High‐density molecular characterization and association mapping in Ethiopian durum wheat landraces reveals high diversity and potential for wheat breeding

Durum wheat (Triticum turgidum subsp. durum) is a key crop worldwide, and yet, its improvement and adaptation to emerging environmental threats is made difficult by the limited amount of allelic variation included in its elite pool. New allelic diversity may provide novel loci to international crop breeding through quantitative trait loci (QTL) mapping in unexplored material. Here, we report the extensive molecular and phenotypic characterization of hundreds of Ethiopian durum wheat landraces and several Ethiopian improved lines. We test 81 587 markers scoring 30 155 single nucleotide polymorphisms and use them to survey the diversity, structure, and genome‐specific variation in the panel. We show the uniqueness of Ethiopian germplasm using a siding collection of Mediterranean durum wheat accessions. We phenotype the Ethiopian panel for ten agronomic traits in two highly diversified Ethiopian environments for two consecutive years and use this information to conduct a genome‐wide association study. We identify several loci underpinning agronomic traits of interest, both confirming loci already reported and describing new promising genomic regions. These loci may be efficiently targeted with molecular markers already available to conduct marker‐assisted selection in Ethiopian and international wheat. We show that Ethiopian durum wheat represents an important and mostly unexplored source of durum wheat diversity. The panel analysed in this study allows the accumulation of QTL mapping experiments, providing the initial step for a quantitative, methodical exploitation of untapped diversity in producing a better wheat.     

Morphological studies of free-threshing wheat ears from a Neolithic site in southwest Germany, and the history of the naked wheats

Two hundred and seventy carbonized naked wheat ears from the Neolithic pile dwelling at Hornstaad-Hörnle IA/Lake Constance (Germany) were analysed. The characteristics of internodes and glumes show that only tetraploid naked wheat types were cultivated. Following the history of free-threshing wheats it seems conceivable that the tetraploid naked wheats spread from their point of origin in the Middle East (7th/8th millennium B.C.) via a Mediterranean route to southwest Europe, reaching the northern foothills of the Alps about 4000 B.C.     

Crop water availability in early agriculture: evidence from carbon isotope discrimination of seeds from a tenth millennium BP site on the Euphrates

The analysis of carbon isotope discrimination (Δ) in crop plant remains from archaeological sites may help to assess water availability for early agriculture. This study presents the analysis of Δ in seeds of naked wheat (Triticum aestivum/durum), lentil (Lens orientalis/culinaris), and flax (Linum sp.) found at the archaeological site of Tell Halula in the valley of the Middle Euphrates (Syria). This Neolithic site is the oldest in this region of the Fertile Crescent where the cultivation of domesticated plants has been reported, with seed remains ranging from 9550 to 8465 BP. Most of the seeds analysed showed Δ values greater than 16 ‰, reaching 20 ‰ for some samples of flax. For wheat, Δ values were much higher than those reported in present-day (1996) durum wheat crops cultivated under rainfed conditions in north-west Syria under environments with somewhat higher rainfall than Tell Halula. Similarly, grains of present-day (1997) barley cultivated in the archaeological site also showed lower values than those found in archaeological kernels. An empirical relationship between Δ of mature kernels and total precipitation (plus irrigation where applicable) from heading to maturity (r² = 0.82, n = 11) was established for durum wheat, currently cultivated in different environments of the Mediterranean basin. The resulting relationship was applied to the data on Δ of wheat fossil kernels from Tell Halula to estimate the accumulated water inputs during the time (about 6 weeks) the kernels were produced. Calculated water inputs for wheat during early agriculture were (over 110 mm) at least 5 times higher than current-day rainfall accumulated in Tell Halula during the same phenological period. These results strongly suggest that early agriculture wheat was cultivated at Tell Halula under much wetter conditions than are currently to be found in the area. The presence of flax and its very high Δ values also support this conclusion. Whether such humid conditions during cultivation were due to moister conditions prevailing at this time, by planting in alluvial areas or by irrigation works is discussed.     

