Carbon Isotope Discrimination, Leaf Ash Content and Grain Yield in Bread and Durum Wheat Grown under Full-Irrigated Conditions

Integrative physiological criteria, such as carbon isotope discrimination (Δ) and (mineral) ash content (m_a) have been found to be very useful, under drought conditions, to elucidate the association between yield gains and variation of photosynthesis‐related traits and orientate future breeding efforts. Information on this association is scarce under irrigated conditions. The relationships between Δ, m_a and yield were studied in bread wheat (Triticum aestivum L.) and durum wheat (Triticum turgidum L. var. durum) under optimal (drip) irrigation in the arid conditions of north‐west Mexico. Carbon isotope discrimination was analysed on leaves at booting stage and anthesis and on grain at maturity, whereas ash content was measured on the flag leaf at anthesis and maturity. At anthesis, there were differences between bread and durum wheat during grain filling for Δ, but not for m_a. No relationship was found between grain yield and Δ. Leaf ash content at anthesis and maturity showed a broad variability within each species and were associated with grain yield. These results suggest that ash content in leaves could be also used as predictive criteria for yield not only under drought, but also under irrigated conditions, particularly when evaporative demand is high.     

Relationships between Grain Yield, Flag Leaf Morphology, Carbon Isotope Discrimination and Ash Content in Irrigated Wheat

The purpose of this study was to examine how differences in leaf angle, leaf rolling (LR) and glaucousness (GL) can modify yield components and leaf physiological traits in wheat. A set of 167 lines derived from a cross between two high‐yielding bread wheat cultivars differing for these traits was grown under flood irrigation and high evaporative demand in the north‐west of Mexico. Area, mass per unit area and chlorophyll content of the flag leaf were assessed. Carbon isotope discrimination (Δ) and ash content (m_a) were also measured. A significant correlation was found between grain yield (GY) and both Δ and m_a suggesting that, despite well‐watered conditions, leaf stomatal conductance was the main yield‐limiting factor. Leaf posture and LR did not significantly affect yield, Δ and m_a. Higher grain weight was noted, however, in lines with droopy flag leaves. Erect leaves had higher mass per unit area. GL was associated with a significant increase in GY and grain weight. Glaucous lines also had higher Δ and m_a, suggesting higher transpiration rate and lower transpiration efficiency. The study confirms that Δ and m_a represent promising criteria for GY in wheat and provides evidence that GL can contribute to higher yield, even under irrigated conditions.     

Relationship between Carbon Isotope Discrimination and Mineral Content in Wheat Grown under Three Different Water Regimes

Carbon isotope discrimination (Δ) has been proposed as an indirect selection criterion for transpiration efficiency and grain yield in wheat. However, because of the high cost for Δ analysis, attempts have been carried out to identify alternative screening criteria. Ash content (m_a) has been proposed as an alternative criterion for Δ in wheat and barley. A pot experiment was conducted to analyse the relationship between Δ and m_a in flag leaf and grain. Plants of 10 genotypes were cultivated under three different water regimes corresponding to moderate, intermediate and severe drought stress obtained by maintaining soil humidity at 75 %, 55 % and 45 % of the humidity at field capacity, respectively. Δ and m_a in flag leaf and grain showed significant differences between the moderate, intermediate and severe drought stress levels. Significant correlations were found among genotypes for Δ and m_a in flag leaf under severe drought stress, and for Δ and m_a in grain under intermediate and moderate drought stress. In flag leaf at anthesis, Δ was negatively associated to K content and positively to Mg content. At maturity, Δ in grain was negatively correlated with Mg and Ca contents in flag leaf and grain, respectively. These results suggested that these traits may be potentially useful traits, which could be surrogates for Δ.     

Relationship between Carbon Isotope Discrimination,Mineral Content and Gas Exchange Parameters in Vegetative Organs of Wheat Grown under Three Different Water Regimes

