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 (ψ₁) 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 ψ₁between 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.     

Contribution of Pre-Anthesis Assimilates and Current Photosynthesis to Grain Yield,and their Relationships to Drought Resistance in Wheat Cultivars Grown Under Different Soil Moisture

We investigated the relative importance of pre-anthesis assimilates stored in plant parts, mainly in the stem, and post-anthesis photosynthesis to drought resistance in wheat (Triticum aestivum L.) cultivars Hongwangmai (drought resistant) and Haruhikari (drought sensitive) subjected to two soil moisture regimes: irrigated and non-irrigated. In the irrigated treatment, soil moisture was maintained near field capacity throughout the growing season, while in the non-irrigated treatment water was withheld from 81 d after sowing until maturity. Drought stress reduced grain yield of Hongwangmai and Haruhikari by 41 and 60 %, respectively. Remobilization of pre-anthesis assimilates to the grain (remobilization) was reduced by drought in Hongwangmai but increased in Haruhikari. The contribution of pre-anthesis assimilates to the grain decreased under non-irrigated treatment in Hongwangmai. However, under water stress, Hongwangmai maintained a higher net photosynthetic rate in the flag leaf than Haruhikari. These results indicated that maintenance of post-anthesis photosynthetic rate was related to drought resistance in Hongwangmai rather than to remobilization under drought stress.     

Gas exchange and photosynthesis of Eucalyptus camaldulensis seedlings inoculated with different ectomycorrhizal symbionts

Eucalyptus camaldulensis Dehnh. seedlings inoculated with Pisolithus tinctorius (Pers.) Coker & Couch and Thelephora terrestris Ehrl. per Fr. were grown in well watered soil (_s –0.03 MPa) or subjected to a long-term soil water stress of up to –1.0 MPa over 13-week period in a glasshouse. After 13 weeks, all seedling containers were watered to field capacity and then water was withheld from the E. camaldulensis seedlings to induce a short-term drought. Diurnal measurements of seedling photosynthesis rate (A), leaf stomatal conductance (g) and leaf water potential (_p) were completed before, during, and after the short term drought. Although they were growing in an equal soil volume, photosynthesis rate (A), leaf stomatal conductance and leaf water potential (_p) of larger seedlings with P. tinctorius ectomycorrhizae were similar to those of smaller seedlings colonized with T. terrestris during the short-term drought period. Seedlings inoculated with Pisolithus tinctorius maintained higher photosynthesis rates over the course of the short-term drought. Thus, P. tinctorius ectomycorrhizae appear to be more efficient than those of T. terrestris in assisting seedlings to maintain gas exchange and photosynthesis under limited soil moisture conditions.     

Drought response of a native and introduced Hawaiian grass

The alien grass, Pennisetum setaceum, dominates many of the lowland arid regions that once supported native Heteropogon contortus grassland on the island of Hawaii. Response to drought in a glasshouse was compared between these C₄ grasses to test if success as an invader is related to drought tolerance or plasticity for traits that confer drought tolerance. Pennisetum produced 51% more total biomass, allocated 49% more biomass to leaves, and had higher net photosynthetic rates (P _n) on a leaf area basis than Heteropogon. Plants of both species under drought produced less total biomass and increased their allocation to roots compared to well-watered plants, but there was no difference between the two species in the magnitude of these responses. The decline in P _n with decreasing leaf water potential (₁) was greater for Pennisetum compared to Heteropogon. Plasticity in the response of P _n to ₁, osmotic potentials, and the water potentials at turgor loss in response to drought were not different between the two species. Stomata were more responsive to w in Heteropogon than in Pennisetum and for well-watered plants compared to droughted plants. Plasticity for the stomatal response to w, however, was not different between the species. There was no evidence that the alien, Pennisetum, had greater plasticity for traits related to drought tolerance compared to the native, Heteropogon. Higher P _n and greater biomass allocation to leaves resulted in greater growth for Pennisetum compared to Heteropogon and may explain the success of Pennisetum as an invader of lowland arid zones on Hawaii.     