Quantifying relationships between rooting traits and water uptake under drought in Mediterranean barley and durum wheat

In Mediterranean regions drought is the major factor limiting spring barley and durum wheat grain yields. This study aimed to compare spring barley and durum wheat root and shoot responses to drought and quantify relationships between root traits and water uptake under terminal drought.One spring barley（Hordeum vulgare L. cv. Rum） and two durum wheat Mediterranean cultivars（Triticum turgidum L. var durum cvs Hourani and Karim） were examined in soil‐column experiments under well watered and drought conditions. Root system architecture traits, water uptake, and plant growth were measured. Barley aerial biomass and grain yields were higher than for durum wheat cultivars in well watered conditions. Drought decreased grain yield more for barley（47%） than durum wheat（30%, Hourani）. Root‐to‐shoot dry matter ratio increased for durum wheat under drought but not for barley, and root weight increased for wheat in response todrought but decreased for barley. The critical root length density（RLD） and root volume density（RVD） for 90% available water capture for wheat were similar to（cv. Hourani） or lower than（cv. Karim） for barley depending on wheat cultivar. For both species, RVD accounted for a slightly higher proportion of phenotypic variation in water uptake under drought than RLD.     

Tetraploid wheat landraces in the Mediterranean basin: taxonomy, evolution and genetic diversity

The geographic distribution of genetic diversity and the population structure of tetraploid wheat landraces in the Mediterranean basin has received relatively little attention. This is complicated by the lack of consensus concerning the taxonomy of tetraploid wheats and by unresolved questions regarding the domestication and spread of naked wheats. These knowledge gaps hinder crop diversity conservation efforts and plant breeding programmes. We investigated genetic diversity and population structure in tetraploid wheats (wild emmer, emmer, rivet and durum) using nuclear and chloroplast simple sequence repeats, functional variations and insertion site-based polymorphisms. Emmer and wild emmer constitute a genetically distinct population from durum and rivet, the latter seeming to share a common gene pool. Our population structure and genetic diversity data suggest a dynamic history of introduction and extinction of genotypes in the Mediterranean fields.     

Global diversity of durum wheat Triticum durum Desf. for alleles of gliadin-coding loci

Genetic diversity for the alleles of gliadin-coding loci was studied with 465 durum wheat accessions from 42 countries. A total of 108 alleles were identified for four loci; 60 alleles were described for the first time. Broad diversity of rare gliadin-coding alleles was observed. The highest genetic diversity was characteristic of durum wheat accessions from the Middle East, Trans-Caucasia, the Pyrenean Peninsula, and the Balkans. Two genetically isolated ancient branches of durum wheat were isolated. A “southern” branch included mostly accessions from the Mediterranean region, the Middle East, and Trans-Caucasia. A “northern” branch included Russian and Ukrainian durum wheat accessions and varieties obtained on their basis. An additional group included durum wheat accessions that had been obtained in several past decades on the basis of the material of international breeding centers (CIMMYT and ICARDA) and had low genetic diversity.     

Linkage Disequilibrium and Genome-Wide Association Mapping in Tetraploid Wheat (Triticum turgidum L.)

Association mapping is a powerful tool for the identification of quantitative trait loci through the exploitation of the differential decay of linkage disequilibrium (LD) between marker loci and genes of interest in natural and domesticated populations. Using a sample of 230 tetraploid wheat lines (Triticum turgidum ssp), which included naked and hulled accessions, we analysed the pattern of LD considering 26 simple sequence repeats and 970 mostly mapped diversity array technology loci. In addition, to validate the potential for association mapping in durum wheat, we evaluated the same genotypes for plant height, heading date, protein content, and thousand-kernel weight. Molecular and phenotypic data were used to: (i) investigate the genetic and phenotypic diversity; (ii) study the dynamics of LD across the durum wheat genome, by investigating the patterns of LD decay; and (iii) test the potential of our panel to identify marker–trait associations through the analysis of four quantitative traits of major agronomic importance. Moreover, we compared and validated the association mapping results with outlier detection analysis based on population divergence. Overall, in tetraploid wheat, the pattern of LD is extremely population dependent and is related to the domestication and breeding history of durum wheat. Comparing our data with several other studies in wheat, we confirm the position of many major genes and quantitative trait loci for the traits considered. Finally, the analysis of the selection signature represents a very useful complement to validate marker–trait associations.     