Carbon isotope discrimination (Δ) has been proposed as an indirect selection criterion for transpiration efficiency and grain yield in wheat. However, because of high cost for Δ analysis, attempts have been made to identify alternative screening criteria. Ash content (m_a) has been proposed as an alternative criterion for Δ in wheat and barley. A pot experiment was conducted to analyse the relationship between Δ, mineral content and gas exchange parameters in seedlings and leaves of bread wheat (Triticum aestivum L.). Plants of 10 genotypes were cultivated under three different water regimes corresponding to moderate (T₃), intermediate (T₂) and severe drought (T₁) stress obtained by maintaining soil humidity at 75 %, 55 % and 45 % of the humidity at field capacity respectively. Δ and m_a in seedlings and leaves showed significant differences among the three water treatments. Significant positive correlations were found between Δ and m_a in seedlings and leaves at elongation and anthesis stages in severe drought stress (T₁). Δ was negatively associated with potassium (K) content in intermediate drought stress (T₂) and positively with magnesium (Mg) content in T₂ and T₃ (moderate drought stress) in flag leaf at anthesis. There were negative correlations between Δ and single‐leaf intrinsic water‐use efficiency (W_T) in T₂ and T₃ at anthesis stage. Stronger positive associations were noted between Δ and stomatal conductance (g_s) in T₁ and T₂ than in T₃ at anthesis. These results suggested that Δ is a good trait as an indirect selection criterion for genotypic improvement in transpiration efficiency, while m_a is a possible alternative criterion of Δ in wheat vegetative organs, especially in stressed environments. Significant association was found between Δ and K, Mg and Ca contents that would merit being better investigated.     

Relationship between Carbon Isotope Discrimination and Grain Yield in Spring Wheat under Different Water Regimes and under Saline Conditions in the Ningxia Province (North-west China)

The relationship between grain yield and carbon isotope discrimination (Δ) was analysed in wheat grown under different water regimes in the Ningxia Province (north‐west of China). When the association was significant, the relationships between grain yield, Δ and other drought tolerance related traits, such as leaf ash content (m_a), chlorophyll concentration (Chl), relative water content (RWC), stomatal conductance (g_S) and the ratio of internal CO₂ leaf concentration to ambient CO₂ concentration (C_i/C_a), were also examined. Using correlation analysis, the relationships were determined during two consecutive years in a set of 20 spring wheat cultivars (landraces, improved varieties and advanced lines) under rainfed and irrigated conditions, including saline conditions. The relationship between Δ and yield within environments highly depended on the quantity of water stored in the soil at sowing, the quantity and distribution of rainfall during the growth cycle, and the irrigation before anthesis. Δ predicted grain yield under limited irrigation (post‐anthesis water stress) but not under pre‐anthesis water stress (rainfed conditions), fully irrigated and saline conditions. Under limited irrigation, grain Δ correlated significantly to grain yield leaf m_a at heading and maturity. It also significantly positively correlated to Chl, RWC, g_S and C_i/C_a assessed at anthesis. A precise characterization of the timing and intensity of the abiotic constraints experienced by the crop is consequently needed before implementing the use of Δ in wheat breeding programmes.     

Association between Yield and Carbon Isotope Discrimination Value in Different Organs of Durum Wheat Under Drought

Carbon isotope discrimination (Δ) has been proposed as a selection criterion for transpiration efficiency and grain yield in drought‐prone environments for several C³ species, including cereals. Δ analysis, however, has mainly been concerned with grain or culm tissues and little work has been devoted to other organs. The objective of this study was to describe Δ variation in different organs and to examine the relationships between Δ and grain yield across environments. Six durum wheat genotypes with contrasted grain Δ were cultivated under rainfed conditions during three successive years at Montpellier (South of France). Δ was measured on flag leaf, stalk, awns, chaff and rachis sampled at anthesis and maturity, and on mature grain. Higher genotypic variation and closer correlation with yield were noted for grain Δ compared to other plant parts. The genotype ranking across years was more consistent for grain Δ than for other organs. Consequently, the grain seems the most effective plant part for Δ analysis in durum wheat under Mediterranean conditions. The study of Δ variation in other organs may be useful, however, to evaluate the contribution of those organs to grain filling and final yield according to environmental conditions.     

Evaluating Leaf Ash Content and Potassium Concentration as Surrogates of Carbon Isotope Discrimination in Grassland Species

Leaf ash content and potassium (K) concentration were evaluated as surrogates of carbon isotope discrimination (Δ) of the dominant species in an upland grassland in a 3-year experiment. Previous results reported positive correlations between Δ and ash content or mineral concentrations, attributing this relationship to the passive accumulation, through the transpiration stream, of minerals in the vegetative plant parts. We found only a weak positive correlation between Δ and ash content or K concentration in 1997 (r=0.79, P < 0.05 and r=0.77, P < 0.05, respectively). No significant correlation was found between the traits in 1998 and 1999. When four of the seven most abundant species ( Poa pratensis , Lolium perenne , Festuca valida and Taraxacum officinale ) were grown under different treatments (mixed sward, monoculture, and nitrogen and P additions in mixed sward), a curvilinear relationship between Δ and ash content or K concentration was evident. Thus, our findings suggest that leaf ash content and K concentration cannot serve as surrogates of carbon isotope discrimination (Δ). Small genotype × environment interactions were reported for the three traits measured; the interactions were smallest for ash content. A positive correlation between leaf ash content and soil water content (at 5 and 15 cm depth) was found for four species ( P. pratensis , L. perenne , F. valida and Achillea millefolium ) and for the community as a whole. No significant correlation was found between leaf K concentration and soil water content.     