Effects of Soil Drought and Atmospheric Humidity on Yield,Gas Exchange,and Stable Carbon Isotope Composition of Barley

The combined effects of water status, vapour pressure deficit (VPD), and elevated temperature from heading to maturity were studied in barley. Plants growing at high VPD, either under well-watered or water deficit conditions, had higher grain yield and grain filling rate than plants growing at low VPD. By contrast, water stress decreased grain yield and individual grain dry matter at any VPD. Water regime and to a lesser extent VPD affected ¹³C of plant parts sampled at mid-grain filling and maturity. The differences between treatments were maximal in mature grains, where high VPD increased ¹³C for both water regimes. However, the total amount of water used by the plant during grain filling did not change as response to a higher VPD whereas transpiration efficiency (TE) decreased. The net photosynthetic rate (P _N) of the flag leaves decreased significantly under water stress at both VPD regimes. However, P _N of the ears was higher at high VPD than at low VPD, and did not decrease as response to water stress. The higher correlation of grain yield with P _N of the ear compared with that of the flag leaf support the role of ear as the main photosynthetic organ during grain filling under water deficit and high VPD. The deleterious effects of combined moderately high temperature and drought on yield were attenuated at high VPD.     

Effect of Growth Regulators on Photosynthesis, Transpiration and Related Parameters in Water Stressed Cotton

Gas exchange in Gossypium hirsutum L. cv. H-777 as affected by water deficit and growth regulators (IAA, GA₃, BAP, ABA, ethrel) was examined. Sixty days after sowing, growth regulators in concentration 50 µM were applied as foliar spray and irrigation was withheld to get desired (moderate and severe) water deficit. All the parameters were measured on the third leaf from the top between 10:00 and 11:00. Net photosynthetic rate (P_N), transpiration rate (E), stomatal conductance (g_s), carboxylation efficiency (CE), and water potential (_w) decreased significantly with the increasing water stress, however, water use efficiency (WUE) was unaffected. Foliar spray with IAA, GA₃ and BAP partially counteracted the effect of water deficit on the above parameters except _w, which became more negative. ABA and up to some extent ethrel increased WUE and maintained higher _w, however, caused further decrease in P_N, E, and g_s.     

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.     

Photosynthesis and the Enzymes of Photosynthetic Carbon Reduction Cycle in Mulberry During Water Stress and Recovery

Net photosynthetic rate (P _N), stomatal conductance (g _S), transpiration rate (E), intercellular CO₂ concentration (C _i), leaf water potential (_w), leaf area, chlorophyll (Chl) content, and the activities of photosynthetic carbon reduction cycle (PCR) enzymes in two mulberry (Morus alba L.) cultivars (drought tolerant Anantha and drought sensitive M-5) were studied during water stress and recovery. During water stress, P _N, g _S, and E declined whereas C _i increased. P _N, g _S, and E were less affected in Anantha than in M-5, which indicates tolerance nature of Anantha over M-5. Activities of ribulose-5-phosphate kinase, NAD- and NADP-glyceraldehyde-3-phosphate dehydrogenases, and 3-phosphoglycerate kinase decreased with increasing stress in both the cultivars. The enzyme activities less affected in tolerant (Anantha) than in sensitive cultivar (M-5) were restored after re-watering to almost initial values in both the cultivars. Re-watering of the plants led to an almost complete recovery of P _N, E, and g _S, indicating that a short-term stress brings about reversible effect in these two cultivars of mulberry.     

Comparison of the Drought Stress Responses of Tolerant and Sensitive Wheat Cultivars During Grain Filling: Changes in Flag Leaf Photosynthetic Activity,ABA Levels,and Grain Yield

Water status parameters, flag leaf photosynthetic activity, abscisic acid (ABA) levels, grain yield, and storage protein contents were investigated in two drought-tolerant (Triticum aestivum L. cv. MV Emese and cv. Plainsman V) and two drought-sensitive (cvs. GK élet and Cappelle Desprez) wheat genotypes subjected to soil water deficit during grain filling to characterize physiological traits related to yield. The leaf water potential decreased earlier and at a higher rate in the sensitive than in the tolerant cultivars. The net CO₂ assimilation rate (P _N) in flag leaves during water deficit did not display a strict correlation with the drought sensitivity of the genotypes. The photosynthetic activity terminated earliest in the tolerant cv. Emese, and the senescence of flag leaves lasted 7 days longer in the sensitive Cappelle Desprez. Soil drought did not induce characteristic differences between sensitive and tolerant cultivars in chlorophyll a fluorescence parameters of flag leaves during post-anthesis. Changes in the effective quantum yield of PSII (Φ_PSII) and the photochemical quenching (qP) depended on the genotypes and not on the sensitivity of cultivars. In contrast, the levels of ABA in the kernels displayed typical fluctuations in the tolerant and in the sensitive cultivars. Tolerant genotypes exhibited an early maximum in the grain ABA content during drought and the sensitive cultivars maintained high ABA levels in the later stages of grain filling. In contrast with other genotypes, the grain number per ear did not decrease in Plainsman and the gliadin/glutenin ratio was higher than in the control in Emese during drought stress. A possible causal relationship between high ABA levels in the kernels during late stages of grain filling and a decreased grain yield was found in the sensitive cultivars during drought stress.     

