Ontogenetic analysis of yield components and yield stability of durum wheat in water-limited environments

Summary One main reason for the slow improvement of durum wheat in water-limited environments is the lack of clear understanding of the interrelationships among yield components and their compensatory changes under low and erratic moisture availability. Five cultivars, varying in many physiological attributes, were tested under different drought-stress conditions in field and greenhouse experiments. The cause-effect relationships of duration of vegetative period, duration of grain-filling period, number of spikes per m², kernels per spike, kernel weight and grain yield per m² were assessed. Furthermore, yield stability was evaluated. Yield reduction was largest under mid-season stress (58%), followed by terminal stress (30%) and early stress (22%). Cultivar Po was very sensitive to terminal stress.Path-coefficient analysis revealed a complex pattern of relationships among the six variables. An increase in vegetative period reduced the grain-filling period under all conditions. It increased number of kernels per spike under non-stress conditions. The direct effect of spikes per m² on grain yield was significantly positive. However, more spikes per m² resulted in fewer kernels per spike and a low kernel weight and, as a result, a negative relationship with grain yield under early stress. Grain-filling period had a strong influence on grain yield via kernel weight. Kernels per spike had the largest direct effect on grain yield. However, it was negatively correlated with kernel weight, especially under terminal stress. Grain yield heavily depended on kernels per spike under early stress and grain-filling period and kernels per spike under terminal stress.Variation in drought susceptibility index among cultivars was significant under early and terminal stress conditions, but not under mid-stress conditions. Yield potential and stability were not correlated for the different drought-stress conditions.Longer grain-filling period, increased number of kernels per spike and limited spike number per m² can be used as selection criteria for sustainable yield in water-limited environments.     

Resistance to stripe rust in Triticum turgidum,T. tauschii and their synthetic hexaploids

Resistance to stripe rust (caused by Puccinia striiformis Westend.) of 34 Triticum turgidum L. var.durum, 278 T. tauschii, and 267 synthetic hexaploid wheats (T. turgidum x T. tauschii) was evaluated at the seedling stage in the greenhouse and at the adult-plant stage at two field locations. Mexican pathotype 14E14 was used in all studies. Seedling resistance, expressed as low infection type, was present in all three species. One hundred and twenty-eight (46%) accessions of T. tauschii, 8 (23%) of T. turgidum and 31 (12%) of synthetic hexaploid wheats were highly resistant as seedlings. In the field tests, resistance was evaluated by estimating area under disease progress curve (AUDPC). Synthetic hexaploid wheats showed a wide range of variability for disease responses in both greenhouse and field tests, indicating the presence of a number of genes for resistance. In general, genotypes with seedling resistance were also found to be resistant as adult plants. Genotypes, which were susceptible or intermediate as seedlings but resistant as adult plants, were present in both T. turgidum and the synthetic hexaploids. Resistances from either T. turgidum or T. tauschii or both were identified in the synthetic hexaploids in this study. These new sources of resistance could be incorporated into cultivated hexaploid wheats to increase the existing gene pool of resistance to stripe rust.     

Effects of dwarfing genes on yield and yield components under irrigated and rainfed conditions in wheat (Triticum aestivum L.)

Summary Near-isogenic tall (no dwarfing gene), semidwarf (Rht1 or Rht2) and dwarf (Rht1 + Rht2 or Rht3) spring wheat lines were evaluated for yield and yield components under irrigated and rainfed conditions. Under irrigated conditions, the dwarf and the semidwarf lines exhibited a significant yield advantage over the tall lines. Under rainfed conditions, the semidwarf lines outyielded the tall as well as the dwarf lines. Percent yield reduction in response to drought stress was highest with the dwarfs and lowest with the tall lines. Dry matter production of the tall lines and that of the semidwarf lines did not differ significantly and both produced significantly more dry matter than the dwarf lines under irrigated as well as rainfed conditions. Plant height and kernel weight decreased with increasing degree of dwarfness while number of kernels per spikelet, harvest index and days to heading increased under both moisture regimes. The dwarfing genes did not have any significant influence on number of tillers/m² and spikelets per spike in either moisture regime.     

