Ear of durum wheat under water stress: water relations and photosynthetic metabolism

The photosynthetic characteristics of the ear and flag leaf of well-watered (WW) and water-stressed (WS) durum wheat (Triticum turgidum L. var. durum) were studied in plants grown under greenhouse and Mediterranean field conditions. Gas exchange measurements simultaneously with modulated chlorophyll fluorescence were used to study the response of the ear and flag leaf to CO₂ and O₂ during photosynthesis. C₄ metabolism was identified by assessing the sensitivity of photosynthetic rate and electron transport to oxygen. The presence of CAM metabolism was assessed by measuring daily patterns of stomatal conductance and net CO₂ assimilation. In addition, the histological distribution of Rubisco protein in the ear parts was studied by immunocytochemical localisation. Relative water content (RWC) and osmotic adjustment (osmotic potential at full turgor) were also measured in these organs. Oxygen sensitivity of the assimilation rate and electron transport, the lack of Rubisco compartmentalisation in the mesophyll tissues and the gas-exchange pattern at night indicated that neither C₄ nor CAM metabolism occurs in the ear of WW or WS plants. Nevertheless, photosynthetic activity of the flag leaf was more affected by WS conditions than that of the ear, under both growing conditions. The lower sensitivity under water stress of the ear than of the flag leaf was linked to higher RWC and osmotic adjustment in the ear bracts and awns. We demonstrate that the better performance of the ear under water stress (compared to the flag leaf) is not related to C₄ or CAM photosynthesis. Rather, drought tolerance of the ear is explained by its higher RWC in drought. Osmotic adjustment and xeromorphic traits of ear parts may be responsible.     

Ear photosynthesis, carbon isotope discrimination and the contribution of respiratory CO2 to differences in grain mass in durum wheat

The role of ear photosynthesis in grain filling was studied in a number of durum wheat (Triticum turgidum var durum L.) landraces and varieties from the Middle East, North Africa, and from the collections of ‘Institut National de la Recherche Agronomique’ (INRA, France) and ‘Centro International de Mejora de Maiz y Trigo’ (CIMMYT, Mexico). Plants were grown in the field in a Mediterranean climate. Flag leaves (blade plus sheath) and ears were kept in the dark from 1 week after anthesis to maturity which reduced grain weight by 22.4% and 59.0%, respectively. In a further experiment, the carbon isotope discrimination ratio (Δ) of ear bracts, awns and flag leaves was measured on samples taken at anthesis and on mature kernels. The mean value of Δ for the water soluble fraction of bracts (17.0‰) and awns (17.7‰) were lower than those of leaves (19.5‰) and fairly similar to those of kernels (17.4‰) averaged across all genotypes. Data indicate that most of the photosynthates in the grain come from ear parts and not from flag leaves. In addition, a higher water use efficiency (WUE) of ear parts than of the flag leaf is suggested by their lower Δ values. Gas exchange in ears and flag leaves was measured during grain filling. Averaged over all genotypes, CO₂ diffusive conductance was about five times higher in the flag leaf than in the spike (with distal portions of awns outside the photosynthetic chamber) 2 weeks after anthesis. In absolute terms, the dark respiration rate (R_d) was greater than the net photosynthesis rate (P_n) by a factor of 1.74 in the spike, whereas R_d was much smaller, only 22.1, 65.7 and 24.8% of P_n in blade, sheath and awns, respectively. Data indicate that photosynthesis, and hence the water use efficiency (photosynthesis/transpiration), is greatly underestimated in ears because of the high rates of respiration which diminish the measured rates of net CO₂ exchange. Results of ¹³C discrimination and gas exchange show that genotypes from North Africa have higher WUE than those from the Middle East. The high R_d values of ears as well as their low diffusive conductance suggest that CO₂ from respiration may be used as source of carbon for ear photosynthesis. In the same way, the anatomy of glumes, for example, supports the role of bracts using internal CO₂ as source of photosynthesis. In the first experiment, the Δ in mature grains from culms with darkened ears compared with control culms provided further evidence in support of this hypothesis. Thus, the Δ from kernels of control plants was 0.40 higher than that from ear-darkened plants, probably because of some degree of refixation (recycling) of respired CO₂ in the grains.     