Arbitrarily primed-PCR based diversity assessment reflects hierarchical groupings of Indian tetraploid wheat genotypes

Genetic diversity analysis using PCR with arbitrary decamer primers (RAPD — random amplified polymorphic DNA) was carried out in a set of 63 tetraploid wheat genotypes which comprised 24 durum landraces, 18 durum cultivars, nine dicoccum cultivars, ten less commonly cultivated species and two wild tetraploid species. The durum and dicoccum wheat genotypes are a part of the germplasm used in Indian tetraploid wheat breeding programs. A total of 206 amplification products were obtained with 21 informative primers, of which 162 were polymorphic. The highest degree of polymorphism was seen in the wild and less commonly cultivated species (68.9%). Durum released cultivars showed greater polymorphism (50.6%) than landraces (44.8%), while dicoccum cultivars showed a considerably low level of polymorphism (23.6%). Cluster analysis led to the separation of wild and cultivated genotypes, and among cultivated emmer wheat distinct groups were formed by the durum cultivars, durum landraces and dicoccum cultivars. The subgroupings of landraces had no relation to their geographical distribution. The durum cultivars formed subgroups based on common parentage in their pedigree. Among species, wild timopheevi wheat (T. araraticum) and its cultivated form (T. timopheevi) formed a distinct group distant from all other genotypes. The present study is a first attempt at determining the genetic variation in Indian tetraploid wheats at the molecular level. Received: 10 January 1999 / Accepted: 30 January 1999     

Relationships between carbon isotope discrimination, dry matter production, and harvest index in durum wheat

Carbon isotope discrimination has been proposed as a criterion for the indirect selection to improve transpiration efficiency and grain yield in bread wheat and barley. Less attention has been devoted to durumwheat (Triticum durumDesf.) despite its economic importance in the Mediterranean basin. The Δ genetic variation and its relationships to dry matter production and harvest index in durum wheat were investigated in this study. For this purpose, field experiments were conducted on 144 durum wheat accessions under Mediterranean conditions (South of France) during three consecutive years with contrasting climatic conditions. Grain yield, above-ground biomass, harvest index, and carbon isotope discrimination of flag leaf and kernel were measured. Differences between years, noted for both leaf and kernel carbon isotope discrimination, were probably related to the variation in water availability from year to year. A large genotypic variation was also noticed for both leaf and kernel carbon isotope discrimination. The two traits were found to be positively correlated with grain yield within and across years, which confirms the interest in carbon isotope discrimination for selection for grain yield improvement under Mediterranean conditions. Both kernel and leaf carbon isotope discrimination correlated better with harvest index than with grain yield, suggesting that carbon isotope discrimination could reflect the efficiency of carbon partitioning to the kernel. The lack of correlation between leaf carbon isotope discrimination and both harvest index and grain yield in favourable water conditions (1996) was probably due to the difference in water availability between the period until flag leaves sampling (favourable conditions) and the strong water stress which accompanied the grain filling. Kernel carbon isotope discrimination correlated better with both harvest index and grain yield than did leaf carbon isotope discrimination. Moreover, a higher broad-sense heritability was obtained for kernel carbon isotope discrimination than for leaf carbon isotope discrimination. As a result, kernel carbon isotope discrimination appeared to be a better predictive criterion for efficiency of the carbon partitioning to the kernel (harvest index), and hence for grain yield, than did flag leaf carbon isotope discrimination.     

Carbon isotope discrimination and mineral composition of three organs in durum wheat genotypes grown under Mediterranean conditions

Carbon isotope discrimination (delta) has been proposed as a good criterion for transpiration efficiency and grain yield improvement. Its measurement, however, remains very expensive. Ash content (ma) has been proposed as an alternative criterion for delta in bread wheat and barley. The aims of this study were (i) to analyse the relationships between delta and mineral composition in different durum wheat plant parts and (ii) to compare the variation of these traits between landraces and improved varieties from different geographic origins. For this purpose, delta, ma, and composition in four minerals (K, Mg, P and Si) were assessed in flag leaves and awns at anthesis, and in mature grains of ten durum wheat genotypes grown under rainfed Mediterranean conditions. The three plant parts differed significantly for the measured traits. Significant correlations were noted between delta and ma in the flag leaf and in the grain. Silicon content in flag leaves and potassium content in awns were also positively related to delta of the considered plant part. The coefficient of correlation between delta and ma was generally higher than that observed between individual mineral content and delta, suggesting that ma is the better alternative criterion for delta. In addition, grain yield was related to grain delta and both ma and potassium content in awns. Harvest index was correlated with delta and ma of grain and flag leaf. These results emphasised that ma values in flag leaf and grain represent the efficiency of carbon partitioning to the grain. Improved varieties showed higher delta and ma values than landraces. Differences between Middle-East and West Mediterranean genotypes for the measured traits were also presented and discussed.     

