambient air and the chloroplasts. The CO₂ draw‐down was strongly associated with photosynthesis reduction of salt‐stressed leaves but also with the variable photosynthesis of controls. The relationship between photosynthesis and CO₂ draw‐down remained unchanged in most of the cultivars, suggesting no or small changes in Rubisco activity of salt‐stressed leaves. The present results indicate that the low chloroplast CO₂ concentration set by both low stomatal and mesophyll conductances were the main limitations of photosynthesis in salt‐stressed olive as well as in cultivars with inherently low photosynthesis. It is consequently suggested that, independently of the apparent sensitivity of photosynthesis to salt, this effect may be relieved if conductances to CO₂ diffusion are restored.     

Photosynthesis in the water-stressed C4 grass Setaria sphacelata is mainly limited by stomata with both rapidly and slowly imposed water deficits

A comparison of the effects of a rapid and a slowly imposed water deficit on photosynthesis was performed in Setaria sphacelata var. splendida (Stapf) Clayton, a C₄ NADP‐ME grass. Gas exchange was measured in rapidly and slowly dehydrated adult leaves either under atmospheric CO₂ partial pressure with an infrared gas analyser or under saturating CO₂ partial pressure with a leaf disc oxygen electrode. These measurements were used to calculate stomatal and non‐stomatal limitations to photosynthesis. These were further investigated using modulated chlorophyll a fluorescence measurements and photosynthetic pigment quantification. The decrease of net photosynthesis, leaf conductance and water use efficiency was more pronounced under rapid stress than in slow stress. However, photosynthesis is always mainly limited by stomata in both types of stress, albeit the contribution of non‐stomatal limitations increases at severe water deficits in slow stress experiments. The substomatal CO₂ partial pressure significantly increased in both types of stress, suggesting an increased resistance due to an internal barrier to CO₂ diffusion. Physical alterations in the structure of the intercellular spaces due to leaf shrinkage may account for these results. The maximal photochemical efficiency of photosystem II (PSII) was remarkably resistant to stress, as the F_v/F_m ratio decreased only at severe water deficit. On the contrary, the effective photochemical efficiency of PSII (ΔF/F′_m) measured under high actinic light decreased linearly in both types of stress, although in a more pronounced way under rapid stress. A similar variation in photochemical quenching suggests that the decrease of ΔF/F′_m is mainly due to the closure of PSII reaction centres. The non‐photochemical quenching did not change significantly except under severe dehydration indicating that the energization state of thylakoids remained stable under stress. The decrease observed in photosynthetic pigments may be an adaptation to stress rather than a limiting factor to photosynthesis. Results suggests that, although intrinsic mesophyll metabolic inhibitions occur, stomatal limitation to CO₂ diffusion is the main reason for the decrease in photosynthesis.     

Diffusion limitations and metabolic factors associated with inhibition and recovery of photosynthesis from drought stress in a C3 perennial grass species