Photosynthesis and yield responses of ozone-polluted winter wheat to drought

Winter wheat (Triticum aestivum L. cv. Jingdong 8) was exposed to short-term high ozone treatment after anthesis and then was either well irrigated with soil water content (SWC) of 80–85 % (O₃+W) or drought treated (SWC 35–40 %, O₃+D). Short-term ozone exposure significantly decreased irradiance-saturated net photosynthetic rate (P _N) of winter wheat. Under good SWC, P _N of the O₃-treated plant was similar to that of control on 2 d after O₃-exposure (6 DAA), but decreased significantly after 13 DAA, indicating that O₃ exposure accelerated leaf senescence. Meanwhile, green flag leaf area was reduced faster than that of control. As a result, grain yield of O₃+W was significantly decreased. P _N of O₃+D was further notably decreased and green flag leaf area was reduced more than that in O₃+W. Consequently, substantial yield loss of O₃+D was observed compared to that of O₃+W. Although P _N was significantly positively correlated with stomatal conductance, it also had notable positive correlation with the maximum photochemical efficiency in the dark adapted leaves (F_v/F_m), electron transport rate (ETR), photochemical quenching (q_P), as well as content of chlorophyll, suggesting that the depression of P _N was mainly caused by non-stomatal limitation. Hence optimal soil water condition should be considered in order to reduce the yield loss caused by O₃ pollution.     

Effects of water deficit on N2(C2H2) fixation in cowpea and groundnut

The effect of water deficit on nodulation, N₂ fixation, photosynthesis, and total soluble sugars and leghemoglobin in nodules was investigated in cowpea and groundnut. Nitrogenase activity completely ceased in cowpea with a decrease in leaf water potential ( _leaf) from –0.4 MPa to –0.9 MPa, while in groundnut it continued down to –1.7 MPa. With increasing water stress, the acetylene reduction activity (ARA) declined very sharply in cowpea, but ARA gradually decreased in groundnut. Even with mild water stress ( _leaf of 0.2 MPa), nodule fresh weight declined 50% in cowpea partly due to a severe nodule shedding whereas nodule fresh weight declined in groundnut only when _leaf decreased by 1.0 MPa. No nodule shedding was noticed even at a higher stress level in groundnut. Photosynthesis and stomatal conductance were also more stable in groundnut than in cowpea under water stress. There was a sharp increase in total soluble sugars and leghemoglobin in the nodules of groundut with water stress, but no definite trend could be found in cowpea.     

Photosynthesis and photoinhibition in two xerophytic shrubs during drought

Seasonal changes in water relations, net photosynthetic rate (P _N), and fluorescence of chlorophyll (Chl) a of two perennial C₃ deciduous shrubs, Ipomoea carnea and Jatropha gossypifolia, growing in a thorn scrub in Venezuela were studied in order to establish the possible occurrence of photoinhibition during dry season and determine whether changes in photochemical activity of photosystem 2 (PS2) may explain variations of P _N in these species. Leaf water potential () decreased from –0.2 to –2.1 MPa during drought in both species. The P _N decreased with in I. carnea and J. gossypifolia by 64 and 74 %, respectively. Carboxylation efficiency (CE) decreased by more than 50 and 70 % in I. carnea and J. gossypifolia, respectively. In I. carnea, relative stomatal limitation (L_s) increased by 17 % and mesophyll limitation (L_m) by 65 % during drought, while in J. gossypifolia L_s decreased by 27 % and L_m increased by 51 %. Drought caused a reduction in quantum yield of PS2 (_PS2) in both species. Drought affected the capacity of energy dissipation of leaves, judging from the changes in the photochemical (q_P) and non-photochemical quenching (NPQ) coefficients. Photoinhibition during drought in I. carnea and J. gossypifolia was evidenced in the field by a drop in the maximum quantum yield of PS2 (F_v/F_m) below 0.8 and also by non-coordinated changes in _PS2 and quantum yield of non-photochemical excitation quenching (Y_n). Total soluble protein content on an area basis increased with but the ribulose-1,5-bisphosphate carboxylase/oxygenase content remained unchanged. A reduction of total Chl content with drought was observed. Hence in the species studied photoinhibition occurred, which imposed an important limitation on carbon assimilation during drought.     

