Grain zinc, iron and protein concentrations and zinc-efficiency in wild emmer wheat under contrasting irrigation regimes

Micronutrient malnutrition, and particularly deficiency in zinc (Zn) and iron (Fe), afflicts over three billion people worldwide, and nearly half of the world’s cereal-growing area is affected by soil Zn deficiency. Wild emmer wheat [Triticum turgidum ssp. dicoccoides (Körn.) Thell.], the progenitor of domesticated durum wheat and bread wheat, offers a valuable source of economically important genetic diversity including grain mineral concentrations. Twenty two wild emmer wheat accessions, representing a wide range of drought resistance capacity, as well as two durum wheat cultivars were examined under two contrasting irrigation regimes (well-watered control and water-limited), for grain yield, total biomass production and grain Zn, Fe and protein concentrations. The wild emmer accessions exhibited high genetic diversity for yield and grain Zn, Fe and protein concentrations under both irrigation regimes, with a considerable potential for improvement of the cultivated wheat. Grain Zn, Fe and protein concentrations were positively correlated with one another. Although irrigation regime significantly affected ranking of genotypes, a few wild emmer accessions were identified for their advantage over durum wheat, having consistently higher grain Zn (e.g., 125 mg kg^?1), Fe (85 mg kg^?1) and protein (250 g kg^?1) concentrations and high yield capacity. Plants grown from seeds originated from both irrigation regimes were also examined for Zn efficiency (Zn deficiency tolerance) on a Zn-deficient calcareous soil. Zinc efficiency, expressed as the ratio of shoot dry matter production under Zn deficiency to Zn fertilization, showed large genetic variation among the genotypes tested. The source of seeds from maternal plants grown under both irrigation regimes had very little effect on Zn efficiency. Several wild emmer accessions revealed combination of high Zn efficiency and drought stress resistance. The results indicate high genetic potential of wild emmer wheat to improve grain Zn, Fe and protein concentrations, Zn deficiency tolerance and drought resistance in cultivated wheat.     

Exploration of Genetic Pattern of Phenological Traits in Wheat (<i>Triticum aestivum</i> L.) under Drought Stress

Drought is the major detrimental environmental factor for wheat (Triticum aestivum L.) production. The exploration of genetic patterns underlying drought tolerance is of great significance. Here we report the gene actions controlling the phenological traits using the line × tester model studying 27 crosses and 12 parents under normal irrigation and drought conditions. The results interpreted via multiple analysis (mean performance, correlations, principal component, genetic analysis, heterotic and heterobeltiotic potential) disclosed highly significant differences among germplasm. The phenological waxiness traits (glume, boom, and sheath) were strongly interlinked. Flag leaf area exhibits a positive association with peduncle and spike length under drought. The growing degree days (heat-units) greatly influence spikelets and grains per spike, however, the grain yield/plant was significantly reduced (17.44 g to 13.25 g) under drought. The principal components based on eigenvalue indicated significant PCs (first-seven) accounted for 79.9% and 73.9% of total variability under normal irrigation and drought, respectively. The investigated yield traits showed complex genetic behaviour. The genetic advance confronted a moderate to high heritability for spikelets/spike and grain yield/plant. The traits conditioned by dominant genetic effects in normal irrigation were inversely controlled by additive genetic effects under drought and vice versa. The magnitude of dominance effects for phenological and yield traits, i.e., leaf twist, auricle hairiness, grain yield/plant, spikelets, and grains/spike suggests that selection by the pedigree method is appropriate for improving these traits under normal irrigation conditions and could serve as an indirect selection index for improving yield-oriented traits in wheat populations for drought tolerance. However, the phenotypic selection could be more than effective for traits conditioned by additive genetic effects under drought. We suggest five significant cross combinations based on heterotic and heterobeltiotic potential of wheat genotypes for improved yield and enhanced biological production of wheat in advanced generations under drought.     

Response of wheat restricted‐tillering and vigorous growth traits to variables of climate change

