B. Ehdaie

Domestication and Crop Physiology: Roots of Green-Revolution Wheat

Background and Aims Most plant scientists, in contrast to animal scientists, study only half the organism, namely above-ground stems, leaves, flowers and fruits, and neglect below-ground roots. Yet all acknowledge roots are important for anchorage, water and nutrient uptake, and presumably components of yield. This paper investigates the relationship between domestication, and the root systems of landraces, and the parents of early, mid- and late green-revolution bread wheat cultivars. It compares the root system of bread wheat and ‘Veery’-type wheat containing the 1RS translocation from rye. Methods Wheat germplasm was grown in large pots in sand culture in replicated experiments. This allowed roots to be washed free to study root characters. Key Results The three bread wheat parents of early green-revolution wheats have root biomass less than two-thirds the mean of some landrace wheats. Crossing early green-revolution wheat to an F2 of ‘Norin 10’ and ‘Brevor’, further reduced root biomass in mid-generation semi-dwarf and dwarf wheats. Later-generation semi-dwarf wheats show genetic variation for root biomass, but some exhibit further reduction in root size. This is so for some California and UK wheats. The wheat–rye translocation in ‘Kavkaz’ for the short arm of chromosome 1 (1RS) increased root biomass and branching in cultivars that contained it. Conclusions Root size of modern cultivars is small compared with that of landraces. Their root system may be too small for optimum uptake of water and nutrients and maximum grain yield. Optimum root size for grain yield has not been investigated in wheat or most crop plants. Use of 1RS and similar alien translocations may increase root biomass and grain yield significantly in irrigated and rain-fed conditions. Root characters may be integrated into components of yield analysis in wheat. Plant breeders may need to select directly for root characters.

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.

GENETIC VARIATION AND INTERRELATIONSHIPS OF AGRONOMIC CHARACTERS IN LANDRACES OF BREAD WHEAT FROM SOUTHEASTERN IRAN

There is renewed interest in wheat landraces as important sources of genetic variation for agronomic characters. Fifty-three pure lines of bread wheat (Triticum aestivum L.) derived from seven landraces collected from southeastern Iran were used to estimate genetic variation and heritability for 13 developmental and quantitative characters. Path-analysis was used to partition the genetic correlations between grain yield and six grain yield-related traits. Mean values of landraces were also compared with three improved cultivars from California and Iran. Genotypic differences among the landraces and among the pure lines collected from the landraces were highly significant for all characters considered. Compared with the modern cultivars, the landrace genotypes were, on average, later in days to heading and taller than the cultivars but had lower values for number of grains per spike, 1000-grain weight, grain yield and harvest index. Some landrace genotypes were similar to the modern cultivars for grain yield. Moderate to high genetic variation was displayed by number of grains per spike, number of spikes per plant, 1000-grain weight, and harvest index. The heritability estimates ranged from 59% for grain yield to 99% for days to anthesis. Expected genetic advance (as % of the mean) was ≈34% for number of spikes per plant, number of grains per spike, and 1000-grain weight. Days to heading and to anthesis correlated positively with number of spikes per plant, shoot biomass, and straw biomass but negatively with number of grains per spike and harvest index. The strong direct effect of number of spikes per plant on grain yield was completely counterbalanced by its indirect negative effects via number of grains per spike and 1000-grain weight. Number of grains per spike and 1000-grain weight were positively correlated with grain yield, and they had large direct effects. These two characters, however, were negatively correlated and exhibited a substantial counterbalance effect via one another and via number of spikes per plant. The landraces could be improved by intercrossing the promising genotypes identified in this study, with simultaneous selection for earliness, fewer number of spikes per plant, greater number of grains per spike and heavier grains. For further improvement, crossing programs between the landraces and introduced germplasm may be necessary.

Dissection of QTL effects for root traits using a chromosome arm-specific mapping population in bread wheat

A high-resolution chromosome arm-specific mapping population was used in an attempt to locate/detect gene(s)/QTL for different root traits on the short arm of rye chromosome 1 (1RS) in bread wheat. This population consisted of induced homoeologous recombinants of 1RS with 1BS, each originating from a different crossover event and distinct from all other recombinants in the proportions of rye and wheat chromatin present. It provides a simple and powerful approach to detect even small QTL effects using fewer progeny. A promising empirical Bayes method was applied to estimate additive and epistatic effects for all possible marker pairs simultaneously in a single model. This method has an advantage for QTL analysis in minimizing the error variance and detecting interaction effects between loci with no main effect. A total of 15 QTL effects, 6 additive and 9 epistatic, were detected for different traits of root length and root weight in 1RS wheat. Epistatic interactions were further partitioned into inter-genomic (wheat and rye alleles) and intra-genomic (rye–rye or wheat–wheat alleles) interactions affecting various root traits. Four common regions were identified involving all the QTL for root traits. Two regions carried QTL for almost all the root traits and were responsible for all the epistatic interactions. Evidence for inter-genomic interactions is provided. Comparison of mean values supported the QTL detection.