Photosynthetic Characteristics of Non-Leaf Organs of Winter Wheat Cultivars Differing in Ear Type and their Relationship with Grain Mass Per Ear

Chlorophyll content, photosystem 2 functioning (F_v/F_m, F_v/F₀), activity of ribulose-1,5-bisphosphate carboxylase/oxygenase, and net photosynthetic rates (P _N) of flag leaf blade, sheath, peduncle, and ear organs were assessed in large-ear type (Pin 7) and small-ear type (ND93) wheat cultivars. Some differences were found in photosynthetic properties between different green plant parts, the values of all studied parameters in ear parts being higher in Pin7 than in ND93. Furthermore, ear surface areas and ear P _N in 26 wheat genotypes measured at anthesis showed highly significant positive correlation with grain mass per ear. Hence a greater capability of ear photosynthesis may result in a greater grain yield in large-ear type cultivars.     

Growth and osmotic adjustment of two tomato cultivars during and after saline stress

The effect of a short period of saline stress was studied in two phenotypically different cultivars, one of normal fruit-size (L. esculentum cv. New Yorker) and one of cherry fruit-size (L. esculentum var.cerasiforme cv. PE-62). In both cultivars the relative growth rate (RGR) and the leaf area ratio (LAR) decreased following salinisation. The leaf turgor potential (_p) and the osmotic potential at full turgor (_os) decreased to the same extent in both cultivars. However, the contributions of organic and inorganic solutes to the osmotic adjustment was different between cultivars. New Yorker achieved the osmotic adjustment by means of the Cl^– and Na⁺ uptake from the substrate, and by synthesis of organic solutes. In the cherry cultivar organic solutes did not contribute to the osmotic adjustment, instead, their contribution decreased after salinisation. After the salt stress was removed, the water stress disappeared, the content of organic solutes decreased in plants of both cultivars and, therefore, their growth was not retarded by the diversion of resources for the synthesis of organic solutes. However, the toxic effects of the Cl^– and Na⁺ did not disappear after removal of the salt stress, and the net assimilation rate (NAR) and the rate of growth (RGR) did not recover.     

Effects of water stress on the gas exchange,the activities of some enzymes of carbon and nitrogen metabolism,and on the pool sizes of some organic acids in maize leaves

Water-stressed maize (Zea mays L.) leaves showed a large decrease in leaf conductance during photosynthesis. Net CO₂ uptake and evaporation declined fast at mild stress (=–0.6 to –1.0 MPa) and slower at more severe stress (=–1.0 to -1.2 MPa), whereas the CO₂ concentration in the intercellular spaces (C_i) did not drop to the CO₂ compensation point. The activities of the enzymes of photosynthetic carbon metabolism tested in this study dropped by approx. 30% at =-1.2 MPa. Glutamine synthetase activity was unaffected by water stress, whereas the activity of nitrate reductase was almost completely inhibited. The decline of enzyme activities in relation to was correlated with a concomitant decrease in the content of total soluble protein of the stressed leaves. The total leaf pools of malate, pyruvate and oxaloacetate decreased almost linearly in relation to , thus obviously contradicting the almost constant C_i. In comparison to the controls (=0.6 MPa) the content of citrate and isocitrate increaed markedly at =-0.9 MPa and decreased again at =-1.2 MPa.Abbreviations PCR photosynthetic carbon reduction cycle - PCO photosynthetic carbon oxidation cycle - PEP phosphoenolypyruvate - RuBP ribulose-1,5-bisphosphate     

Adaptive photosynthetic strategies of the Mediterranean maquis species according to their origin

In consideration of their origin the adaptive strategies of the evergreen species of the Mediterranean maquis were analysed. Rosmarinus officinalis L., Erica arborea L., and Erica multiflora L. had the lowest net photosynthetic rate (P_N) in the favourable period [7.8±0.6 mol(CO₂) m^–2s^–1, mean value], the highest P_N decrease (on an average 86 % of the maximum) but the highest recovery capacity (>70 % of the maximum) at the first rainfall in September. Cistus incanus L. and Arbutus unedo L. had the highest P_N during the favourable period [15.5±5.2 mol(CO₂) m^–2s^–1, mean value], 79 % decrease during drought, and a lower recovery capacity (on an average 54 %). Quercus ilex L., Phillyrea latifolia L., and Pistacia lentiscus L. had an intermediate P_N in the favourable period [9.2±1.3 mol(CO2) m^–2s^–1, mean value], a lower reduction during drought (on an average 63 %), and a range from 62 % (Q. ilex and P. latifolia) to 39 % (P. lentiscus) of recovery capacity. The Mediterranean species had higher decrease in P_N and stomatal conductance during drought and a higher recovery capacity than the pre-Mediterranean species. Among the pre-Mediterranean species, P. latifoliahad the best adaptation to long drought periods also by its higher leaf mass per area (LMA) which lowered leaf temperature thus decreasing transpiration rate during drought. Moreover, its leaf longevity determined a more stable leaf biomass during the year. Among the Mediteranean species, R. officinalis was the best adapted species to short drought periods by its ability to rapidly recover. Nevertheless, R. officinalis had the lowest tolerance to high temperatures by its P_N dropping below half its maximum value when leaf temperature was over 33.6°C. R. officinalismay be used as a bioindicator species of global change.This revised version was published online in March 2005 with corrections to the page numbers.     

