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.     

The inheritance and chromosomal location of morphological traits in wild wheat, Triticum turgidum L. ssp. dicoccoides

The homoeologous groups of chromosomes carrying the genes for some morphological traits in wild wheat Triticum turgidum L. ssp. dicoccoides (T. dicoccoides) were determined, but not the actual chromosomes carrying them. The objectives of this study were to investigate the modes of inheritance, and determine the chromosomes carrying some morphological traits in wild emmer (2n = 28; AABB), the progenitor of most cultivated wheats. To investigate the inheritance of morphological traits, crosses were made between T. turgidum L. ssp. durum (T. durum cultivar Chartokhmi (IR10) and T. dicoccoides accessions TA1150 and TA1131. F₂ seeds from each cross were grown in the field and six qualitative characters were investigated. Purple coleoptile, purple auricle, purple culm, hairy auricle, hairy rachilla, and fragility of spike were controlled by single dominant genes. To determine the chromosomal locations, accession TA1131 was crossed with the complete set of LDN D-genome disomic substitution lines. Assessments of F₂ populations showed that chromosomes 7A, 6A, 7B, 5B, 5A and 3B carried genes for purple coleoptile, purple auricle, purple culm, hairy auricle, hairy rachilla and brittle rachis, respectively.     

Mapping of a gene (Vir) for a non-glaucous, viridescent phenotype in bread wheat derived from Triticum dicoccoides, and its association with yield variation

The introgression of desirable genes or alleles from the wild relatives of hexaploid wheat can be a valuable source of genetic variation for wheat breeders to enhance modern varieties. The UK Group 1 bread making variety Shamrock is an example where the introgression of genetic material from wild emmer (Triticum dicoccoides) has been used to develop a modern cultivar. A striking character of Shamrock is its unique viridescent colour compared to other UK wheats, a trait that coincides with a non-glaucous phenotype. A doubled haploid population segregating for the trait (Shamrock × Shango) was examined to map the location of Vir, and analyse any associated pleiotropic effects. The viridescence gene located to the distal end of the short arm of chromosome 2B. QTL analysis of productivity traits shows an association between Vir and a significant delay in senescence, resulting in an extension of the grain filling period. A stable yield QTL, accounting for up to a quarter of the variation in one case, was also identified at or near Vir, indicating significant yield benefits either by linkage or pleiotropy.     

Identification of Resistance Genes Against Powdery Mildew in Four Accessions of Hordeum Vulgare SSP. Spontaneum

Summary Four newly detected accessions of wild barley (Hordeum vulgare ssp. spontaneum) resistant to powdery mildew caused by Blumeria graminis f. sp. hordei were studied with the aim of finding the number of genes/loci conferring the resistance of individual accessions, the type of inheritance of the genes and their relationships to the Mla locus. F₂ populations after crosses between the winter variety ‘Tiffany’ and four wild barley accessions and use of microsatellite DNA markers were focused on the identification of individual resistance genes/loci by means of their chromosomal locations. In PI466495, one locus conferring powdery mildew resistance was identified in highly significant linkage with the marker Bmac0213. This location is consistent with the known locus Mla on chromosome 1HS. In the other three accessions the resistance was determined by two independent loci. In PI466197, PI466297 and PI466461, one locus was identified on chromosome 1HS and three new loci were revealed on chromosomes 2HS (highly significant linkage with Bmac0134), 7HS (highly significant linkage with Bmag0021) and 7HL (significant linkage with EBmac0755). Our prospective aim is identification of further linked DNA markers and the exact location of the resistance genes on the barley chromosomes.     

Morphology,geography and infraspecific taxonomics of Triticum dicoccoides Körn. A retrospective of 80 years of research

Summary Research on the morphology, geography and infraspecific taxonomics of Triticum dicoccoides Körn. performed during the 80 years since its rediscovery by Aaronsohn is reviewed. T. dicoccoides is a vluable genetic resource for the improvement of cultivated wheat. Its intraspecific structure is, however, not known sufficiently. It is evident that T. dicoccoides is subdivided into two morphologically and geographically distinct forms. The intepretations of these forms by various authors are diverse. There is also no consensus concerning their geographical within the general range of T. dicoccoides. It is necessary to carry out a detailed morphological study in order to define their taxonomical rank and specify their ranges. It would be expedient to perform comparative genetic, biochemical and agronomic investigations of the two T. dicoccoides infraspecific types.     

Wild emmer: genetic resources,gene mapping and potential for wheat improvement

Wild emmer, Triticum dicoccoides, the progenitor of cultivated wheat, harbors rich genetic resources for wheat improvement. They include many agronomic traits such as abiotic stress tolerances (salt, drought and heat), biotic stress tolerances (powdery mildew, rusts, and Fusarium head blight), grain protein quality and quantity, and micronutrient concentrations (Zn, Fe, and Mn). In this review, we summarize (1) traits and controlling genes identified and mapped in T. dicoccoides; and (2) the genes transferred to cultivated wheat from T. dicoccoides. These genes, controlling important agronomic traits such as disease resistance, high protein and micronutrient content, should contribute to wheat production and food nutrition. However, most of the rich genetic reservoir in wild emmer remains untapped, highlighting the need for further exploration and utilization for long-term wheat breeding programs.     