Stomatal frequency and size differentiate ploidy levels in Aegilops neglecta

Aegilops neglecta Req. ex Bertol. is a forage goatgrass that has tetraploid and hexaploid forms. No morphological trait is known to distinguish between the two forms. Accessions of Ae. neglecta representing diverse germplasm were characterized for stomatal frequency and size on the adaxial surface of leaves. Stomatal frequency among tillers within a plant and among leaves on a tiller was measured. Significant variation was found in stomatal frequency between the ploidy levels for basal leaves 2 and 3 and the penultimate and flag leaves. Mean stomatal frequency for basal leaves ranged from 50.24 to 54.80 per mm2 for tetraploid and from 39.24 to 40.50 per mm2 for hexaploid accessions. For upper leaves, it varied from 52.88 to 53.50 per mm2 for tetraploid and from 46.12 to 46.40 per mm2 for hexaploid accessions. The two ploidy forms also exhibited clear differences for stomatal size on the basal and upper leaves of the main tiller. Mean stomatal size for the basal leaves ranged from 41.42 to 46.09 μm for tetraploid and from 53.59 to 54.97 μm for hexaploid accessions. For upper leaves, stomatal size varied from 47.29 to 53.29 μm in tetraploid and from 64.76 to 66.84 μm for hexaploid accessions. None of the individual accession ranges overlapped between the ploidy levels. Stomatal frequency and size were highly negatively correlated. Increased ploidy in Ae. neglecta has resulted in fewer but larger stomata per unit leaf area. Stomatal size and/or frequency on the adaxial surface of leaves could be used to distinguish tetraploid from hexaploid cytotypes in Ae. neglecta in fresh material from Turkey collected in the field.

Genetic analysis of carbon isotope discrimination and agronomic characters in a bread wheat cross

Carbon isotope discrimination (Δ) has been suggested as a selection criterion to improve transpiration efficiency (W) in bread wheat (Triticum aestivum L.). Cultivars ‘Chinese Spring’ with low A (high W) and ‘Yecora Rojo’ with high Δ (low W) were crossed to develop F1, F2, BC1, and BC2 populations for genetic analysis of Δ and other agronomic characters under well-watered (wet) and water-stressed (dry) field conditions. Significant variation was observed among the generations for Δ only under the wet environment. Generation x irrigation interactions were not significant for Δ. Generation means analysis indicated that additive gene action is of primary importance in the expression of Δ under nonstress conditions. Dominance gene action was also detected for Δ, and the direction of dominance was toward higher values of Δ. The broad-sense and the narrow-sense heritabilities for Δ were 61 % and 57% under the wet conditions, but were 48% and 12% under the draughted conditions, respectively. The narrow-sense heritabilities for grain yield, above-ground dry matter, and harvest index were 36%, 39%, and 60% under the wet conditions and 21%, 44%, and 20% under dry conditions, respectively. The significant additive genetic variation and moderate estimate of the narrow-sense heritability observed for Δ indicated that selection under wet environments should be effective in changing Δ in spring bread wheat.

Genetic diversity in populations of wild diploid wheat Triticum urartu Tum. ex. Gandil. revealed by isozyme markers

Genetic variation and its distribution within and among 23 populations of Triticum urartu collected from Syria, Lebanon, Turkey, Armenia, and Iran was estimated using isozyme markers at eight polymorphic loci. The number of alleles per locus (A= 1.21), percentage polymorphic loci (P= 20.1%), and mean gene diversity (He= 0.024) were relatively low. In a population from Lebanon, a high number of alleles per locus (A= 2.13) and percentage polymorphic loci (P= 87.5%) was found. On average, genetic variation among populations (GST= 0.407) was smaller than within-population variation (0.593). However, different patterns of genetic structure were found among various geographic regions. Interpopulation variation was highest for the Iranian populations (0.89) followed by the Turkish populations (0.66). A reverse pattern was observed for the Syrian (0.11) and for the Lebanese (0.13) populations. The Armenian populations exhibited similar interpopulation and within-population variation. Principal component and cluster analyses resulted in distinct grouping of the geographically proximal populations, with the exception of the two Iranian populations. The Turkish populations were different from the neighboring Armenian populations compared to other countries. The populations from southern Syria and those from Lebanon also exhibited a high degree of genetic diversity. The two most heterozygous loci, Mdh-2 and Pgi-2, separated the populations along the first and second principal components, respectively. Most of the rare alleles were scattered sporadically throughout the geographic regions. Rare alleles with high frequencies were found in the Turkish and Armenian populations. These results indicated that different geographic regions require specific sampling procedures in order to capture the range of genetic variation observed in T. urartu populations.