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.     

Lack of C4 photosynthetic metabolism in ears of C3 cereals

There is continuing controversy over whether a degree of C₄ photosynthetic metabolism exists in ears of C₃ cereals. In this context, CO₂ exchange and the initial products of photosynthesis were examined in flag leaf blades and various ear parts of two durum wheat (Triticum durum Desf.) and two six-rowed barley (Hordeum vulgare L.) cultivars. Three weeks after anthesis, the CO₂ compensation concentration at 210 mmol mol^?1 O₂ in durum wheat and barley ear parts was similar to or greater than that in flag leaves. The O₂ dependence of the CO₂ compensation concentration in durum wheat ear parts, as well as in the flag leaf blade, was linear, as expected for C₃ photosynthesis. In a complementary experiment, intact and attached ears and flag leaf blades of barley and durum wheat were radio-labelled with ¹⁴CO₂ during a 10s pulse, and the initial products of fixation were studied in various parts of the ears (awns, glumes, inner bracts and grains) and in the flag leaf blade. All tissues assimilated CO₂ mainly by the Calvin (C₃) cycle, with little fixation of ¹⁴CO₂ into the C₄ acids malate and aspartate (about 10% or less). These collective data support the conclusion that in the ear parts of these C₃ cereals C₄ photosynthetic metabolism is nil.     

Photosynthesis and conductance of spring wheat ears: field response to free-air CO2 enrichment and limitations in water and nitrogen supply

The mid-day responses of wheat ear CO₂ and water vapour exchange to full-season CO₂ enrichment were investigated using a Free-Air CO₂ Enrichment (FACE) apparatus. Spring wheat [Triticum aestivum (L). cv. Yecora Rojo] was grown in two experiments under ambient and elevated atmospheric CO₂ (C_a) concentrations (approximately 370 μ mol mol ^{− 1} and 550 μ mol mol ^{− 1}, respectively) combined first with two irrigation (Irr) schemes (Wet: 100% and Dry: 50% replacement of evapotranspiration) and then with two levels of nitrogen (N) fertilization (High: 350, Low: 70 kg ha ^{− 1} N). Blowers were used for C_a enrichment. Ambient C_a plots were exposed to blower induced winds as well the C_a × N but not in the C_a × Irr experiment. The net photosynthesis for the ears was increased by 58% and stomatal conductance (g_s) was decreased by 26% due to elevated C_a under ample water and N supply when blowers were applied to both C_a treatments. The use of blowers in the C_a-enriched plots only during the C_a × Irr experiment (blower effect) and Low N supply restricted the enhancement of net photosynthesis of the ear due to higher C_a. In the latter case, the increase of net photosynthesis of the ear amounted to 26%. The decrease in g_s caused by higher C_a was not affected by the blower effect and N treatment. The mid-day enhancement of net photosynthesis due to elevated C_a was higher for ears than for flag leaves and this effect was most pronounced under ample water and N supply. The contribution of ears to grain filling is therefore likely to increase in higher C_a environments in the future. In the comparison between Wet and Dry, the higher C_a did not alter the response of net photosynthesis of the ear and g_s to Irr. However, C_a enrichment increased the drought tolerance of net photosynthesis of the glume and delayed the increase of the awn portion of net photosynthesis of the ear during drought. Therefore, the role of awns for maintaining high net photosynthesis of the ear under drought may decrease when C_a increases.     

In vitro enzyme activities and products of 14CO2 assimilation in flag leaf and ear parts of wheat (Triticum aestivum L.)