The response of wheat to the variables of climate change includes elevated CO₂, high temperature, and drought which vary according to the levels of each variable and genotype. Independently, elevated CO₂, high temperature, and terminal drought affect wheat biomass and grain yield, but the interactive effects of these three variables are not well known. The aim of this study was to determine the effects of elevated CO₂ when combined with high temperature and terminal drought on the high‐yielding traits of restricted‐tillering and vigorous growth. It was hypothesized that elevated CO₂ alone, rather than combined with high temperature, ameliorates the effects of terminal drought on wheat biomass and grain yield. It was also hypothesized that wheat genotypes with more sink capacity (e.g. high‐tillering capacity and leaf area) have more grain yield under combined elevated CO₂, high temperature, and terminal drought. Two pairs of sister lines with contrasting tillering and vigorous growth were grown in poly‐tunnels in a four‐factor completely randomized split‐plot design with elevated CO₂ (700 µL L^?1), high day time temperature (3 °C above ambient), and drought (induced from anthesis) in all combinations to test whether elevated CO₂ ameliorates the effects of high temperature and terminal drought on biomass accumulation and grain yield. For biomass and grain yield, only main effects for climate change variables were significant. Elevated CO₂ significantly increased grain yield by 24–35% in all four lines and terminal drought significantly reduced grain yield by 16–17% in all four lines, while high temperature (3 °C above the ambient) had no significant effect. A trade‐off between yield components limited grain yield in lines with greater sink capacity (free‐tillering lines). This response suggests that any positive response to predicted changes in climate will not overcome the limitations imposed by the trade‐off in yield components.     

Salinity and drought interaction in wheat (Triticum aestivum L.) is affected by the genotype and plant growth stage

Salinity and drought are important agro-environmental problems occurring separately as well as together with the combined occurrence increasing with time due to climate change. Screening of bread wheat genotypes against salinity or drought alone is common; however, little information is available on the response of wheat genotypes to a combination of these stresses. This study investigates the response of a salt-resistant (SARC-1) and a salt-sensitive (7-Cerros) wheat genotype to drought at different growth stages under non-saline (ECe 2.1 dS m^?1) and saline soil (ECe 15 dS m^?1) conditions. Drought was applied by withholding water for 21 days at a particular growth stage viz. tillering, booting, and grain filling stages. At booting stage measurements regarding water relations, leaf ionic composition and photosynthetic attributes were made. At maturity grain yield and different yield, components were recorded. Salinity and drought significantly decreased grain yield and different yield components with a higher decrease in the case of combined stress of salinity × drought. The complete drought treatment (drought at tillering + booting + grain filling stages) was most harmful for wheat followed by drought at booting stage and grain filling–tillering stages, respectively. The salt-resistant wheat genotype SARC-1 performed better than the salt-sensitive genotype 7-Cerros in different stress treatments. A decrease in the water and turgor potentials, photosynthetic and transpiration rates, stomatal conductance, leaf K⁺, and increased leaf Na⁺ were the apparent causes of growth and yield reduction of bread wheat due to salinity, drought, and salinity × drought.     

A proteomics view on the role of drought-induced senescence and oxidative stress defense in enhanced stem reserves remobilization in wheat

Drought is one of the major factors limiting the yield of wheat (Triticum aestivum L.) particularly during grain filling. Under terminal drought condition, remobilization of pre-stored carbohydrates in wheat stem to grain has a major contribution in yield. To determine the molecular mechanism of stem reserve utilization under drought condition, we compared stem proteome patterns of two contrasting wheat landraces (N49 and N14) under a progressive post-anthesis drought stress, during which period N49 peduncle showed remarkably higher stem reserves remobilization efficiency compared to N14. Out of 830 protein spots reproducibly detected and analyzed on two-dimensional electrophoresis gels, 135 spots showed significant changes in at least one landrace. The highest number of differentially expressed proteins was observed in landrace N49 at 20days after anthesis when active remobilization of dry matter was observed, suggesting a possible involvement of these proteins in effective stem reserve remobilization of N49. The identification of 82 of differentially expressed proteins using mass spectrometry revealed a coordinated expression of proteins involved in leaf senescence, oxidative stress defense, signal transduction, metabolisms and photosynthesis which might enable N49 to efficiently remobilized its stem reserves compared to N14. The up-regulation of several senescence-associated proteins and breakdown of photosynthetic proteins in N49 might reflect the fact that N49 increased carbon remobilization from the stem to the grains by enhancing senescence. Furthermore, the up-regulation of several oxidative stress defense proteins in N49 might suggest a more effective protection against oxidative stress during senescence in order to protect stem cells from premature cell death. Our results suggest that wheat plant might response to soil drying by efficiently remobilize assimilates from stem to grain through coordinated gene expression.     