A genome‐wide association study of 23 agronomic traits in Chinese wheat landraces

Uncovering the genetic basis of agronomic traits in wheat landraces is important for ensuring global food security via the development of improved varieties. Here, 723 wheat landraces from 10 Chinese agro‐ecological zones were evaluated for 23 agronomic traits in six environments. All accessions could be clustered into five subgroups based on phenotypic data via discriminant function analysis, which was highly consistent with genotypic classification. A genome‐wide association study was conducted for these traits using 52 303 DArT‐seq markers to identify marker‐trait associations and candidate genes. Using both the general linear model and the mixed linear model, 149 significant markers were identified for 21 agronomic traits based on best linear unbiased prediction values. Considering the linkage disequilibrium decay distance in this study, significant markers within 10 cM were combined as a quantitative trait locus (QTL), with a total of 29 QTL identified for 15 traits. Of these, five QTL for heading date, flag leaf width, peduncle length, and thousand kernel weight had been reported previously. Twenty‐five candidate genes associated with significant markers were identified. These included the known vernalization genes VRN‐B1 and vrn‐B3 and the photoperiod response genes Ppd and PRR. Overall, this study should be helpful in elucidating the underlying genetic mechanisms of complex agronomic traits and performing marker‐assisted selection in wheat.     

The Triticeae genetic resources of central Italy: collection,evaluation and conservation

One hundred and six landraces belonging to 7 species of the Triticeae tribe were collected in central Italy by DBVBA (Perugia University), DIBIAGA (Ancona University) and ARSSA (Abruzzo Region Agricultural Development Agency) in different individual and joint missions. A few accessions were supplied by private and other public organisations. Triticum dicoccum Schubler is the most widespread species, followed by T. aestivum L., T. monococcum L., T. spelta L., T. turgidum var. durum Desf., Secale cereale L. and Hordeum vulgare L. Besides the presence of landraces reproduced by farmers over generations, information related to on-farm management and to qualitative/organoleptic traits as well as information related to their local names, uses, traditions and social context was gathered during the missions. The majority of the accessions was characterised by morphological and phenological traits and molecular markers. This work shows the presence of morpho-phenologic and genetic differences among landraces and the importance of some species in the agricultural systems and food customs of the investigated area. Particularly for emmer three well distinct landraces are present, "Farro Italia Centrale", "Farro della Garfagnana" and "Farro Italia Meridionale". Other interesting and traditional landraces are the "Solina" common wheat in Abruzzo and the "Orzo mondo" naked barley in Marche. Most of the populations are still cultivated in marginal lands and under low input or organic agronomic conditions; nevertheless, in many cases, they are found near modern varieties in conventional agriculture systems. Moreover, the in situ (on-farm) conservation of Triticeae landraces in central Italy is strictly linked to elderly farmers.     

Effects of nitrogen rates on grain yield and nitrogen agronomic efficiency of durum wheat genotypes under different environments

Durum wheat is an important staple food crop in Tunisia and other Mediterranean countries and is grown in various climatic conditions. Production and yield are however severely limited not only by drought events but also by reduced levels of nitrogen fertilisation. A study was carried out at two locations in the sub‐humid area of Tunisia: Mateur in 2009–10 and 2010–11 and Beja in 2011–12 and 2012–13 under rainfed conditions. Four durum wheat genotypes (landraces: Bidi, Azizi; improved: Om Rabia, Khiar) were evaluated for nitrogen agronomic efficiency and related agronomic traits under various nitrogen rates: 0, 50, 100, 150, 200 and 250 kg N ha^?1, with three replications. There was a significant interaction effect (P ≤ 0.001) environments × genotypes × N treatments for grain yield (GY), biomass yield (BY), harvest index (HI), partial factor productivity of applied nitrogen (PFP_N) and nitrogen agronomic use efficiencies (NAE). GY was the most affected trait by nitrogen applied showing an increase of 94% under high N treatment (250 kg N ha^?1) compared to control plots without N treatments. A significant linear regression exists between GY (0 N) and GY for the different N rates (r = 0.70; P < 0.001). This effect was more pronounced for improved genotypes than landraces for all parameters excepting BY and NAE_BY. BY showed +11% increase in landraces than improved genotypes. PFP_N showed an average decrease of 65% under high‐N fertilisation with 10% prevalence for improved genotypes. Landraces tend to promote vegetative growth while grain filling efficiency was higher for improved genotypes.     