Effect of Glaucousness on Carbon Isotope Discrimination and Grain Yield in Durum Wheat

The effect of glaucousness on yield and transpiration efficiency was studied in durum wheat under Mediterranean conditions. Two groups of 16 genotypes, with contrasting glaucousness, were compared over 2 years for flag leaf and kernel carbon isotope discrimination, residual transpiration, specific leaf dry weight, biomass and grain yield. Significantly higher carbon isotope discrimination and lower specific leaf dry weight were noted in the glaucous lines. No significant effect of glaucousness was found on residual transpiration. Glaucous genotypes also exhibited higher grain yield, which was strongly correlated to the carbon isotope discrimination values. Results suggest that glaucousness has a negative effect on transpiration efficiency, which could be related to modifications of leaf energy balance. Higher stomatal conductance, reflected by the high carbon isotope discrimination, could mainly explain the higher grain yield found in the glaucous lines.     

Analysis of Wheat Cultivar Differences in Grain Yield, Grain Nitrogen Yield and Nitrogen Utilization Efficiency.

More detailed information on the causes of yield variability among wheat cultivars is needed to further increase wheat yield. Field studies were conducted in Northern Greece over the two cropping seasons of 1985—1986 and 1986—1987 to assess the effects of nitrogen fertilizer and application timing of the various component traits that determine grain yield, grain nitrogen yield and nitrogen utilization efficiency of two bread ( Triticum aestivum L.) and two durum ( Triticum durum Desf.) wheat cultivars, using yield and yield component analysis. Nitrogen at a rate of 150 kg ha^-1 was applied before planting or 100 N kg ha^-1 before planting and then 50 N kg ha^-1 top dressed at early boot stage. Nitrogen and cultivars affected all traits examined, while split nitrogen application affected only some of the traits. Grain yields in the most cases were correlated with number of grains per unit area and grain weight and grain nitrogen yields in all cases with grain number per unit area. The contribution of the number of grains per spike to total variation in grain yield among cultivars was almost consistent (37 to 55 %), while the contribution of grain weight was more significant (up to 55 %) in high yields (>6.500kg ha^-1) and number of spikes per unit area (>500). The number of grains per spike contributed from 60 to 83 % to the total variation in grain nitrogen per spike. Increased grain nitrogen concentration resulted in a reduction of its contribution in grain nitrogen yield variation. Nitrogen utilization efficiency was higher during grain filling than during vegetative biomass accumulation. The contribution of nitrogen harvest index to the variation of utilization efficiency for grain yield was higher in plants receiving nitrogen application.     

Carbon Isotope Discrimination and Grain Yield Variations among Tetraploid Wheat Species Cultivated under Contrasting Precipitation Regimes

Nineteen accessions belonging to six tetraploid wheat species were evaluated and compared for carbon isotope discrimination and grain yield during two successive cropping seasons (1995 and 1996) under Mediterranean rainfed conditions. The two seasons differed markedly in precipitation regime. A wide variation for the measured traits was observed among the 19 accessions for carbon isotope discrimination and grain yield in both years. Significant differences were also noted between species accessions for Δ and grain yield. In both years, Triticum timopheevi (AG genome) showed lower Δ values than the other tetraploid species (all carrying the AB genome). Positive correlations were observed between Δ and grain yield in both years. The use of carbon isotope discrimination as an indirect selection criterion for yield under stress and the potential benefits of some alien tetraploid species in improving drought tolerance in durum wheat are discussed.     