Stomatal closure and metabolic impairment under drought stress limits photosynthesis. The objective of this study was to determine major stomatal and metabolic factors involved in photosynthetic responses to drought and recovery upon re‐watering in a C₃ perennial grass species, Kentucky bluegrass (Poa pratensis L.). Two genotypes differing in drought resistance, ‘Midnight’ (tolerant) and ‘Brilliant’ (sensitive), were subjected to drought stress for 15 days and then re‐watered for 10 days in growth chambers. Single‐leaf net photosynthetic rate (A), stomatal conductance (g_s) and transpiration rate (Tr) decreased during drought, with a less rapid decline in ‘Midnight’ than in ‘Brilliant’. Photochemical efficiency, Rubisco activity and activation state declined during drought, but were significantly higher in ‘Midnight’ than in ‘Brilliant’. The relationship between A and internal leaf CO₂ concentration (A/Ci curve) during drought and re‐watering was analyzed to estimate the relative influence of stomatal and non‐stomatal components on photosynthesis. Stomatal limitation (Ls %), non‐stomatal limitation (Lns %), CO₂ compensation point (CP) and dark respiration (Rd) increased with stress duration in both genotypes, but to a lesser extent in ‘Midnight’. Maximum CO₂ assimilation rate (A_max), carboxylation efficiency (CE) and mesophyll conductance (g_m) declined, but ‘Midnight’ had significantly higher levels of A_max, CE and g_m than ‘Brilliant’. Maximum carboxylation rate of Rubisco (V_cmax) and ribulose‐1,5‐bisphospate (RuBP) regeneration capacity mediated by maximum electron transport rate (J_max) decreased from moderate to severe drought stress in both genotypes, but to a greater extent in ‘Brilliant’ than in ‘Midnight’. After re‐watering, RWC restored to about 90% of the control levels in both genotypes, whereas A, g_s, Tr and Fv/Fm was only partially recovered, with a higher recovery level in ‘Midnight’ than in ‘Brilliant’. Rubisco activity and activation state restored to the control level after re‐watering, with more rapid increase in ‘Midnight’ than in ‘Brilliant’. The values of Ls, Lns, CP and Rd declined, and A_max, CE, V_cmax, J_max and g_m increased after re‐watering, with more rapid change in all parameters in ‘Midnight’ than in ‘Brilliant’. These results indicated that the maintenance of higher A and A_max under drought stress in drought‐tolerant Kentucky bluegrass could be attributed to higher Rubico activation state, higher CE and less stomatal limitation. The ability to resume metabolic activity (A_max, CE, Fv/Fm and Rubisco) was observed in the drought‐tolerant genotype and is the most likely cause for the increased recuperative ability of photosynthesis. Incomplete recovery of photosynthesis upon re‐watering could be attributable to lasting stomatal limitations caused by severe drought damage in both genotypes. Promoting rapid stomatal recovery from drought stress may be critical for plants to resume full photosynthetic capacity in C₃ perennial grass species.     

Photosynthesis, non-photochemical pathways and activities of antioxidant enzymes in a resilient evergreen oak under different climatic conditions from a valley-savanna in Southwest China

Plants in the savanna‐valleys in Southwest China are annually exposed to different combinations of multiple stresses from the hot‐rainy, to chill‐dry, and to warm‐dry seasons. This study monitored seasonal changes in photosynthesis and photoprotection in an evergreen oak (Cyclobalanopsis helferiana) from one of these valleys for four years during which usual and abnormal drought occurred. In general, during the study period with decreasing xylem water potential (Ψx), photosynthetic gas exchange, quantum yield of photosystem II (PSII) photochemistry and activities of most of the measured antioxidant enzymes decreased, while activities of the xanthophyll cycle and associated non‐photochemical energy dissipation and glutathione peroxidase (GP) (EC 1.11.1.9) increased. In a fairly severe chill period, high concentration of reactive oxygen species induced high activities of most of the antioxidant enzymes and relatively stronger decrease in gas exchange. In the most severe dry period, even when predawn Ψx decreased down to ?4 MPa, considerable Pn (maximum photosynthetic rate) (4 μmol m^–2 s^–1) was still maintained in midmorning. At this time, most of the antioxidant enzyme activities decreased to the lowest values, whereas the xanthophyll cycle and associated non‐photochemical energy dissipation and GP activities increased to their highest levels. High predawn antheraxanthin and zeaxanthin contents were observed in the severe and very severe drought periods. Superoxide dismutase maintained high and fairly constant activity (1500–1800 U mg^?1 protein) and predawn maximum photochemistry efficiency of PSII was always above 0.8 throughout the whole study period. These results indicated that the photosynthetic apparatus of the oak leaves was highly capable of maintaining its function under the multiple stresses in different seasons in the present valley‐savanna.     

Effects of Drought on Photosynthetic Performance and Water Relations of Four Vigna Genotypes

The effect of drought on plant water relations and photosynthesis of Vigna glabrescens (Vg) and Vigna unguiculata (cvs. 1183, EPACE-1 and Lagoa), which differ in their drought resistance, was compared. With the increase of drought severity, Vg showed a more gradual stomatal closure and maintained significantly higher levels of stomatal conductance (g_s) and photosynthetic activity (P_N) than the other genotypes even when minimum relative water content (RWC) values were observed. Furthermore, Vg was the only genotype able to accumulate significant amounts of proline already under moderate water deficit, what could explain the lower osmotic potential (ψ_s) values observed in these plants. The three V. unguiculata cultivars presented a similar stomatal control under increasing water deficit. A mesophyllic impairment of photosynthetic capacity (P_max) was detected for cv. 1183 from the beginning of drought onset (85-75 % RWC) while in the Vg plants the values remained unaffected along the whole drought period, indicating that P_N decrease observed in this genotype is mainly a consequence of stomatal closure. Such P_max maintenance suggests the existence of a high mesophyllic ability to cope with increasing tissue dehydration in Vg. This revised version was published online in September 2006 with corrections to the Cover Date.     