Activities of key enzymes of Calvin cycle and C₄ metabolism, rate of ¹⁴CO₂ fixation in light and dark and the initial products of photosynthetic ¹⁴CO₂ fixation were determined in flag leaf and different ear parts of wheat viz. pericarp, awn and glumes. Compared to the activities of RuBP carboxylase and other Calvin cycle enzymes viz. NADP-glyceraldehyde-3-phosphate dehydrogenase, NAD-glyceraldehyde-3-phosphate dehydrogenase and ribulose-5-phosphate kinase, the levels of PEP carboxylase and other enzymes of C₄ metabolism viz. NADP-malate dehydrogenase, NAD-malate dehydrogenase, NADP-malic enzyme, NAD-malic enzyme, glutamate oxaloacetate transaminase genase, NADP-malic enzyme, NAD-malic enzyme, glutamate oxaloacetate transaminase and glutamate pyruvate transaminase, were generally greater in ear parts than in the flag leaf. In contrast to CO₂ fixation in light, the various ear parts incorporated CO₂ in darkness at much higher rates than flag leaf. In short term assimilation of ¹⁴CO₂ by illuminated ear parts, most of the ¹⁴C was in malate with less in 3-phosphoglyceric acid, whereas flag leaves incorporated most into 3-phosphoglyceric acid. It seems likely that ear parts have the capability of assimilating CO₂ by the C₄ pathway of photosynthesis and utilise PEP carboxylase for recapturing the respired CO₂.     

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 awns and drought on the supply of photosynthate and its distribution within wheat ears

The presence of awns doubled the net photosynthetic rate of wheat ears and also increased the proportion of ¹⁴CO₂ assimilated by the ear that moved to the grain. The effect of water supply on photosynthesis and movement of assimilates was greater for leaves than ears, so that drought increased the proportion of assimilate contributed by ear photosynthesis to grain filling from 13% to 24% in the awnless ears, and from 34% to 43% in the awned ears. ¹⁴C assimilated by the ears was most important to the economy of the upper spikelets and to the distal florets in each spikelet, whereas flag leaf assimilate went mainly to the spikelets in the lower half of the ear, and to the proximal florets. Awns increased grain yield in the dry but not in the irrigated treatment, despite the large contribution of awned ears to grain filling. Either the supply of assimilate did not limit grain yield when water supply was not limiting, or there were compensating disadvantages to awns. However, they did not seem to have any adverse effect on the development of the upper florets, nor did they reduce grain number per ear.     

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).     

Effect of assimilate utilization on photosynthetic rate in wheat

Summary Two weeks after anthesis, when the grain is filling rapidly, the rate of photosynthesis by flag leaves of wheat cv. Gabo was between 20 and 30 mg CO₂ dm^-2 leaf surface hour^-1 under the conditions used. About 45% of flag-leaf assimilates were translocated to the ear, and only about 12% to the roots and young shoots.On removing the ear, net photosynthesis by the flag leaves was reduced by about 50% within 3–15 hours, and there was a marked reduction in the outflow of ¹⁴C-labelled assimilates from the flag leaves.Subsequent darkening of all other leaves on plants without ears led to recovery of flag-leaf photosynthesis, as measured by gas analysis and ¹⁴CO₂ fixation, and to increased translocation of assimilates to the roots and young shoots. Minor changes in the rates of dark respiration accompanied these major, reversible changes in photosynthetic rate.After more than a week in continuous, high-intensity light, the rate of photosynthesis by flag leaves of intact plants had fallen considerably, but could be restored again by a period in darkness, or by inhibiting photosynthesis in the ears by spraying them with DCMU. The inhibition of ear photosynthesis increased translocation of labelled assimilates from the flag leaf to the ears, without affecting leaf sugar levels.The application of TIBA to the culm below the ear inhibited auxin movement throught the culm, but had no influence on flag-leaf photosynthesis.These results suggest that, at least in this system, photosynthesis by the flag leaf is regulated directly by the demand for assimilates from the flag leaf and not indirectly through action in the leaf of auxins produced by the sink organs.     

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.     