Proteomic response of barley leaves to salinity

Drought and salinity stresses are adverse environmental factors that affect crop growth and yield. Proteomic analysis offers a new approach to identify a broad spectrum of genes that are expressed in living system. We applied this technique to investigate protein changes that were induced by salinity in barley genotypes (Hordeum vulgare L.), Afzal, as a salt-tolerant genotype and L-527, as a salt-sensitive genotype. The seeds of two genotypes were sown in pot under controlled condition of greenhouse, using a factorial experiment based on a randomized complete block design with three replications. Salt stress was imposed at seedling stage and leaves were collected from control and salt-stressed plant. The Na⁺ and K⁺ concentrations in leaves changed significantly in response to short-term stress. About 850 spots were reproducibly detected and analyzed on 2-DE gels. Of these, 117 proteins showed significant change under salinity condition in at least one of the genotypes. Mass spectrometry analysis using MALDI-TOF/TOF led to the identification some proteins involved in several salt responsive mechanisms which may increase plant adaptation to salt stress including higher constitutive expression level and upregulation of antioxidant, upregulation of protein involved in signal transduction, protein biosynthesis, ATP generation and photosynthesis. These findings may enhance our understanding of plant molecular response to salinity.     

Genetic diversity of bread wheat genotypes in Iran for some nutritional value and baking quality traits

Genetic variation among 78 irrigated bread wheat genotypes was studied for their nutritional value and baking quality traits as well as some agronomic traits. The experiment was conducted in a randomized complete block design with three replicates under normal and terminal drought stress conditions in Kermanshah, Iran during 2012–2013 cropping season. The results of combined ANOVA indicated highly significant genotypic differences for all traits. All studied traits except grain yield, hectoliter weight and grain fiber content were significantly affected by genotype × environment interaction. Drought stress reduced grain yield, thousand kernel weight, gluten index, grain starch content and hectoliter weight and slightly promoted grain protein and fiber contents, falling number, total gluten and ratio of wet gluten to grain protein content. Grain yield by 31.66% and falling number by 9.20% attained the highest decrease and increase due to drought stress. There were negative and significant correlations among grain yield with grain protein and fiber contents under both conditions. Results of cluster analysis showed that newer genotypes had more grain yield and gluten index than older ones, but instead, they had the lower grain protein and fiber contents. It is thought that wheat breeders have bred cultivars with high grain yield, low protein content, and improved bread-making attributes during last seven decades. While older genotypes indicated significantly higher protein contents, and some of them had higher gluten index. We concluded from this study that it is imperative for breeders to pay more attention to improve qualitative traits coordinated to grain yield.     

Chlorophyll a fluorescence during flag leaf senescence of field-grown winter wheat plants under drought conditions

Improving wheat grain yield plays a significant role in ensuring global food security. Wheat production could be increased by the genetic improvement of wheat genotypes where delayed senescence with enhanced post-anthesis capacity and staygreen traits could have an important role. In this study, chlorophyll a fluorescence (ChlF) rise kinetics from the early until late senescence of flag leaves, grain yield and other agro-morphological characteristics were compared for three winter wheat advanced lines (Osk.4.312/10-18, Osk.4.330/6-18 and Osk.4.354/12-18) under natural drought conditions. The differences between lines were observed when considering the heading date which was 1 and 4 days earlier for the line Osk.4.354/12-18, than lines Osk.4.312/10-18 and Osk.4.330/6-18, respectively. Furthermore, line Osk.4.354/12-18 had the highest test weight (kg hl⁻¹), while line Osk.4.330/6-18 showed a tendency of decreased grain yield, compared to the other two lines. Analysis of ChlF transients and several JIP-test parameters indicated that all three lines had a generally similar course of changes in the photosynthetic performance of flag leaves during senescence under drought conditions. However, at the point when a decrease in photosynthetic performance was initiated, it was slightly less intensive in line Osk.4.354/12-18 accompanied by longer preservation of functionality and connectivity of PSII units, than in the other two lines, which contributed to its better agronomical performance. These results indicated that even delicate variations in the functioning of the photosynthetic apparatus of the flag leaf during grain filling were agronomically important, especially when plants were exposed to drought stress, and could be used to differentiate otherwise similar wheat genotypes. Even small genotype-specific differences in the photosynthetic performance of senescing flag leaves, along with earlier heading dates, could assist in the selection of genotypes with a better ability to cope with unfavourable environmental conditions.     

Gene action of canopy temperature in bread wheat under diverse environments

While canopy temperature (CT) shows a strong and reliable association with yield under drought and heat stress and is used in wheat breeding to select for yield, little is known of its genetic control. The objective of this study was to determine the gene action controlling CT in five wheat populations grown in diverse environments (heat, drought, and well-irrigated conditions). CT showed negative phenotypic correlations with grain yield under drought and well-irrigated environments. Additive × additive effects were most prevalent and significant for all crosses and environments. Dominance and dominance × dominance gene actions were also found, though the significance and direction was specific for each environment and genotypic cross. The use of CT as a selection criterion to improve tolerance to drought was supported by its significant association with grain yield and the genotype differences observed between cultivars. Our results indicated that genetic gains for CT in wheat could be achieved through conventional breeding. However, given some dominance and epistatic effects, it would be necessary to delay the selection process until the frequency of heterozygous loci within families is reduced.     

