Relationship between Carbon Isotope Discrimination, Ash Content and Grain Yield in Wheat in the Peninsular Zone of India

Carbon isotope discrimination (Δ) and ash content (m_a) have been proposed as indirect selection criteria for grain yield in wheat. The associations between Δ, m_a and grain yield were found, however, to depend highly on the environmental conditions, the organ sampled and the time of sampling. In this study, carried out in the warm conditions of the Peninsular Zone of India, the relationship between Δ, m_a and yield was studied in 30 bread and durum wheat cultivars under residual soil moisture stress (RSMS), post‐anthesis water stress (PAWS) and well‐watered (WW) conditions. Both Δ and m_a were analysed in young seedlings (four‐leaf stage), leaves at anthesis and grain at maturity. Ash content was also evaluated in leaves at booting stage and maturity. Grain Δ was lower under PAWS and RSMS than under WW, while seedling and leaf Δ did not significantly differ among water regimes. Grain yield was positively correlated to grain Δ under PAWS and negatively correlated to grain m_a under RSMS. A significant positive correlation was noted under RSMS and WW treatments between m_aL_m and grain yield. Ash content in leaf at maturity consequently appears to be a useful indirect selection criterion in environments where Δ does not show any correlation with yield. The results highlight the potential of Δ and m_a as indirect selection criteria for wheat yield in the conditions of the Peninsular Zone of India.     

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.     

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.     

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.     

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.     

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.     

Bread and Durum Wheat under Heat Stress: A Comparative Study on the Photosynthetic Performance

The photosynthetic responses to heat stress, during grain filling, in four genotypes of Triticum aestivum L. (Sever and Golia) and Triticum turgidum subsp. durum (Acalou and TE 9306), chosen according to its genetic background diversity, were investigated. All wheat genotypes (excepting Golia) showed synergistic trends implicating the internal CO₂ concentration, net photosynthesis and stomatal conductance. Additionally, the modifications of net photosynthesis were associated with changes in stomatal control. Chlorophyll a fluorescence parameters (minimal fluorescence, maximal and variable fluorescence, intrinsic efficiency of PSII in darkness, non‐photochemical quenching, photochemical quenching and energy‐dependent chlorophyll fluorescence quenching) further pointed heat protective mechanisms, implicating F_v/F_m stabilization (i.e. maintaining the efficiency of PS II) and electron transport rate preservation. It is concluded that, comparatively to bread wheat, the photosynthetic performance of durum wheat is more tolerant to heat stress, as stomatal conductance and transpiration are less affected.     

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.     

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.     

Flag Leaf Variation and the Analysis of Diversity in Durum Wheat

Flag leaf in wheat (Triticum spp.) is a major source of photosynthate for the developing grain and its size contributes to interplant competition. Flag leaf length, width, and area in a sample of 778 entries of durum wheat (T. turgidum L. durum group) germplasm obtained from the U.S. National Small Grains Collection showed considerable phenotypic variation. Correlations between flag leaf and eight spike quantitative characters showed significant values greater than 0.30 for kernel weight per spike with all three flag leaf measurements and for awn length with flag leaf length and area. Larger flag leaf area appeared to be associated with accessions having a history of some deliberate plant breeding for high grain yield. A multivariate approach was used to assess how discrimination among 26 countries of origin obtained with spike characters in a previous study was affected by reducing the sample size from about 3,000 to 735 and by the addition of flag leaf measurements to the analysis. The reduced sample size had little impact on the results of the previous analysis on the whole collection of about 3,000 entries. The addition of flag leaf characters to the analysis caused a rearrangement in the clusters of countries, mostly from Asia, where less germplasm modified by plant breeding was included in the collection. Five clusters were identified: (1) 11 Mediterranean basin countries, (2) USA, UdSSR, and Yugoslavia, (3) four Near East Countries and Bulgaria, (4) three Middle East countries and Greece, India and Ethiopia, and (5) Afghanistan. Phenotypic differentiation of durum wheat, and probably many other crops, on a geographic basis is more pronounced in areas where plant breeding has had little impact.     