The temporal and species dynamics of photosynthetic acclimation in flag leaves of rice (Oryza sativa) and wheat (Triticum aestivum) under elevated carbon dioxide

In this study, we tested for the temporal occurrence of photosynthetic acclimation to elevated [CO₂] in the flag leaf of two important cereal crops, rice and wheat. In order to characterize the temporal onset of acclimation and the basis for any observed decline in photosynthetic rate, we characterized net photosynthesis, g_s, g_m, C_i/C_a, C_i/C_c, V_cmax, J_max, cell wall thickness, content of Rubisco, cytochrome (Cyt) f, N, chlorophyll and carbohydrate, mRNA expression for rbcL and petA, activity for Rubisco, sucrose phosphate synthase (SPS) and sucrose synthase (SS) at full flag expansion, mid‐anthesis and the late grain‐filling stage. No acclimation was observed for either crop at full flag leaf expansion. However, at the mid‐anthesis stage, photosynthetic acclimation in rice was associated with RuBP carboxylation and regeneration limitations, while wheat only had the carboxylation limitation. By grain maturation, the decline of Rubisco content and activity had contributed to RuBP carboxylation limitation of photosynthesis in both crops at elevated [CO₂]; however, the sharp decrease of Rubisco enzyme activity played a more important role in wheat. Although an increase in non‐structural carbohydrates did occur during these later stages, it was not consistently associated with changes in SPS and SS or photosynthetic acclimation. Rather, over time elevated [CO₂] appeared to enhance the rate of N degradation and senescence so that by late‐grain fill, photosynthetic acclimation to elevated [CO₂] in the flag leaf of either species was complete. These data suggest that the basis for photosynthetic acclimation with elevated [CO₂] may be more closely associated with enhanced rates of senescence, and, as a consequence, may be temporally dynamic, with significant species variation.     

Photosynthesis of Ears and Flag Leaves of Wheat and Barley

Immediately after anthesis ears of spring wheat absorbed lessthan 0.5 mg CO₂, per hour in daylight and later evolved CO₂,in the light and in the dark. The rate of apparent photosynthesisof the combined flag-leaf lamina and sheath and peduncle (collectivelycalled flag leaf) of two spring wheat varieties, Atle and JufyI, was 3–4 mg per hour; the rates of the flag leaf andthe ear of two spring barleys, Plumage Archer and Proctor, wereeach about 1 mg per hour. The gas exchange of ears and flag leaves between ear emergenceand maturity accounted for most of the final grain dry weight.The CO₂, fixed by the wheat ear was equivalent to between 17and 30 per cent of the grain weight, but more than this waslost by respiration, so assimilation in the flag leaf was equivalentto 110–20 per cent of the final grain weight. In barley,photosynthesis in the flag leaf and the net CO₂ uptake by theear each provided about half of the carbohydrate in the grain. Barley ears photosynthesized more than wheat ears because oftheir greater surface, and flag leaves of wheat photosynthesizedmore than those of barley because they had more surface anda slightly greater rate of photosynthesis per dm².     

Carbon isotope ratios in ear parts of triticale : influence of grain filling

Four triticale (×Triticosecale Wittmack) genotypes were grown under rainfed conditions with limited irrigation support in Lleida in northeast Spain. For each variety, samples consisting of 10 tillers with half-sterilized spikes were taken three times from anthesis to maturity. Carbon isotope ratios (δ¹³C) were then determined in water extracts from ear bracts (glumes, paleas, and lemmas), awns and flag leaves, and in powdered kernels. For the half-sterilized spikes, carbon isotope analysis was carried out separately in bracts and awns from fertile and nonfertile spikelets. The δ¹³C in the water-soluble fraction of awns, glumes, and glumells from fruitless spikelets was significantly higher than that from fertile spikelets sampled at mid-grain filling. Differences in δ¹³C among sterile and fertile spikelets were not significant in samples taken a few days after anthesis or at maturity. These results are in accordance with some degree of refixation by awns and ear bracts of the CO₂ respired by grains during grain filling. There was progressively higher δ¹³C from flag leaf blades to awns, glumes, and glumells. This variation in δ¹³C along plant parts may be caused by differences in the ratio of assimilation rate to CO₂-diffusive conductance. Values of δ¹³C of mature kernels were between the values at anthesis and mid-grain filling for the water-soluble fraction of flag leaves and inner bracts and were fairly similar to those of glumes and awns.     