Applicability of Aegilops tauschii drought tolerance traits to breeding of hexaploid wheat

Few genes are available to develop drought-tolerant bread wheat (Triticum aestivum L.) cultivars. One way to enhance bread wheat’s genetic diversity would be to take advantage of the diversity of wild species by creating synthetic hexaploid wheat (SW) with the genomic constitution of bread wheat. In this study, we compared the expression of traits encoded at different ploidy levels and evaluated the applicability of Aegilops tauschii drought-related traits using 33 Ae. tauschii accessions along with their corresponding SW lines under well-watered and drought conditions. We found wide variation in Ae. tauschii, and even wider variation in the SW lines. Some SW lines were more drought-tolerant than the standard cultivar Cham 6. Aegilops tauschii from some regions gave better performing SW lines. The traits of Ae. tauschii were not significantly correlated with their corresponding SW lines, indicating that the traits expressed in wild diploid relatives of wheat may not predict the traits that will be expressed in SW lines derived from them. We suggest that, regardless of the adaptability and performance of the Ae. tauschii under drought, production of SW could probably result in genotypes with enhanced trait expression due to gene interactions, and that the traits of the synthetic should be evaluated in hexaploid level.     

Assessment of genetic diversity in synthetic hexaploid wheats and their Triticum dicoccum and Aegilops tauschii parents using AFLPs and agronomic traits

Synthetic hexaploid wheats are of interest to wheat breeding programs, especially for introducing new genes that confer resistance to biotic and abiotic stresses. A group of 54 synthetic hexaploid wheats derived from crosses between emmer wheat(Triticum dicoccum, source of the A and B genomes) and goat grass (Aegilops tauschii, D genome donor) were investigated for genetic diversity. Using the AFLP technique, dendrograms revealed clear grouping according to geographical origin for the T. dicoccum parents but no clear groups for the Ae. tauschii parents. The geographical clustering of the T. dicoccum parents was also reflected in the dendrogram of their derived synthetic hexaploids. Diversity of the T. dicoccum parents and their derived synthetic hexaploids was further evaluated by measuring 18morphological and agronomic traits on the plants. Clustering based on morphological and agronomic data also reflected geographical origin. However, comparison of genetic distances obtained from AFLP and agronomic data showed no correlation between the two diversity measurements. Nevertheless, similarities among major clusters with the two systems could be identified. Based on percentage of polymorphic markers, the synthetic hexaploids had a considerably higher level of AFLP diversity (39%) than normally observed in cultivated hexaploid wheat (12–21%). This suggests that synthetic hexaploid wheats can be used to introduce new genetic diversity into the bread wheat gene pool. This revised version was published online in July 2006 with corrections to the Cover Date.     

Effects of moisture stress on leaf appearance,tillering and other aspects of development in Triticum tauschii

Summary A glasshouse study was conducted to describe the dynamics of leaf and tiller appearance of four accessions of T. tauschii (Tt 04, Tt 17, Tt 65 and Tt 74) and to determine the influence of moisture stress (treatments were high and low moisture, imposed seven days after transplanting) on these and other aspects of development in this wild wheat.Under high moisture conditions, accessions differed greatly in flag leaf dimensions, culm length and seed number per spike, the values being lower in Tt 04 than in the other accessions. Low moisture strongly reduced values for these traits, with Tt 04 being least affected, but overall, there was no apparent association between the values obtained for these variables in the high moisture conditions and the effects of moisture stress. For three of the four accessions, final leaf number on the main culm was significantly lower in the low moisture treatment than in the respective control (P<0.05), but the differences between treatments (ca. 0.5 leaves or less) were very small. Maximum tiller number, on the other hand, was strongly reduced by low moisture, and initiation of tillering was inhibited until water was reapplied. There were no apparent after-effects of the moisture regime on the rate of subsequent tiller appearance.The four accessions differed in their leaf appearance rates, giving phyllochron values (117–142° Cd leaf^-1) within the range reported for hexaploid wheat. Low moisture tended to increase phyllochron, but in only one accession was this effect significant. Thus, depending on the accession, low moisture did not affect, or slightly decreased (by ca. 15–20%) the rate of leaf appearance. These effects were similar to those reported for cultivated wheat suggesting that there would be little scope for using these accessions of T. tauschii in breeding for stress tolerance.     