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.     

Effects of Reddening of Cotton (Gossypium hirsutum L.) Leaves on Functional Activity of Photosynthetic Apparatus

Strong inhibition of rates of CO₂ assimilation and transpiration, stomatal conductance, and water use efficiency as well as photosystem 2 (PS2) photochemical activity were related to the severity of reddening. The inhibition of photosynthesis in red cotton leaves was due to both decreased photochemical activity and stomatal limitation. Lowered photosynthetic capacity could be one of the main factors of reduced yield in reddening cotton.     

Effect of arbuscular mycorrhizal inoculation on water status and photosynthesis of Populus cathayana males and females under water stress

Drought is one of the most serious environmental limitations for poplar growth. Although the ways in which plants deal with water stress and the effects of arbuscular mycorrhizal (AM) formation have been well documented, little is known about how the male and female plants of Populus cathayana respond to drought and AM formation. We also aimed to investigate the potential role of AM fungi in maintaining gender balance. We tested the impact of drought and AM formation on water status and photosynthesis. The results suggested that both sexes showed similar responses to water stress: drought decreased the growth of stem length (GSL), growth of ground diameter (GGD), relative water content (RWC), increased the relative electrolyte leakage (REL), and limited the photosynthesis and chlorophyll fluorescence indexes. However, the responses of the two sexes to drought and AM formation differed to some extent. AM formation had positive effects on RWC, photosynthesis and the intrinsic water use efficiency (WUEi) but negative effects on the REL of males and females, especially under drought. AM formation enhanced the maximum quantum yield of photosystem II (PSII) (Fv/Fm), the actual quantum yield of PSII (ΦPSII), non‐photochemical quenching (qN) and photochemical quenching (qP) under drought conditions, and had no significant effects under well‐watered conditions except on the qP of males. Principal component analysis showed that males were significantly more drought tolerant than females, and AM formation enhanced drought tolerance, particularly among males, which suggested that AM fungi are beneficial for ecological stability and for P. cathayana survival under drought conditions.     

Chemical and hydraulic signals regulate stomatal behavior and photosynthetic activity in maize during progressive drought

The objectives of this study were to investigate stomatal regulation in maize seedlings during progressive soil drying and to determine the impact of stomatal movement on photosynthetic activity. In well-watered and drought-stressed plants, leaf water potential (Ψ _leaf), relative water content (RWC), stomatal conductance (g _s), photosynthesis, chlorophyll fluorescence, leaf instantaneous water use efficiency (iWUE_leaf), and abscisic acid (ABA) and zeatin-riboside (ZR) accumulation were measured. Results showed that g _s decreased significantly with progressive drought and stomatal limitations were responsible for inhibiting photosynthesis in the initial stages of short-term drought. However, after 5 days of withholding water, non-stomatal limitations, such as damage to the PSII reaction center, became the main limiting factor. Stomatal behavior was correlated with changes in both hydraulic and chemical signals; however, changes in ABA and ZR occurred prior to any change in leaf water status. ABA in leaf and root tissue increased progressively during soil drying, and further analysis found that leaf ABA was negatively correlated with g _s (R ² = 0.907, p < 0.05). In contrast, leaf and root ZR decreased gradually. ZR in leaf tissue was positively correlated with g _s (R ² = 0.859, p < 0.05). These results indicate that ABA could induce stomatal closure, and ZR works antagonistically against ABA in stomatal behavior. In addition, the ABA/ZR ratio also had a strong correlation with g _s, suggesting that the combined chemical signal (the interaction between ABA and cytokinin) plays a role in coordinating stomatal behavior. In addition, Ψ _leaf and RWC decreased significantly after only 3 days of drought stress, also affecting stomatal behavior.     