Evaluation of Grain Filling Rate and Duration in Bread and Durum Wheat, under Heat Stress after Anthesis

Bread and durum wheat genotypes were submitted to heat stress during the grain filling period, and relationships between grain weight and accumulated time from anthesis until maturity, using days after anthesis and growing degree days, were described by cubic polynomials. Maximum grain weight and the duration and rate of grain filling were estimated from the fitted curves. It was found that bread and durum wheat exposure to high temperatures significantly decreased grain weight and hastens physiological maturity (shortening the grain filling period). High temperatures significantly affected the rate (on a growing degree day basis) and duration (on Julian day units) of grain filling. The grain filling rate, on a thermal time basis, was positively associated with the final grain weight and the estimated maximum grain weight. The duration of grain filling does not appear to be a limiting factor for genotype grain weight stability, being mainly fixed by temperature. Grain weight of the controlled plants was positively correlated with the final and maximum grain weight of heat stressed plants. It was concluded that a high grain filling rate and a high potential grain weight are major traits that can be useful to improve heat tolerance of Triticum under Mediterranean environments.     

Contribution of Different Organs to Grain Filling in Durum Wheat under Mediterranean Conditions I. Contribution of Post‐Anthesis Photosynthesis and Remobilization

Under Mediterranean conditions, drought affects cereals production principally through a limitation of grain filling. In this study, the respective role of post‐anthesis photosynthesis and carbon remobilization and the contribution of flag leaf, stem, chaff and awns to grain filling were evaluated under Mediterranean conditions in durum wheat (Triticum turgidum var. durum) cultivars. For the purpose, we examined the effects of shading and excision of different parts of the plant and compared carbon isotope discrimination (Δ) in dry matter of flag leaf, stem, chaff, awns and grain at maturity and in sap of stem, flag leaf, chaff and awns, this last measurement providing information on photosynthesis during a short period preceding sampling. Source–sink manipulations and isotopic imprints of different organs on final isotope composition of the grain confirmed the high contribution of both carbons assimilated by ears and remobilized from stems to grain filling, and the relatively low contribution of leaves to grain filling. Grain Δ was highly and significantly associated with grain yield across treatments, suggesting the utilization of this trait as an indicator of source–sink manipulations effects on grain yield. Chaff and awns Δ were better correlated with grain Δ than stem and leaf Δ, indicating that chaff were more involved in grain filling than other organs. Moreover, in chaff, sap Δ was highly significantly correlated with dry matter Δ. These results suggest the use of Δ for a rapid and non‐destructive estimation of the variation in the contribution of different organs to grain filling.     

Effects of Zinc Deficiency and Drought on Grain Yield of Field-grown Wheat Cultivars in Central Anatolia

Drought stress and zinc (Zn) deficiency are serious abiotic stress factors limiting crop production in Turkey, especially in Central Anatolia. In this study, the effects of Zn deficiency and drought stress on grain yield of 20 wheat cultivars (16 bread wheat, Triticum aestivum; four durum wheat, Triticum durum cultivars) were investigated over 2 years under rainfed and irrigated conditions in Central Anatolia where drought and Zn deficiency cause substantial yield reductions. Plants were treated with (+Zn: 23 kg Zn ha⁻¹, as ZnSO₄·7H₂O) and without (−Zn) Zn under rainfed and irrigated conditions. Both Zn deficiency and rainfed treatments resulted in substantial decreases in grain yield. Significant differences were determined between both bread wheat and durum wheat cultivars in terms of drought stress tolerance. Considering drought sensitivity indices over 2 years, the bread wheat cultivars Yayla‐305, Gerek‐79, Dagdas‐94 and Bolal‐2973 were found to be more drought‐tolerant than the other cultivars under both −Zn and +Zn treatments. Especially the durum wheat cultivars Cakmak 79 and Selcuklu 97 showed much greater drought susceptibility under Zn deficiency, and irrigation alone was not sufficient to obtain satisfying grain yield without Zn application. The results indicate that sensitivity to Zn deficiency stress became more pronounced when plants were drought‐stressed. The effect of irrigation on grain yield was maximized when Zn was adequately supplied, leading to the suggestion that efficient water use in Central Anatolia seems to be highly dependent on the Zn nutritional status of plants.     

Relationship of Carbon Isotope Discrimination to Water Use Efficiency and Productivity of Barley Under Field and Greenhouse Conditions

This study was conducted to evaluate the application of carbon isotope discrimination (CID) as a selection criterion for improving water use efficiency (WUE) and productivity of barley (Hordeum vulgare L.) under field and drought‐stress conditions in a greenhouse. A total of 54 genotypes were screened for variability in CID under field conditions, while 23 genotypes were evaluated under water‐deficit conditions in the greenhouse. A survey of leaf CID of 54 genotypes at two field locations showed more than 2.14‰ difference between extreme genotypes. Significant (P ≤ 0.05) genotypic variation was found in WUE and CID that had a negative strong correlation. There was a negative correlation between leaf CID and aerial biomass in the greenhouse and among six‐row genotypes in the field. Correlations between leaf CID across field locations and across irrigation regimes in the greenhouse were significant (experiment 1, r = 0.79 and 0.94 for six‐ and two‐row genotypes), suggesting stability of the CID trait across different environments. Overall, these results indicate the potential of leaf CID as a reliable method for selecting for high WUE and productivity in barley breeding programmes in the Canadian prairies. Further work is currently underway to determine heritability/genetics of leaf CID and application of molecular marker‐assisted selection for the traits in barley breeding programmes.     