Responses of Ribulose-1,5-Bisphosphate Carboxylase,Protein Content,and Stomatal Conductance to Water Deficit in Maize,Tomato, and Bean

We compared responses of maize, tomato, and bean plants to water stress. Maize reached a severe water deficit (leaf water potential –1.90 MPa) in a longer period of time as compared with tomato and bean plants. Maize stomatal conductance (g _s) decreased at mild water deficit. g _s of tomato and bean decreased gradually and did not reach values as low as in maize. The protein content was maintained in maize and decreased at low water potential (_w); in tomato it fluctuated and also decreased at low _w; in bean it gradually decreased. Ribulose-1,5-bisphosphate carboxylase/oxygenase activity remained high at mild and moderate stress in maize and tomato plants; in bean it remained high only at mild stress.     

Elevated CO2 concentration enhances the role of the ear to the flag leaf in determining grain yield of wheat

Net photosynthetic rate (P _N) of ear and flag leaf during grain filling stage and grain yield of plants with non-darkened or darkened flag leaf or darkened ear were examined in two different CO₂ concentrations: ambient (AC) and AC+200 μmol mol⁻¹ (EC). Ear showed much higher enhancement (56 %) of P _N than flag leaf (23 %) under EC. Moreover, CO₂ enrichment shortened the photosynthetic duration of flag leaf relative to ear. In this way the ratio of ear to flag leaf contribution to grain yield increased from 1.18 (AC) to 1.39 (EC).     

Growth,N2 fixation and photosynthesis in a cyanobacterium,Trichodesmium sp., under Fe stress

Trichodesmium sp., isolated from the Great Barrier Reef lagoon, was cultured in artificial seawater media containing a range of Fe concentration. Fe additions stimulated growth, N₂ fixation, cellular chlorophyll a content, light-saturated chlorophyll a-specific gross photosynthetic capacity (P_m ^chla) and the dark respiration rate (R_d ^chla). Cell yields only doubled for 9 nM Fe relative to zero added Fe, whereas N₂ fixation increased 11-fold considerably for 450 nM Fe. The results suggest that N₂ fixation of Trichodesmium is more sensitive to Fe limitation than are the cell yields.     

Leaf water relations and anatomy of a tropical rainforest tree species vary with crown position

Summary Leaf water potentials, osmotic properties and structural characteristics were examined in the Australian tropical rainforest tree species, Castanospermum australe. These features were compared for individuals growing in the understorey and canopy of the undisturbed forest and in an open pasture from which the forest had been cleared. Leaf water potentials during the day declined to significantly lower values in the open-grown and canopy trees than in the understorey trees. During most of the day the opengrown tree experienced the lowest water potentials. These differences were paralleled by significant differences in tissue osmotic properties. The tissue osmotic potential at full hydration was lowest in the open-grown tree (-1.80 MPa), intermediate in the canopy trees (-1.38 MPa), and highest in the understorey trees (-0.80 MPa). As a result, the degree to which high and positive turgor pressures were maintained as water potentials declined was highest in the open-grown tree, intermediate in the canopy trees, and lowest in the understorey trees. The differences in tissue osmotic properties between individuals in the three crown positions were paralleled, in turn, by differences in leaf structual characteristics. Relative to leaves of the canopy and open-grown trees, leaves of the understorey trees had significantly larger epidermal cells with thinner cell walls, larger specific leaf areas and turgid weight: dry weight ratios, and a higher proportion of intercellular air space.Abbreviations ₁ Leaf tissue water potential - _min Lowest value of ₁ during the day ( noon) - _{P=0 1} zero turgor - R Relative water content - P Tissue turgor pressure - Tissue osmotic potential - ₀ at full hydration