Photosynthetic Responses of a Temperate Liana to Xylella fastidiosa Infection and Water Stress

Xylella fastidiosa is a xylem‐limited bacterial plant pathogen that causes bacterial leaf scorch in its hosts. Our previous work showed that water stress enhances leaf scorch symptom severity and progression along the stem of a liana, Parthenocissus quinquefolia, infected by X. fastidiosa. This paper explores the photosynthetic gas exchange responses of P. quinquefolia, with the aim to elucidate mechanisms behind disease expression and its interaction with water stress. We used a 2 × 2‐complete factorial design, repeated over two growing seasons, with high and low soil moisture levels and infected and non‐infected plants. In both years, low soil moisture levels reduced leaf water potentials, net photosynthesis and stomatal conductance at all leaf positions, while X. fastidiosa‐infection reduced these parameters at basally located leaves only. Intercellular CO₂ concentrations were reduced in apical leaves, but increased at the most basal leaf location, implicating a non‐stomatal reduction of photosynthesis in leaves showing the greatest disease development. This result was supported by measured reductions in photosynthetic rates of basal leaves at high CO₂ concentrations, where stomatal limitation was eliminated. Repeated measurements over the summer of 2000 showed that the effects of water stress and infection were progressive over time, reaching their greatest extent in September. By reducing stomatal conductances at moderate levels of water stress, P. quinquefolia maintained relatively high leaf water potentials and delayed the onset of photosynthetic damage due to pathogen and drought‐induced water stress. In addition, chlorophyll fluorescence measurements showed that P. quinquefolia has an efficient means of dissipating excess light energy that protects the photosynthetic machinery of leaves from irreversible photoinhibitory damage that may occur during stress‐induced stomatal limitation of photosynthesis. However, severe stress induced by disease and drought eventually led to non‐stomatal decreases in photosynthesis associated with leaf senescence.     

Physiological and anatomical changes induced by drought in two olive cultivars (cv Zalmati and Chemlali)

Photosynthetic gas exchange, vegetative growth, water relations and fluorescence parameters as well as leaf anatomical characteristics were investigated on young plants of two Olea europaea L. cultivars (Chemlali and Zalmati), submitted to contrasting water availability regimes. Two-year-old olive trees, grown in pots in greenhouse, were not watered for 2 months. Relative growth rate (RGR), leaf water potential (Ψ_LW) and the leaf relative water content (LWC) of the two cultivars decreased with increasing water stress. Zalmati showed higher values of RGR and LWC and lower decreased values of Ψ_LW than Chemlali, in response to water deficit, particularly during severe drought stress. Water stress also caused a marked decline on photosynthetic capacity and chlorophyll fluorescence. The net photosynthetic rate, stomatal conductance, transpiration rate, the maximal photochemical efficiency of PSII (F _v/F _m) and the intrinsic efficiency of open PSII reaction centres (F′ _v/F′ _m) decreased as drought stress developed. In addition, drought conditions, reduced leaf chlorophyll and carotenoids contents especially at severe water stress. However, Zalmati plants were the less affected when compared with Chemlali. In both cultivars, stomatal control was the major factor affecting photosynthesis under moderate drought stress. At severe drought-stress levels, the non-stomatal component of photosynthesis is inhibited and inactivation of the photosystem II occurs. Leaf anatomical parameters show that drought stress resulted in an increase of the upper epidermis and palisade mesophyll thickness as well as an increase of the stomata and trichomes density. These changes were more characteristic in cv. ‘Zalmati’. Zalmati leaves also revealed lower specific leaf area and had higher density of foliar tissue. From the behaviour of Zalmati plants, with a smaller reduction in relative growth rate, net assimilation rate and chlorophyll fluorescence parameters, and with a thicker palisade parenchyma, and a higher stomatal and trichome density, we consider this cultivar more drought-tolerant than cv. Chemlali and therefore, very promising for cultivation in arid areas.     