Variation in fructooligosaccharide contents during plant development and in different cultivars of durum wheat

Abstract

Fructooligosaccharides (FOS) have received a great deal of attention for their properties as prebiotic components. Several plants store FOS in their tissues as carbohydrate source or as osmoregulators. In comparison with the other natural FOS sources, cereals, in particular durum wheat, store a significant amount of these oligosaccharides during their growth. To gain a deeper insight on the relationships between FOS and plant/kernel development, FOS levels were analysed in the grains and stems of a widely cultivated durum wheat cultivar (Simeto), during the whole period of grain filling. In addition, a broad analysis of FOS contents was carried out by comparing their levels in the stems and grains of 45 cultivars of durum wheat harvested at two development stages. Results show a large variation among cultivars in their capability to metabolise FOS.     

Genetic parameters estimation for some wild wheat species and their F1 hybrids grown in different regions of Saudi Arabia

Wheat (Triticum aestivum L.) is the most important crop for human nutrition that underpins the food safety of Saudi Arabia. The investigation here was to determine heterosis effects using different genetic methods: heterosis over better, mid parents, the genetic advance, and genotype, phenotypic coefficient of variation for estimation some traits among six wheat landraces and their F₁ hybrids. In 2019, these landraces were sown using hand and after 100 days, the emasculation and crossing were made among these six landraces using hand emasculation of anthers. In 2020, seeds for these genotypes (six wheat landraces and their F₁) were sown under normal irrigation accordingly done in 2019. The results showed that the most important parent was Mabia resulted with the highest value in number of tiller/ plant, 1,000-grain weight, and fresh shoot weight. The highest value of plant height among six parents was Naqra while highest value at the same trait among F₁ hybrids was P₃ XP₆. The estimations of heterosis showed that out of 15 crosses, one cross (P₁XP₅) was significantly better yield than all crosses for these four traits. The genotype coefficient of variation (GCV) ranged from 12.5% to 8.7% while phenotypic coefficient of variation ranged from 17.7% to 11.3%. The correlation coefficients was found between fresh shoot weight and number of tiller and plant height and umber of tiller. Wild wheat still serve as a source of useful germplasm with proven adaption and productivity and thus assembles of the wild wheat assortments are the initial step of breeding program.     

Domesticated emmer wheat [T. turgidum L. subsp. dicoccon (Schrank) Thell.] as a source for improvement of zinc efficiency in durum wheat

Modern durum wheat (AABB) is more sensitive to zinc (Zn) deficiency than bread wheat (AABBDD). One strategy to increase productivity and expansion of durum wheat industry in Zn-deficient soils is to improve its ability to grow and yield in such soils. This ability is termed Zn efficiency. In a growth room experiment using soil culture, we assessed the potential of Triticum turgidum L. subsp. dicoccon (Shrank) Thell. (domesticated emmer wheat, AABB) as a genetic resource for further improvement of Zn efficiency in modern durum wheat. Twenty four accessions of domesticated emmer wheat, four durum landraces/cultivars, and two bread wheat cultivars/ advanced breeders lines of known Zn efficiency were tested under Zn deficiency and Zn sufficiency. Significant variation was observed among genotypes in Zn deficiency symptoms, dry matter production, shoot Zn concentration, shoot Zn content and Zn utilisation efficiency (physiological efficiency). We identified domesticated emmer wheat accessions with greater Zn efficiency than modern durum wheat and even bread wheat genotypes. These accessions could be used in breeding programs to improve Zn efficiency of durum wheat. The results suggest that Zn efficiency of durum or bread wheat is likely to be determined collectively by its progenitors.     