Variation in the holm oak leaf proteome at different plant developmental stages, between provenances and in response to drought stress

Major proteins of the holm oak leaf proteome have been previously identified using a combination of 2-DE, MS analysis and BLAST similarity search (Jorge et al., Proteomics 2005, 5, 222-234). That study, conducted with field samples from mature trees, revealed the existence of a great variability in the 2-DE protein map, with qualitative as well as quantitative changes, both analytical and biological. A similar study has been carried out with 2-year-old seedlings to analyze and study: (i) changes in the 2-DE protein profile at different tree developmental stages; (ii) the 2-DE protein map variability between three different Spanish provenances; and (iii) variations in the 2-DE protein profile in response to drought stress. Although the protein profile of leaves from seedlings and mature trees was fairly similar, the biological variance found was lower in the former. In the present study, new proteins have been identified. At least four different protein spots differentiated Spanish provenances, two of them identified as an ATP synthase alpha chain, and a 2,3-bisphosphoglycerate-independent phosphoglycerate mutase. Fourteen different protein spots were qualitatively variable between well-watered and drought-stressed seedlings, with some of them corresponding to enzymes of carbohydrate and protein metabolism. Data presented indicated the mobilization of storage proteins and carbohydrates, as well as photosynthesis inhibition under drought conditions.     

Proteomic analysis of wheat embryos with 2-DE and liquid-phase chromatography (ProteomeLab PF-2D) — A wider perspective of the proteome

Cereal embryos are a model system to study desiccation tolerance due to their ability to survive extreme water loss during late embryogenesis. To identify proteins accumulating in mature embryos which can be used as potential markers for dehydration tolerance, we compared the embryo proteome from two durum wheat genotypes (Triticum durum Desf.), Mahmoudi (salt and drought sensitive) and Om Rabia3 (salt and drought tolerant). Total protein extracts from wheat embryos were analyzed by using conventional 2-DE and ProteomeLab PF-2D. Analysis using different pH ranges showed that a larger number of fractions were solved by LC, than by conventional 2-DE at extreme technical pHs (pH 4.0–5.0 and pH 6.5–8.0). In contrast, at intermediate pHs (pH 5.0–6.5), resolution was better in 2-DE gels. The two techniques were used in parallel to analyze total protein extracts from embryos of the two wheat varieties. Several proteins belonging to the seed storage family, LEA-type/heat shock proteins, enzyme metabolism and radical scavengers were identified by analysis of trypsin digested peptides via mass spectrometry. These proteins accumulate in different amounts in embryos of tolerant and sensitive wheat varieties. The differences in expression pattern were further validated by enzyme activity, western blotting analysis and correlated with their corresponding mRNA expression by RT-PCR analyses for the corresponding protein. We suggest that the differential expression pattern could be used as a basis for a biochemical screen of tolerance/sensitivity to drought and salt stress in wheat embryos and germplasm.     

Comparative physiological response of wheat genotypes under terminal heat stress

Wheat (Triticum aestivum L.), a staple food crop, is of great commercial importance. Its production is restricted due to multiple environmental stresses. There are indications that the wheat production is consistently limited by terminal heat stress. Previous studies revealed a varied response of different wheat genotypes under heat stress conditions. Here, comparative physiological changes in wheat genotypes viz., DBW-140, Raj-3765, PBW-574, K-0-307 and HS-240 were evaluated under timely and late sown conditions in rabi season. We observed that heat stress dramatically affects chlorophyll content and leaf area index (LAI) in sensitive genotypes whereas proline and malondialdehyde (MDA) content were higher in tolerant genotypes under late sown conditions. Further, the heat susceptibility index (HIS) for 1,000-grain weight, grain weight and grain yield of wheat genotypes viz., HS 240 and K-0-307 was highest as compared with DBW 140, Raj 3765 and PBW 574 genotypes. This finding suggests that wheat genotypes are found to differ in their ability to respond to heat, thereby tolerance, which could be useful as genetic stock to develop wheat tolerant varieties in breeding programs.     