Drought-inhibition of photosynthesis in C3 plants: stomatal and non-stomatal limitations revisited

There is a long-standing controversy as to whether drought limits photosynthetic CO2 assimilation through stomatal closure or by metabolic impairment in C3 plants. Comparing results from different studies is difficult due to interspecific differences in the response of photosynthesis to leaf water potential and/or relative water content (RWC), the most commonly used parameters to assess the severity of drought. Therefore, we have used stomatal conductance (g) as a basis for comparison of metabolic processes in different studies. The logic is that, as there is a strong link between g and photosynthesis (perhaps co-regulation between them), so different relationships between RWC or water potential and photosynthetic rate and changes in metabolism in different species and studies may be 'normalized' by relating them to g. Re-analysing data from the literature using light-saturated g as a parameter indicative of water deficits in plants shows that there is good correspondence between the onset of drought-induced inhibition of different photosynthetic sub-processes and g. Contents of ribulose bisphosphate (RuBP) and adenosine triphosphate (ATP) decrease early in drought development, at still relatively high g (higher than 150 mmol H20 m(-2) s(-1)). This suggests that RuBP regeneration and ATP synthesis are impaired. Decreased photochemistry and Rubisco activity typically occur at lower g (<100 mmol H20 m(-2) s(-1)), whereas permanent photoinhibition is only occasional, occurring at very low g (<50 mmol H20 m(-2) s(-1)). Sub-stomatal CO2 concentration decreases as g becomes smaller, but increases again at small g. The analysis suggests that stomatal closure is the earliest response to drought and the dominant limitation to photosynthesis at mild to moderate drought. However, in parallel, progressive down-regulation or inhibition of metabolic processes leads to decreased RuBP content, which becomes the dominant limitation at severe drought, and thereby inhibits photosynthetic CO2 assimilation.     

Lower leaf gas‐exchange and higher photorespiration of treated wastewater irrigated Citrus trees is modulated by soil type and climate

Water quality, soil and climate can interact to limit photosynthesis and to increase photooxidative damage in sensitive plants. This research compared diffusive and non‐diffusive limitations to photosynthesis as well as photorespiration of leaves of grapefruit trees in heavy clay and sandy soils having a previous history of treated wastewater (TWW) irrigation for >10 years, with different water qualities [fresh water (FW) vs TWW and sodium amended treated wastewater (TWW + Na)] in two arid climates (summer vs winter) and in orchard and lysimeter experiments. TWW irrigation increased salts (Na⁺ and Cl⁻), membrane leakage, proline and soluble sugar content, and decreased osmotic potentials in leaves of all experiments. Reduced leaf growth and higher stomatal and non‐stomatal (i.e. mesophyll) limitations were found in summer and on clay soil for TWW and TWW + Na treatments in comparison to winter, sandy soil and FW irrigation, respectively. Stomatal closure, lower chlorophyll content and altered Rubisco activity are probable causes of higher limitations. On the other hand, non‐photochemical quenching, an alternative energy dissipation pathway, was only influenced by water quality, independent of soil type and season. Furthermore, light and CO₂ response curves were investigated for other possible causes of higher non‐stomatal limitation. A higher proportion of non‐cyclic electrons were directed to the O₂ dependent pathway, and a higher proportion of electrons were diverted to photorespiration in summer than in winter. In conclusion, both diffusive and non‐diffusive limitations contribute to the lower photosynthetic performance of leaves following TWW irrigation, and the response depends on soil type and environmental factors.     

24-epibrassionlide improves photosynthetic response of Rhododendron delavayi to drought

Rhododendron delavayi is an alpine evergreen ornamental plant with strong tolerance to drought stress. Brassinosteroids are promising agents for alleviating the negative effects of drought on plants, but the mechanism by which BRs induce plant resistance to drought is not well understood. The present study investigated the effects of exogenous spray of 24-epibrassionlide (EBR) at different concentrations (0~1 mg l⁻¹) on the physiological response of R. delavayi to drought caused by no watering for 10 days. With the increase in EBR concentration, net photosynthetic rate, stomatal conductance, transportation rate, light saturated photosynthetic rate, light compensation point, light saturation point, excitation energy capture efficiency of reaction center, actual photochemical efficiency of photosystem II (PSII), photochemical quenching and electron transport rate significantly increased, but there were no significant effects on photosynthetic pigment content. These results suggested that the EBR-induced improvement in CO₂ assimilation under drought was mainly related to stomatal and non-stomatal factors, and partially attributed to the increased photochemical efficiency of PSII. In addition, the leaf water potential increased with the increase in EBR concentration, while the malondialdehyde, superoxide dismutase, catalase, proline and soluble protein decreased. The results suggested EBR application partially alleviated the negative effect of drought on R. delavayi by improving water relations and decreasing lipid peroxidation and reactive oxygen species production. We concluded that exogenous application of EBR improved photosynthesis and alleviated the negative effects of drought-induced membrane peroxidation and severe oxidative stress.     