Persistence of photosynthetic components and photochemical efficiency in ears of water-stressed wheat (Triticum aestivum)

Ear photosynthesis may be an important source of C for grain growth in water-stressed plants of cereals. The main objectives of this work were to determine the stability of the photosynthetic apparatus and the photochemical efficiency of ears in plants subjected to post-anthesis drought. Plants of wheat ( Triticum aestivum L. cv. Granero INTA) were grown in pots under a rain shelter and subjected to water stress (soil water potential around −0.6 to −0.8 MPa) starting 4  days after anthesis. Post-anthesis drought substantially accelerated the loss of chlorophyll, Rubisco and the light-harvesting complex of photosystem II (LHCII) in the flag leaf, but the degradation of these photosynthetic components was much less affected by water deficit in awns and ear bracts. Quantum yield of PSII (Φ_PSII) decreased in leaves of water-stressed plants. In contrast, ear bracts had a higher Φ_PSII than leaves, and Φ_PSII of ear bracts did not decrease at all in response to drought. Removing the grains immediately before fluorescence measurements (less than 30 min) slightly reduced Φ_PSII, indicating that CO₂ supplied by grain respiration may contribute to the high photochemical efficiency of ears in droughted plants. However, other factors may be involved in maintaining high Φ_PSII, since even in the absence of grains Φ_PSII remained much higher in ear bracts than in the flag leaf. The relative stability of ear photosynthetic components and their relatively high photochemical efficiency may help to maintain ear photosynthesis during the grain filling period in droughted plants.     

Wheat flag leaf epicuticular wax morphology and composition in response to moderate drought stress are revealed by SEM,FTIR‐ATR and synchrotron X‐ray spectroscopy

Wheat (Triticum aestivum L.) is the largest cereal crop grown in Western Canada where drought during late vegetative and seed filling stages affects plant development and yield. To identify new physiochemical markers associated with drought tolerance, epidermal characteristics of the flag leaf of two wheat cultivars with contrasting drought tolerance were investigated. The drought resistant ‘Stettler’ had a lower drought susceptibility index, greater harvest index and water‐use efficiency than the susceptible ‘Superb’. Furthermore, flag leaf width, relative water content and leaf roll were significantly greater in Stettler than in Superb at moderate drought stress (MdS). Visible differences in epicuticular wax density on the adaxial flag leaf surfaces and larger bulliform cells were identified in Stettler as opposed to Superb. Mid‐infrared attenuated total internal reflectance spectra revealed that Stettler flag leaves had increased asymmetric and symmetric CH₂ but reduced carbonyl esters on its adaxial leaf surface compared to Superb under MdS. X‐ray fluorescence spectra revealed a significant increase in total flag leaf Zn concentrations in Stettler in response to MdS. Such information on the microstructural and chemical features of flag leaf may have potential as markers for drought tolerance and thereby accelerate the selection and release of more drought‐resistant cultivars.     

Modelling C3 and C4 photosynthesis under water-stressed conditions

Despite the observed impact of water stress on photosynthesis, some of the most used models of CO₂ assimilation in C₃ and C₄ functional types do not directly account for it. We discuss an extension of these models, which explicitly includes the metabolic and diffusive limitations due to water stress on photosynthesis. Functional relationships describing the photosynthetic processes and CO₂ diffusion inside leaves are modified to account for leaf water status on the basis of experimental results available in the literature. Extensive comparison with data shows that the model is suitable to describe the reduction in CO₂ assimilation rate with decreasing leaf water potentials in various species. A simultaneous analysis of photosynthesis, transpiration and soil moisture dynamics is then carried out to explore the actual impact of drought on different photosynthesis processes and on the overall plant activity. The model well reproduces measured CO₂ assimilation rate as a function of soil moisture and could be useful to formulate hypotheses for detailed experiments as well as to simulate in detail transpiration and photosynthesis dynamics under water stress.     