Genetics of morphological traits in wild wheat,Triticum turgidum var.dicoccoides

Summary The mode of inheritance, linkage groups, and chromosomal location of 23 morphological and 4 biochemical traits were characterized in the wild tetraploid emmer wheat,Triticum turgidum var.dicoccoides. These traits were described and their mode of inheritance was determined by their segregation in four F₂ populations derived from crosses between four var.dicoccoides accessions and a tetraploiddurum cultivar. Linkage groups among the genes encoding for these traits were determined or postulated, and their chromosomal location was deduced by linkage to previously located genes. The genetic control of the following traits was characterized and is first reported here: black keel; hairy leaf sheath; hairy auricles; hairy rachilla; hairy kernel brush; obtuse flag leaf; and curved neck/peduncle. The linkage data indicated that developmentally-related genes tended to occur in clusters.     

Selection for yield in small plots of spring wheat

Summary Four selection experiments were sown: single-row plots with single row-spacing (20 cm), single-row plots with double row spacing (40 cm), three-row plots and six-row plots both with single row-spacing (20 cm). Selection entries were mimiced by 16 different varieties or advanced breeding lines, which were also sown in a yield trial. Each experiment was laid out as a 4-times replicated randomized block design. Row length was 2 m. Alley borders and border-rows of multiple-row plots were harvested separately to evaluate the effects of different harvesting procedures on the selection efficiency. Removal of alley borders was found to be disadvantageous, since the gain in precision was more than offset by the loss in sampled area. Wide spacing of single-row plots improved the selection efficiency in comparison with normal spaced single-row plots. In multiple-row plots the selection efficiency was not improved by harvesting only the central rows.For gross plot yield (= yield of net plot + yield of alley borders) the differences in selection efficiency between the various selection plot-types were explained on the basis of the genetic variance, the environmental variance and the coefficient of genetic correlation with farm yield as determined in the yield trial.     

Use of the gene pools of Triticum turgidum ssp. dicoccoides and Aegilops squarrosa for the breeding of common wheat (T. aestivum), through chromosome-doubled hybrids

Summary Triticum turgidum ssp. dicoccoides (wild emmer wheat, AABB, 2n=28) and Aegilops squarrosa (goat grass, DD, 2n=14) comprise a rich reservoir of valuable genetic material, which could be useful for the breeding of common wheat (T. aestivum, AABBDD, 2n=42). Many accessions of both wild species, most of them selected for resistance to stripe rust, were used to make amphiploids. Two strategies were applied: (1) the production of autopolyploid cytotypes of the wild species, followed by hybridisation, and (2) the production of allotriploid interspecific hybrids, followed by doubling of the number of chromosomes. The first route was unsuccessful because of failure of the crosses between the autopolyploid cytotypes, possibly due to incongruity between the two species and to reduced fertility in the autopolyploid cytotypes. The second route yielded the desired synthetic hexaploids. However, the rate of success of the crosses was low and there were great differences between years, and within years between crosses. Embryo rescue was applied to obtain the primary hybrids (2n=21), which were highly sterile and had on average 0.3 bivalents and 20.4 univalents per pollen mother cell. Various abnormalities were recorded. Doubling of the number of chromosomes sometimes occurred spontaneously or was brought about by colchicine treatment. The large scale of the interspecific hybridisation programme ensured that one-third of the female and one-sixth of the male accessions were represented in the synthetic hexaploids.     