Carbon Isotope Discrimination and Productivity in Field-Grown Barley Genotypes

Five barley genotypes of different origins were used to examine the relationships between carbon isotope discrimination (Δ) and grain yield, yield components, dry aerial biomass and harvest index under rainfed and irrigated Mediterranean conditions. High positive correlations were found between Δ and grain yield and harvest index under both water conditions, suggesting that Δ may represent efficiency of dry matter partitioning to the grain. This hypothesis was also supported by the positive correlation found between Δ and thousand-grain weight when all the data were considered. The two drought-tolerant genotypes LM2887 and Tadmor presented the smallest difference between the two treatments for traits related to productivity as well as the highest Δ values. These results highlight the potential of Δ as a selection criterion in barley breeding in Mediterranean regions. They also encourage the study of the genetics of Δ and the identification of molecular markers linked to its variation in available progenies derived from crosses between the genotypes used in the present paper.     

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.     

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.     

Grain Yield and Quality of Durum Wheat Landraces in a Dry Mediterranean Region of Northern Syria

Twenty-two durum wheat landraces of various origins were evaluated for performance and stability of grain yield, protein content and gluten strength (SDS sedimentation volume) at two sites of Northern Syria situated in the drought-prone part of the local durum wheat belt. Yield was recorded in three seasons, while protein content and SDS volume were assessed in two seasons only. A high “entry X environment” interaction was observed for all traits, which was significantly attributable to heterogeneity of entry regressions on the environmental mean for grain yield only. Compared to the recently-bred, widely-grown variety ‘Sham I’, some landraces showed somewhat similar yield responses, whereas others proved more stable without being significantly lower yielding. Significant variation was found for SDS volume, though only few of the best-ranking landraces for this trait also possessed an acceptable yield level. The entries did not significantly differ for protein content. The broad sense heritability estimates obtained for grain yield (0.59) and SDS volume (0.89) highlighted the interest in some of the landraces examined for improving grain yield and quality in the region.     

Performance of Selected Strains of 'Farro' (Triticum monococcum L., Triticum dicoccum Schübler, Triticum Spelta L.) and Durum Wheat (Triticum Durum Desf. cv. Trinakria) in the Difficult Flat Environment of Southern Italy

The aim of this study was to evaluate the performance of three species of unhulled-wheat (Triticum monococcum L., Triticum dicoccum Schubler and Triticum spelta L.) with respect to durum wheat (Triticum durum Desf. cv. Trinakria) in southern Italy's flat environment.
As far as yield is concerned, the mean yield was significantly different among species. Durum wheat yielded 16.1 %, 37.6% and 69.5 % more than dicoccum, spelta and monococcum species, respectively.
Among unhulled-wheat species, only Triticum dicoccum showed some similarity with durum wheat, such as one-thousand kernels weight, plant height and number of spikes per m².
Among the factors studied, the days to reach heading represented the factor that mostly influenced yield     

Long-term Low Radiation Decreases Leaf Photosynthesis, Photochemical Efficiency and Grain Yield in Winter Wheat

Low radiation reduces wheat grain yield in tree-crop intercropping systems in the major wheat planting area of China. Here, two winter wheat ( Triticum aestivum L) cultivars, Yangmai 158 (shading tolerant) and Yangmai 11 (shading sensitive), were shaded from jointing to maturity to evaluate the impact of low radiation on crop growth, photosynthesis and yield. Grain yield losses and leaf area index (LAI) reduction were less than the reduction in solar radiation under both shading treatment in both cultivars. Compared with the control (S0), grain yield only reduced 6.4 % and 9.9 % under 22 % shading treatment (S1), while 16.2 % and 25.8 % under 33 % shading (S2) in Yangmai 158 and Yangmai 11 respectively. The reduction in LAI was 6.0 % and 9.2 % (S1), and 18.2 % and 22.2 % (S2) in Yangmai 158 and Yangmai 11 respectively. However, decline in canopy apparent photosynthetic rate (CAP) was 15.0–22.9 % (S1) and 29.5–49.6 % (S2), which was consistent with the reduction in radiation. The reduction in LAI was partially compensated by increases in the fraction of the top and bottom leaf area to the total leaf area, which facilitated to intercept more solar radiation by the canopy. The decrease in photosynthetic rate ( Pn ) of flag leaf was partially compensated by the increase in Pn of the third leaf from the top. In addition, an inconsistency between the low Pn and the high Chl content in flag leaf was observed at 30 DAA. This could be explained that more excitation energy was dispersed via the non-photochemical approaches in the photosystem II (PSII) of flag leaf after long-term shading.