Gas exchange and leaf water relations in two peanut cultivars of different drought tolerance

In summer seasons of 1991 and 1992 the gas exchange and leaf water relations were analysed in two peanut cultivars: drought tolerant cv. GG 2 (DT) and drought sensitive cv. JL 24 (DS). Soil moisture stress was imposed by withholding irrigation at pod development phase. The decrease in photosynthesis (P_N) under stress was associated with a decrease in stomatal conductance (g_s) and relative water content (RWC). The P_N and RWC were significantly higher under stress in DT than DS. On relief of stress the g_s and RWC recovered more quickly in DT than DS. The maintenance of higher RWC (>80 %), g_s and P_N under stress appears to be imparting drought tolerance in peanut.     

The apparent feed-forward response to vapour pressure deficit of stomata in droughted, field-grown Eucalyptus globulus Labill

This study tested a multiplicative model of stomatal response to environment for drought‐affected trees of Eucalyptus globulus Labill. growing in southern Australia. The model incorporates a feed‐forward response to vapour pressure deficit of ambient air (δe_a) and performed well if evaluated using reduced major axis regression and log‐transformed data. There was strong evidence from gas‐exchange data, leaf water potentials and sapflow measurements of the feed‐forward response by stomata to leaf‐to‐air vapour pressure deficit (δe_l). The response of stomata to δe_l was irreversible. Stomatal conductance and the rate of net photosynthesis were highly correlated and declined, together with the rate of transpiration, throughout the afternoon as δe_a increased despite increasing leaf water potentials. The concentration of CO₂ inside leaves (c_i) increased as stomatal conductance declined indicating increasing non‐stomatal limitations to photosynthesis. The stomatal response to δe_l of E. globulus in the field is best described as an ‘apparent feed‐forward response’ that probably results from both slowly reversible depression of net photosynthesis and abscisic acid accumulation in guard cells. We suggest that the stomatal response to c_i may strengthen the link between photosynthetic capacity and stomatal conductance during leaf drying as a result of either drought or large δ e_l.     

Photosynthesis by six Portuguese maize cultivars during drought stress and recovery

Photosynthesis, chlorophyll fluorescence, and leaf water parameters were measured in six Portuguese maize (Zea mays L.) cultivars during and following a period of drought stress. The leaf relative water content (RWC) responded differently among cultivars but except for cultivar PB369, recovered close to initial values after watering was restored. Photosynthetic rate and stomatal conductance decreased with drought but more slowly in cultivars PB269 and PB260 than in cultivars AD3R, PB64, PB304 and PB369. Water use efficiency (WUE) decreased during the water stress treatment although with cultivar PB260 the decrease was marked only when the RWC fell below 40%. Recovery of WUE was seen with all cultivars except PB369. The maximum quantum efficiency of photosystem II, the photochemical quenching coefficient, the electron transport rate in PSII and the estimated functional plastoquinone pool tended to decrease with drought, while the non-photochemical quenching coefficient increased. The parameters estimated from chlorophyll fluorescence did not recover in PB369, during re-watering. The results show that PB260 and PB269 were the most tolerant and PB369 was the least tolerant cultivars to water stress. The variation found among the cultivars tested suggests the existence of valuable genetic resources for crop improvement in relation to drought tolerance.     

Effect of low boron supply in turnip plants under drought stress

Turnip (Brassica rapa L.) plants were grown in Perlite with low (< 2.5 μM) or adequate (25 μM) boron supply under well-watered and drought conditions for 12 weeks. Dry mass of leaves and roots was reduced under drought by about 61 and 56 % in plants supplied adequately with B, while up to 84 and 74 % under B starvation. Drought reduced B content by about 70 and 82 % for B-sufficient and B-deficient plants, respectively. According to the chlorophyll fluorescence parameters, the photosynthesis processes conserved their normal activities under low B supply in well-watered plants, while a serious damage to photosystem 2 occurred under drought stress. Stomatal limitation was the most important cause for a 17 % lower net photosynthetic rate (P_N) of drought stressed B-sufficient plants. In B-deficient plants, however, both stomatal and non-stomatal limitations were involved in 53 % reduction of P_N. Low B supply reduced strongly leaf water potential.