河南省花生农家品种资源农艺和品质性状分析

综合分析河南省花生农家品种资源的农艺和品质性状,为花生的遗传育种提供理论依据。以128个不同地域来源的河南省农家品种为材料,田间调查株型、分枝型和开花习性等植物学性状,收获考种测定主茎高、侧枝长、总分枝数、结果枝数、单株产量、百果重和百仁重农艺性状,并测定蛋白质、脂肪、油酸和亚油酸含量。结果表明,农家品种以密枝型为主,农艺性状中百仁重的变异系数最大,为31.1%,其次为单株产量和总分枝数,分别为27.5%和22.2%,变异系数最小的为侧枝长,为12.5%;品质性状方面,河南农家品种资源的脂肪含量较高,脂肪含量55%以上的有10个花生品种。蛋白质含量偏低,最高仅25.3%,油酸含量中等,平均45.4%,最高的52.4%。本研究表明河南省农家品种的农艺性状表现丰富的遗传多样性,品质方面脂肪含量相对较高,合理利用河南省农家品种资源,可为花生品质改良提供优质性状的亲本。     

重庆大豆地方资源多样性评价及群体表型特点研究

对收集于重庆市的103份大豆地方品种进行了表型遗传多样性分析,并利用主成分分析和聚类分析对其进行了群体表型特点的研究,研究结果表明,重庆地区大豆地方种质资源以灰色茸毛、黄粒、褐色脐居多,个体之间数量性状上存在较大变异。主成分分析以3个主成分反映了10个农艺性状的大部分信息,将主要农艺性状归纳成产量因子、生长势因子及籽粒因子。聚类分析将103份地方种质资源聚为6个类群, 同一地区的大部分地方品种表现较为相似,少部分地方品种差异大,品种类群间的表型分化与地理分布既有一定的联系又不绝对相关。利用这些种质选配育种亲本应关注各类群间性状差异作选择,而不能仅关注地理来源选择。     

Overexpression of Puroindoline a gene in transgenic durum wheat (Triticum turgidum ssp. durum) leads to a medium–hard kernel texture

Durum wheat is the second-most widely grown wheat species, and is primarily used in the production of pasta and couscous. The grain utilization of durum wheat is partly related to its very hard kernel texture because of the lack of the D genome and consequentially the Puroindoline genes. Our previous study reported the transformation of durum wheat with the Puroindoline a (Pina) gene. Here, we characterized the transgenic durum wheat lines expressing the Pina gene, and studied the effects of PINA on grain texture and other kernel characteristics. SDS-PAGE and Western blotting results demonstrated that starch-bound PINA levels of Pina-overexpressing lines were lower than that of Pina-positive control, common wheat cv. Chinese Spring, suggesting a weak association of PINA protein with starch granules in the absence of Pinb. Grain hardness analysis and flour milling tests indicated that the overexpression of PINA resulted in decreased grain hardness and increased flour yield in transgenic durum wheat lines. The agronomic performance of the transgenic and control lines was also examined and it was found that no significant differences in measured traits were observed between Pina-overexpressing and control lines in the 2-year field trials. Since grain hardness strongly affects milling and end-use qualities, the development of medium–hard-textured durum wheat lines is not only of significance for our knowledge of grain hardness and Puroindolines, but also has practical implications for plant breeders and food technologists for the expansion of utilization of durum wheat.     

Identification of QTL and association of a phytoene synthase gene with endosperm colour in durum wheat

The yellow colour of durum wheat (Triticum turgidum L. var durum) semolina is due in part to the presence of carotenoid pigments found in the endosperm and is an important end-use quality trait. We hypothesized that variation in the genes coding for phytoene synthase (Psy), a critical enzyme in carotenoid biosynthesis, may partially explain the phenotypic variation in endosperm colour observed among durum cultivars. Using rice sequence information, primers were designed to PCR clone and sequence the Psy genes from Kofa (high colour) and W9262-260D3 (medium colour) durum cultivars. Sequencing confirmed the presence of four Psy genes in each parent, corresponding to a two member gene family designated as Psy1-1, Psy1-2 and Psy2-1 and Psy2-2. A genetic map was constructed using 155 F1-derived doubled haploid lines from the cross W9262-260D3/Kofa with 194 simple sequence repeat and DArT markers. Using Psy1-1 and Psy2-1 allele-specific markers and chromosome mapping, the Psy1 and Psy2 genes were located to the group 7 and 5 chromosomes, respectively. Four quantitative trait loci (QTL) underlying phenotypic variation in endosperm colour were identified on chromosomes 2A, 4B, 6B, and 7B. The Psy1-1 locus co-segregated with the 7B QTL, demonstrating an association of this gene with phenotypic variation for endosperm colour. This work is the first report of mapping Psy genes and supports the role of Psy1-1 in elevated levels of endosperm colour in durum wheat. This gene is a target for the further development of a molecular marker to enhance selection for endosperm colour in durum wheat breeding programs.