SOWING DATE AND NITROGEN INPUT INFLUENCE NITROGEN-USE EFFICIENCY IN SPRING BREAD AND DURUM WHEAT GENOTYPES

Environmental and economic considerations require the effective use of nitrogen (N) fertilizer in wheat production. This study evaluated genetic variation in nitrogen-use efficiency (NUE) for grain yield (NUEY) and protein (NUEP), and their components using nine bread wheat (Triticum aestivum L.) genotypes, five durum wheat (T. turgidum, var. durum Desf.) genotypes and a triticale (× Triticosecale Wittmak) genotype under 105 (N1) and 170 (N2) kg N ha −1 sown in October (early) December (optimum),and February (late) 1993 at the Moreno Valley Field Station of University of California, Riverside. Late and early sowing decreased mean grain yield and harvest index, respectively. Mean NUEY and NUEP was greater at optimum and early than at late sowing, but their components, N-uptake efficiency, N-production efficiency, and N-partitioning efficiency exhibited different patterns. Greater N supply, on average,increased shoot biomass by 29%, grain yield by 16%, and percent protein by 5%, but decreased harvest index by 10%, NUEY by 28%, NUEP by 26%, N-uptake efficiency by 18%, N-production efficiency by 12%, and N-partitioning efficiency by 8%. However, genotypes responded differently to sowing date and N rate for different characters. None of the genotypes examined consistently performed satisfactorily over different sowing dates and all, but one at early sowing, had similar or reduced NUEY and NUEP at N2 than at N1. Genotypic variation in modern bread and durum wheat was relatively small for NUEY and NUEP. Nitrogen-uptake efficiency was the most important component of both NUEY and NUEP. Correlation coefficients between NUEY and NUEP ranged from 0.83 to 0.98, whereas those between the components were nonsignificant in all cases, except one. Selection for N-uptake efficiency under low N input should improve both NUEY and NUEP in wheat. Wheat genotypes with desirable characteristics under low N input were identified for early, optimum, and late sowing.

Chromosomal location of genes influencing plant characters and evapotranspiration efficiency in bread wheat

Little is known of genes that influence root development and drought resistance in bread wheat. The evapotranspiration efficiency (ETE = ratio of vegetative dry weight to total water used) of spring bread wheat tall landrace ‘Chinese Spring’ is relatively high. We used 42 ditelosomic and dimonotelosomic lines of Chinese Spring to identify chromosome arms that influence plant characters and ETE. Multiple regression analyses indicated that 96% of the variation observed in ETE was explained by variation in vegetative dry weight and total water used. Variation in plant height, number of spikes (tillers), root dry weight and shoot dry weight (excluding grains) together explained 88% of the variation observed in plant vegetative dry weight. Chromosome arms involved in expression of days to heading and maturity, plant height, number of spikes, root dry weight, shoot dry weight, number of grains, grain weight, and carbon isotope discrimination (Δ) were identified. Specifically, both arms of chromosome 2A, the long arm of chromosome 2B, and the short arm of chromosome 2D might carry genes with positive effects on number of spikes, root dry weight, and shoot dry weight. None of the aneuploids produced grain yield greater than Chinese Spring. The short arms of chromosomes 6A and 4D might carry genes that suppress Δ. Chromosome 1D might carry genes that increase relative water loss. The chromosome arms belonging to homoeologous group 2 might carry genes with positive effects on ETE. The genetic basis of ETE in modern wheats could be broadened by substituting specific chromosome arms of landrace wheats carrying desired characters into their genomes.

Dosage effect of the short arm of chromosome 1 of rye on root morphology and anatomy in bread wheat

The spontaneous translocation of the short arm of chromosome 1 of rye (1RS) in bread wheat is associated with higher root biomass and grain yield. Recent studies have confirmed the presence of QTL for different root morphological traits on the 1RS arm in bread wheat. This study was conducted to address two questions in wheat root genetics. First, does the presence of the 1RS arm in bread wheat affect its root anatomy? Second, how does root morphology and anatomy of bread wheat respond to different dosages of 1RS? Near-isogenic plants with a different number (0 to 4 dosages) of 1RS translocations were studied for root morphology and anatomy. The F1 hybrid, with single doses of the 1RS and 1AS arms, showed heterosis for root and shoot biomass. In other genotypes, with 0, 2, or 4 doses of 1RS, root biomass was incremental with the increase in the dosage of 1RS in bread wheat. This study also provided evidence of the presence of gene(s) influencing root xylem vessel number, size, and distribution in bread wheat. It was found that root vasculature follows a specific developmental pattern along the length of the tap root and 1RS dosage tends to affect the transitions differentially in different positions. This study indicated that the inherent differences in root morphology and anatomy of different 1RS lines may be advantageous compared to normal bread wheat to survive under stress conditions.