河北省冬小麦丰产抗旱性表型鉴定指标分析

以河北省审定的85个冬小麦品种为材料,采用防雨棚春季干旱和露地灌溉2个处理,分别于开花期、成熟期调查株高等27个表型性状,分析了各表型性状与单株子粒产量的相关性。结果表明,单株成穗数等12个性状与单株子粒产量抗旱系数或抗旱指数呈显著或极显著相关;结合表型性状变异系数,明确了提高单株成穗数、穗粒数、灌溉条件下较长的旗叶长度和干旱条件下较短的旗叶长度是培育丰产抗旱小麦新品种的主攻方向;子粒比重、子粒长度及干旱条件下的结实率和每穗小穗数可作为河北省小麦种质资源丰产抗旱性的鉴定依据;河北省小麦品种丰产性高,而抗旱性尚需进一步改善。     

Genetics of pre-harvest sprouting resistance in a cross of Canadian adapted durum wheat genotypes

Severe losses attributable to pre-harvest sprouting (PHS) have been reported in Canada in recent years. The genetics of PHS resistance have been more extensively studied in hexaploid wheat and generally not using combinations of elite agronomic parents. The objective of our research was to understand the genetic nature of PHS resistance in an elite durum cross. A doubled haploid (DH) population and checks were phenotyped in replicated trials for grain yield and PHS traits over 3 years in western Canada. The response of intact spikes to sprouting conditions, sampled over two development time points, was measured in a rain simulation chamber. The DH population was genotyped with simple sequence repeat and Diversity Arrays Technology markers. Genotypes were a significant source of variation for grain yield and PHS resistance traits in each tested environment. Transgressive segregant DH genotypes were identified for grain yield and PHS resistance measurements. Low or no correlation was detected between grain yield and PHS, while correlation between PHS resistance measurements was moderate. The heritability of PHS resistance was moderate and higher than grain yield. Significant quantitative trait loci with small effect were detected on chromosomes 1A, 1B, 5B, 7A and 7B. Both parents contributed to the PHS resistance. Promising DH genotypes with high and stable grain yield as well as PHS resistance were identified, suggesting that grain yield and PHS can be improved simultaneously in elite genetic materials, and that these DH genotypes will be useful parental material for durum breeding programs.     

Wheat-Aegilops biuncialis amphiploids have efficient photosynthesis and biomass production during osmotic stress

Osmotic stress responses of water content, photosynthetic parameters and biomass production were investigated in wheat-Aegilops biuncialis amphiploids and in wheat genotypes to clarify whether they can use to improve the drought tolerance of bread wheat. A decrease in the osmotic pressure of the medium resulted in considerable water loss, stomatal closure and a decreased CO₂ assimilation rate for the wheat genotypes, while the changes in these parameters were moderate for the amphiploids. Maximal assimilation rate was maintained at high level even under severe osmotic stress in the amphiploids, while it decreased substantially in the wheat genotypes. Nevertheless, the effective quantum yield of PS II was higher and the quantum yield of non-photochemical quenching of PS II and PS I was lower for the amphiploids than for the wheat cultivars. Parallel with this, higher cyclic electron flow was detected in wheat than in the amphiploids. The elevated photosynthetic activity of amphiploids under osmotic stress conditions was manifested in higher biomass production by roots and shoots as compared to wheat genotypes. These results indicate that the drought-tolerant traits of Ae. biuncialis can be manifested in the wheat genetic background and these amphiploids are suitable genetic materials for improving drought tolerance of wheat.     

Proteomic Analysis of Osmotic Stress-Responsive Proteins in Sugarcane Leaves

Osmotic stress-related proteins in sugarcane were identified using proteomics approach based on two-dimensional polyacrylamide gel electrophoresis (2-DE) and matrix-assisted laser desorption/ionization-time of flight mass spectrometry (MALDI-TOF-MS). Sugarcane settlings were subjected to osmotic stress in the nutrient solution containing 10% (w/v) PEG 6000 for 14 h. Total proteins were extracted from leaves, and separated by 2-DE. Four typical spots exhibited significant changes in PEG treatment compared to control, which were identified using MALDI-TOF-MS successfully. The drought inducible 22 kDa protein and Rubisco small subunit were up-regulated while isoflavone reductase-like (IRLs, related to antioxidant defense system) protein and delta chain of ATP synthase were down-regulated by the osmotic stress. Analysis of the results showed that the most differential proteins under osmotic stress were acidic, unstable and transmembrane proteins, enriched with hydrophobic amino acids such as leucine and alanine which are extremely important for structural stabilization of proteins by hydrophobic interaction. However, the drought inducible 22 kDa protein was a hydrophile and non-transmembrane protein enriched with glutamic acid. These results provide new insight into the part of regulatory mechanism of adaptations to osmotic stress through differential expression of specific proteins and implicate several previously unrecognized proteins to osmotic stress.