Production of haploids in bread wheat, durum wheat and hexaploid triticale crossed with pearl millet

Pearl millet is an efficient alternative to maize as a pollen source for haploid production in bread wheat. To compare haploid production frequencies in other Triticeae species, the crossabilities of two genotypes each of bread wheat, durum wheat and hexaploid triticale with four pearl millet genotypes and a maize control were examined. Embryos were obtained from crosses of all three species with both pearl millet and maize. However, significant differences in crossability were found among the three species (10.5–79.8% seed development and 1.4–15.8% embryo formation), as well as among genotypes of durum wheat (7.2–23.7% and 2.1–6.4%) and hexaploid triticale (0.3–20.6% and 0.1–2.7%). Crossability of bread wheat with pearl millet was relatively high. Haploid plants were regenerated from crosses of all three species with pearl millet. As in the case of maize crosses, low crossabilities of durum wheat and hexaploid triticale with pearl millet can be attributed to the absence of D-genome chromosomes.     

Leaf Carbon Isotope Discrimination as an Accurate Indicator of Water‐Use Efficiency in Sunflower Genotypes Subjected to Five Stable Soil Water Contents

Leaf carbon isotope discrimination (CID) has been suggested as an indirect tool for breeding for water‐use efficiency (WUE) in various crops. This work focused on assessing phenotypic correlations between WUE and leaf CID and analysing genotypic variability in four sunflower genotypes grown in a greenhouse in pots with five different stable levels of soil water content (SWC). We measured WUE at whole plant and leaf (intrinsic) level. At whole plant level, WUE was derived from the ratio of total dry aerial biomass (BM) to cumulative water transpired (CWT). At leaf level, intrinsic WUE was calculated as the ratio of light‐saturated CO₂ assimilation to stomatal conductance (A/g_s) in younger expanded leaves. Significant differences among the four genotypes and the five SWCs were observed for whole plant and leaf WUE and CID. Strong negative correlations were observed between whole plant WUE and CID as well as between intrinsic WUE and CID with decreasing water availability. No relationships appeared between BM production and WUE or CID. Our results can help agronomists and breeders to evaluate sunflower lines with high WUE for adaptation to drought conditions and for reducing water consumption and crop water needs. Leaf CID appears to be a pertinent and valuable trait to select sunflower genotypes with high WUE.     

Water Stress, Water Use Efficiency, Carbon Isotope Discrimination and Leaf Gas Exchange Relationships of the Bush Bean

Water use efficiencies for the whole plant (WUE₁) and single leaves (WUE₁) were studied in a greenhouse as a function of soil moisture during four phenological stages of bush bean growth. WUE₁ increased significantly with soil moisture stress and attained its maximum value before the flowering stage. WUE₁ and WUE₁ were linearly related ( r = 0.92), and WUE₁ was correlated with the transpiration rate ( r = -0.87), stomatal conductance ( r = -0.80) and photosynthetic rate ( r = 0.81). Carbon isotope discrimination.), decreased as soil moisture decreased, and) was negatively correlated with both WUE₁ ( r = -0.92) and WUE₁ ( r = -0.88). There were significant differences in leaf N among water regimes.     

小麦籽粒淀粉分支酶同工酶基因型与酶活性关系研究

测定了71个小麦品种的籽粒淀粉分支酶活性,并采用非变性聚丙烯酰胺凝胶电泳（Native-PAGE）,鉴定了分支酶同工酶的基因型,以明确小麦籽粒淀粉分支酶（SBE）各同工型等位基因的分布特点和基因型组成,阐明分支酶活性与分支酶同工酶基因型的相关性。结果表明,分支酶有SBEⅠ和SBEⅡ两个基因位点,SBEⅡ位点未显现多态性,SBEⅠ位点存在多样性。在SBEⅠ位点鉴定出A、B、D_ⅰ和D_ⅱ共4个等位基因位点,各等位基因位点出现频率分别为9.9%、76.1%、95.8%和57.8%;共检测到9种基因型,其中基因型D_ⅰD_ⅱB、D_ⅰB和D_ⅰD_ⅱ有较高的分布频率,分别为45.1%、28.2%和8.5%。从单一位点分析,分支酶活性与SBEⅠ位点有显著的相关性,与SBEⅡ位点相关性不显著。由此可见,小麦籽粒胚乳淀粉分支酶同工酶不同基因位点和基因型对酶活性具有不同的遗传效应。     