Evaluation of proline analysis and chlorophyll fluorescence quenching measurements as drought tolerance indicators in durum wheat (Triticum turgidum L. var. durum)

Summary The possibility of using proline accumulation and fluorescence inhibition as predictive tests for drought tolerance in durum wheat has been investigated. The drought susceptibility of 25 genotypes was evaluated by comparing yields and yield components in irrigated and non irrigated conditions in the field. A drought susceptibility index (DSI) was calculated based on yields from irrigated and dry treatments and compared with the results obtained using the two physiological criteria. Proline accumulation and chlorophyll fluorescence inhibition were found to be significantly and negatively correlated with DSI of grain yield, biological yield, and thousand kernel weight, and tiller index. The use of both criteria for breeding durum wheat in Mediterranean dryland is discussed.     

Morpho-physiological traits to complement grain yield selection under semi-arid Mediterranean conditions in each of the durum wheat types mediterraneum typicum and syriacum

Summary Some 238 landraces from Algeria and Tunisia representative of the mediterraneum typicum durum wheat type and 265 landraces from Syria and Jordan representing the syriacum type were grown in a semi-arid Mediterranean environment of Northern Syria characterized by moderate drought stress. The germplasm types were compared for mean value, level of variation and relationships with grain yield of various morpho-physiological traits possibly usable for indirect selection of best yielding materials. The syriacum germplasm showed higher yield mainly due to greater earliness of cycle, slightly longer grain filling period, shorter stature, lower early growth vigour and higher drought tolerance expressed by a visual score recorded in another, more stressful environment in the region. It also showed lower variation for all morpho-physiological characters except plant glaucousness, for which it was more variable. The wheat types differed not only for architecture but also for optima of individual morpho-physiological traits required to maximize the yield response in the given environment. Higher yield of syriacum materials was attained at same heading and three to four days delayed maturity with respect to average phenology of three well-adapted control cultivars, and it was favoured by increasing number of kernels per spike, early vigour and drought tolerance. Higher yield of mediterraneum typicum landraces was related to heading and maturity dates approaching those of the control cultivars and to increasing kernel weight, early vigour and drought tolerance. Plant stature hardly affected the yield. Both absent and strong glaucousness could confer a yield advantage in syriacum materials.     

Development of diversity array technology (DArT) markers for assessment of population structure and diversity in Aegilops tauschii

Aegilops tauschii Coss. is the D-genome donor to hexaploid bread wheat (Triticum aestivum) and is the most promising wild species as a genetic resource for wheat breeding. To study the population structure and diversity of 81 Ae. tauschii accessions collected from various regions of its geographical distribution, the genomic representation of these lines were used to develop a diversity array technology (DArT) marker array. This Ae. tauschii array and a previously developed DArT wheat array were used to scan the genomes of the 81 accessions. Out of 7500 markers (5500 wheat and 2000 Ae. tauschii), 4449 were polymorphic (3776 wheat and 673 Ae. tauschii). Phylogenetic and population structure studies revealed that the accessions could be divided into three groups. The two Ae. tauschii subspecies could also be separately clustered, suggesting that the current taxonomy might be valid. DArT markers are effective to detect very small polymorphisms. The information obtained about Ae. tauschii in the current study could be useful for wheat breeding. In addition, the new DArT array from this Ae. tauschii population is expected to be an effective tool for hexaploid wheat studies.     