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.     

Water use efficiency in C3 cereals under Mediterranean conditions: a review of physiological aspects

In this review, we will discuss physiological traits of C₃ cereals related to water use efficiency (WUE) in Mediterranean environments, from leaf (WUE_{instantaneous}) to crop level (WUE_yield or ‘water productivity’). First, we analyse the WUE_{instantaneous} and the possible trade‐off between improving this parameter and growth/yield performance. Ways to ameliorate WUE without penalties are discussed. We also analyse in what cases breeding by high or low WUE_{instantaneous} is a suitable criterion to maintain grain yield under drought (Mediterranean) conditions. This question is approached in the framework of carbon isotope discrimination, (Δ¹³C), the main indirect parameter used to integrate (at time and space scale) the WUE_{instantaneous} in C₃ plants. A negative correlation between these two parameters has been confirmed by several studies. The relationship between Δ¹³C and grain yield, however, is more complex, and may differ from one environment to another. In Mediterranean conditions with moderate or no water stress, a positive correlation between Δ¹³C and grain yield is found in barley and wheat, whereas in ‘stored‐water’ crops (such as in some regions of Australia), lower Δ¹³C (i.e. higher WUE_{instantaneous}) is associated with higher grain yield, particularly in more stressful conditions. These apparent inconsistencies and their possible implications for plant breeding are discussed. One physiological trait that has received minor attention in attempts to improve WUE_{instantaneous} is the role of ear photosynthesis. Ears of barley and durum wheat have a higher WUE_{instantaneous} than the flag leaf, both in well‐watered and in drought conditions. The underlying causes of the higher WUE_{instantaneous} of ears are not fully understood, but their refixation capacity (i.e. the capacity to re‐assimilate respired carbon dioxide) could be important. Although the genotypic variability of this trait has not been extensively studied, some data support the idea that variation in refixation capacity may be attributable to genetic factors. At the crop level, decreasing soil evaporation is a crucial factor in efforts to improve the WUE_yield in Mediterranean conditions, and fast initial growth of the crop (i.e. early vigour) seems to be relevant. In wheat, modern varieties with dwarfing genes (giberellic acid – insensitive) have higher yields but, concomitantly, they have lower initial growth performance. Recently, semi‐dwarf cultivars (giberellic acid – sensitive) with high grain yield and simultaneously high early vigour were found, opening new avenues to increase WUE_yield in wheat. The negative effects of futile water loss by cuticular and nocturnal transpiration are also commented. Finally, we discuss some agronomic practices (in particular, ‘deficit irrigation’ systems) linked to physiological traits that confer higher WUE_yield,, in particular, in the cases of Mediterranean regions.     

Contribution of the ear and the fl ag leaf to grain fi lling in durum wheat inferred from the carbon isotope signature: Genotypic and growing conditions eff ects FA

The ear, together with the flag leaf, is believed to play a major role as a source of assimilates during grain filling in C3cereals. However, the intrusive nature of most of the available methodologies prevents reaching conclusive results in this regard. This study compares the carbon isotope composition（d13C） in its natural abundance in the water‐soluble fractions of the flag leaf blade and the ear with the d13C of mature kernels to assess the relative contribution of both organs to grain filling in durum wheat（Triticum turgidum L. var.durum）. The relative contribution of the ear was higher in landraces compared to modern cultivars, as well as in response to nitrogen fertilization and water stress. Such genotypic and environmentally driven differences were associated with changes in harvest index（HI）, with the relative contribution of the ear being negatively associated with HI. In the case of the genotypic differences, the lower relative contribution of the ear in modern cultivars compared with landraces is probably associated with the appearance in the former of a certain amount of source limitation driven by a higher HI. In fact, the relative contribution of the ear was far more responsive to changes in HI in modern cultivars compared with landraces.