Water Use Efficiency, Carbon Isotope Discrimination and Biomass Production of Two Sugar Beet Varieties Under Well-Watered and Dry Conditions

Two sugar beet (Beta vulgaris sp.) varieties, which were supposed to differ in drought tolerance, were exposed to drought stress in a growth chamber and a container experiment in field. The aim was to test for (i) differences between the varieties in water use efficiency (WUE), biomass production and distribution and (ii) the relationship between WUE and carbon isotope discrimination (Δ), and between biomass production and Δ. Significant differences in WUE were detected between plants of well‐watered and drought treatments in both experiments, but not between the varieties. Production losses due to drought were large for both varieties in both experiments. Losses in the growth chamber were up to 50 % of plant dry weight and the corresponding value in the field was 24 %, when plants were given 60 and 30 %, respectively, of the full‐watered treatments. Significant negative correlations between WUE and Δ were found, but not between biomass production and Δ, when both varieties were included. Negative correlations between WUE and Δ were also found for each variety separately. The results suggest that Δ estimates from leaf tissue of sugar beet may provide a useful tool for genetic selection of drought‐tolerant sugar beet varieties.     

Irrigation and Nutrient Effects on Growth and Water–Yield Relationship of Wheat (Triticum aestivum L.) in Central India

Increasing production of wheat from a limited water supply can result from efficient irrigation and nutrient management. A 3‐year field experiment was conducted at the Indian Institute of Soil Science, Bhopal, to study the growth, yield, seasonal evapotranspiration (ET) and water use efficiency (WUE), and the water–yield relationship of wheat in a soybean–wheat cropping system on vertisols. Three levels of irrigation, viz. I₀, no post‐sowing irrigation; I₁, two irrigations [crown root initiation (CRI) and flowering stage]; and I₂, three irrigations (CRI, maximum tillering and flowering stage) and three nutrient management treatments, viz. F₀, control (without fertilizer/manure); F₁, 100 % NPK (100–21.5–24.9 kg ha^?1); and F₂, 100 % NPK + farmyard manure (FYM‐10 t ha^?1) were tested in a split‐plot design with three replication. It has been established (through anova ) that the year effect was rather negligible and the interaction effects of irrigation and nutrient management on the growth parameters, ET, yield components, yield and WUE were significant. Plant height, progressive leaf area index, dry matter accumulation and crop growth rate were higher in I₂F₂, and I₂F₁ and I₁F₂ were statistically at par. The seasonal ET increased significantly with the increase in water supply in every nutrient treatment and it was highest in I₂F₂ and lowest in I₀F₀. The highest grain yield was obtained in I₂F₂; and a similar yield was recorded in I₃F₁ and I₂F₂. This shows a strong interaction effect between irrigation and nutrients. Yield components, viz. number of ears m^?2, number of grains ear^?1 and 1000‐grain weight were significant. The higher number of ears m^?2 containing greater number of grains with relatively heavier weights appeared to have contributed to the higher yield in I₁F₂, I₂F₁ and I₂F₂. The highest WUE obtained in I₀F₂ did not correspond to the highest yield and maximum ET, but a WUE of 10.43 kg ha^?1 mm^?1 in the I₂F₂ combination corresponded with the highest yield and the seasonal ET requirement was 391.8, which was 137 % greater than the water use at maximum WUE. The ET–grain yield relationship was linear, with a lowest regression slope (i.e. marginal WUE) and elasticity of water production (E_wp) in F₀ and a considerably higher slope and E_wp in F₁ and F₂. As the E_wp is positive and close to one in 100 % NPK treatment, the scope of improving WUE and yield with only inorganic fertilizer is very little, and relatively greater scope exists in the integrated management of organic manure and inorganic fertilizer. The results suggest that integrated nutrient management (100 % NPK + FYM) in conjunction with three irrigations maximized yield of wheat with concomitant improvement in ET and WUE under limited water availability.