Increase in grain protein percentage in high-yielding common wheat breeding lines by genes from wild tetraploid wheat

Summary Forty-one breeding lines of common wheat, derived from crosses between the Israeli cultivars Miriam and Lakhish and high-protein lines of wild tetraploid wheat, Triticum turgidum var. dicoccoides, were tested for various protein and yield parameters in field trials, under typical agronomic conditions. All lines had a higher grain protein percentage (GPP) than the leading Israeli cultivar Deganit, which was grown as a control. Grain yield (GY) ranged in the breeding lines from a low of 2.44 t/ha to as high as that of Deganit (6.95 t/ha). Despite the weak negative correlation between GPP and GY, several lines excelled both in GPP and in GY. The grain protein yield (GPY) of some of these selected breeding lines was higher than that of Deganit; e.g., 1.19 t/ha in the best line vs. 1.02 t/ha in Deganit. The 16.7% increase in GPY in this line reflected a more efficient utilization of nitrogen.     

Influence of genotype and culture conditions on the production of embryos from anthers of tetraploid wheat (Triticum turgidum)

Summary Anther culture of 10 tetraploid wheat (Triticum turgidum) genotypes and two backcross lines representing a wide range of genetic variation was studied in a randomized block design with three replications. Each replication consisted of 2 pots with 3 plants. The day length was 16h and temperature 25° C/15°C for day/night in a controlled greenhouse where the anther donor plants were grown. Two different treatments were used for anther culture. The first one was potato 2 medium (Chuang et al., 1978) modified by adding 0.5 mg/l glutamine and solidified by gelrite (4g/l) (Henry & De Buyser, 1981). Cultures were incubated in light (15 E m^–2 S^–1) at 26°C at 16h day length. The second medium was described by Fadel & Wenzel (1990), differing from the first by the nature of the sugar (maltose) and consistency of the medium (semiliquid by ficoll). Anther cultures were incubated in the dark at 28°C. The study of about 1300 anthers per genotype and treatment showed that both genotype and treatment affected embryo formation of tetraploid wheat. The backcross lines exhibited significant differences for androgenic abilities when compared to their common parent. Most of the genotypes were medium dependent for androgenesis and revealed significant interactions with the two treatments. Five green plantlets were regenerated and fertile doubled haploid plants were obtained from three out of the 12 studied genotypes.     

Location of genes for high grain protein percentage and other quantitative traits in wild wheat Triticum turgidum var. dicoccoides

Summary The genetic control of grain protein percentage (GPP) in the wild tetraploid wheat, Triticum turgidum var. dicoccoides, was determined by crossing four accessions of this taxonomic variety with durum cultivar Inbar, and analyzing the parents, F₁ and F₂ populations. Reciprocal crosses indicated no cytoplasmic effect on GPP. The F₂ variation was continuous in all crosses, showing no transgressive segregation. However, crosses between different accessions of var. dicoccoides showed transgressive segregation indicating the presence of different genes for high GPP in these accessions. Grain protein percentage was mostly codominant with high GPP, showing either no dominance, or a weak dominance. Heritability coefficients (broad sense) ranged from 0.30 to 0.53. Correlation coefficients between GPP and yield components were usually significantly negative, with the exception of the number of spikelets per spike, and in some crosses, grain weight.The number and chromosomal location of genes coding for high GPP were determined by the association between GPP and 27 markers (23 morphological and 4 biochemical markers). For this purpose, the genetic control of these markers, their linkage groups and chromosomal location were studied. At least four loci for high GPP that segregated in the F₂ populations are suggested: one on chromosome arm 1AS, marked by the black glume gene (Bg); one on 1BS, marked by the HMW gliadin locus Gli-B1; one on group 5, marked by the genes for beaked glume (Bkg) and toothed palea (Tp); and one on group 7, marked by the kinky neck gene (Kn). The relationship between GPP and several yield components was studied in a similar manner. In general, loci of markers that correlated positively with high GPP were not correlated with a decrease in yield components. Moreover, several loci of var. dicoccoides were associated with an increase in yield components.The utilization of markers for chromosomal location of genes coding for quantitative traits is compared to the technique of aneuploid analysis, commonly used in wheat. The significance of the above findings for breeding is discussed.     

Flag leaf area in five spring wheat crosses and the relationship to grain yield

Summary Five spring wheat crosses were evaluated over a 6-year period using comparisons between F₂ and F₃ data and between near-isogenic F₄ populations selected for flag leaf area. Nonsignificant r values for F₂ vs. F₃ flag leaf measurements may be due to the effect of environment on flag leaf area, but are probably also an indication of low heritability for this plant character. Near-isogenic populations selected on the basis of flag leaf area showed little difference in grain yield, an indication that other plant parts must be more influential in determining grain yield. Flag leaf area, by itself, appears not to be a good index to plant performance.Joint contribution of ARS/USDA and the Montana Agricultural Experiment Station. Published with approval of the Director of the Montana Agricultural Experiment Station as Paper No. 725. Journal Series. Use of data from Research Centers at Moccasin. Havre. Huntley, and Kalispell is gratefully acknowledged.     

Selection for yield in wheat breeding

Summary If selection based on F₃ yield tests is to be effective, the yield tests must be successful in discriminating among yield genotypes. The available literature indicates that simple tests with limited or no replication are not very effective, although more extensive, replicated tests may be.Data from an experiment comparing F₃ yield tests with a single seed descent procedure showed that F₃ selection based on a two-replicate test with single seed descent procedure did not justify the extra work involved.     

Relationship between relative water content during reproductive development and winter wheat grain yield

Summary Water is often the most limiting factor to winter wheat (Triticum aestivum L.) production in the southern Great Plains of the U.S.A., yet the lack of reliable screening criteria has precluded direct selection for drought resistance in breeding programs. Previous work showed that leaf relative water content (RWC) was highly heritable when measured under field-drought conditions, but its adoption as a screening tool for yield improvement requires further investigation of the genetic relationship between grain yield and RWC. Plants representing high and low yield potential under drought stress, and a random group of plants, were selected from an F₂ population having the pedigree, TAM W-101/Sturdy. Two sets of entries, each comprised of the two parents and 24 F₂-derived lines, were evaluated under a rainshelter in the F₃ (1986) and F₄ (1987) generations to determine differences in leaf RWC during reproductive development. One set of entries did not receive any water after the jointing stage, and the other set was grown under well-watered conditions. A positive relationship was observed between grain yield and RWC measured during anthesis and mid-grain fill, as the high-yield selections maintained a significantly higher RWC than the low-yield selections. Grain yield and RWC were also positively associated among random selections segregating for both traits. Subsequent adjustment of genotype means for differences in reproductive development at time of sampling underscored the need to consider differences in maturity when RWC is the selection criterion.     

Genetic analysis and selection for wheat yield in drought-stressed and irrigated environments

Summary Wheat (Triticum aestivum L.) cultivars grown in the southern Great Plains of the U.S.A. are exposed to a wide range of moisture conditions due to large fluctuations in the amount and frequency of rainfall. Yield stability under those conditions is therefore a desirable trait for wheat breeders. Our primary objective was to quantify various genetic parameters for grain production in drought-stressed and irrigated environments. We also attempted to predict and measure yield responses when selection is practiced in either drought-stressed or irrigated environments, or both. Seventy F₂-derived lines from the cross, TAM W-101/Sturdy, were evaluated at Goodwell, OK, under irrigated and naturally drought-stressed conditions in 1987 and 1988. Genetic variance and heritability estimates were higher in the irrigated environment than in the drought-stressed environment. The genetic correlation coefficient for yields in the two environments was 0.20±0.16, indicating that selection of widely adapted genotypes requires testing in both environments. Based on the genetic variance/covariance structure of this particular population, the linear index which maximized the combined expected gain in both environments was 0.66Y₁ + 0.34Y₂, in which Y₁ and Y₂ are yields in the irrigated and drought-stressed environments. This index is not expected to apply across all populations; rather, it further supports the hypothesis that testing in either environment alone (drought stressed or irrigated) may not be most effective for increasing either mean productivity or yield under drought stress.