Genetics of five hairy phenotypes and a linkage group of Pennisetum americanum

Summary In pearl millet hairy lamina, hairy sheath and hairy stem were inherited as monofactorial recessives while hairy leaf margin and hairy node were inherited as monogenic dominant traits. The gene for hairy lamina hl showed independent assortment from the gene for hairy node, Hn, and showed linkage with the genes hst (hairy stem), hs (hairy sheath) and Hm (hairy leaf margin). Furthermore, Hl was observed to have an epistatic effect on the expression of hs. The percentages of recombination between the gene pairs hl-hst, hl-hs and hl-Hm were 0.0, 8.30±0.44 and 19.81±0.98 respectively. Thus the genes hl-hst-hs Hm form one linkage group.     

A comparison of marker-assisted and phenotypic selection for high grain protein content in spring wheat

Wheat grain protein content (GPC) is a primary end-use quality determinant for hard spring wheat (Triticum aestivum L.), and marker-assisted selection (MAS) could help plant breeders to develop high GPC cultivars. Two experiments were conducted using two populations developed by crossing low GPC cultivars (Ember) and (McVey) with (Glupro), which contains a high GPC QTL from Triticum dicoccoides (DIC). In one experiment, MAS and phenotypic selection (PS) were employed to select high GPC genotypes, and the selected genotypes were grown in six North Dakota (ND), USA environments. In a second experiment, molecular markers were used to select BC₂F₂ plants from each marker class for the DIC allele from each population. These plants were twice self-pollinated to produce BC₂F₄ plants, which were grown in single ND and Minnesota (MN) environments. Mean GPC was highest among lines using PS at two environments and not significantly different between MAS and PS in the other four environments. Lines presumably homozygous for DIC alleles had significantly higher GPC than their respective low GPC parents. The phenotypic GPC variation explained by the markers (r ²) was 30% at the ND and 15% at the MN environment. The use of PS was as effective as MAS in selecting for high GPC genotypes and more effective in some environments. This likely can be attributed to PS enabling selection for both the major QTL and other genes contributing to GPC. The use of molecular markers might be more advantageous for transferring the high GPC DIC QTL in a backcrossing program during parent development.     

Resistance of Triticum dicoccoides to infection with Erysiphe graminis tritici

Summary The reactions of 233 Triticum dicoccoides acessions, collected at 10 sites in Israel and elsewhere, to infection with cultures of Erysiphe graminis tritici, were determined. The reactions indicated that the number of sources of resistance to E. graminis tritici which can be obtained from T. dicoccoides plants growing wild in Israel and elsewhere is almost unlimited. One hundred and fourteen or 49% of the accessions were resistant, and 137 or 59% of the accessions were resistant or moderately resistant to infection with four cultures of E. graminis tritici which possess the virulence genes corresponding to most of the identified resistance genes in wheat. Accessions collected at sites with marginal habitats where T. dicoccoides grows poorly and has lower grain weight, were more susceptible than were accessions collected at sites with an optimal habitat for growth of T. dicoccoides. The results agreed with those in a previous study with Hordeum spontaneum, and indicate that to obtain H. spontaneum or T. dicoccoides accessions with the highest level of resistance to the powdery mildew pathogens, plants should be collected at sites in ecological and geographic regions where those two species occupy optimum habitats and are exposed to the powdery mildew pathogens.     

Cytogenetic studies in wheat XVII. Monosomic analysis and linkage relationships of gene Yr15 for resistance to stripe rust

Summary The highly effective stripe rust resistance gene, Yr15, derived from Triticum dicoccoides, was located in chromosome 1BS. Yr15 showed linkage of 0.30 (34 cM) with Yr10 and 0.07 with the centromere. Yr15 was preferentially transmitted relative to its alternate allele.     

Increased grain protein content and its association with agronomic and end-use quality in two hard red spring wheat populations derived from Triticum turgidum L. var. dicoccoides

Suitability of wheat (Triticum aestivum L.) for many food products depends on its unique protein. Elevated grain protein content (GPC) and its quality influences the bread making properties of wheat. The objective of this study was to examine the association of elevated GPC with agronomic and end-use quality in two hard red spring wheat recombinant inbred (RI) populations derived from wild emmer (Triticum turgidum L. var. dicoccoides). The two hard red spring populations (ND683/Bergen and Glupro/Bergen) were developed using a single-seed-descent method. ND683 and ‘Glupro’ are high in GPC (180 g kg^-1), presumably due to the introgression of gene(s) from Triticum turgidum L. var. dicoccoides and ‘Bergen’ is low in GPC (145 g kg^-1). From each of the two populations 12 high- and 12 low-GPC RI lines (F_5:7) were selected for replicated testing at two North Dakota (ND) locations in 1995. In both populations, the high-GPC lines had significantly (p < 0.05) higher values compared to the low-GPC lines for mean GPC and water absorption. Mean grain yield of the high-GPC lines was not significantly different from the low-GPC lines in both populations. In the ND683/Bergen population, the high-GPC lines had significantly (p < 0.05) higher values than the low-GPC lines for mean plant height, days to heading, and flour extraction. GPC was significantly (p < 0.05)and negatively associated with test weight and also significantly (p < 0.01) and positively associated with water absorption in the Glupro/Bergen population. In these populations, results suggested that it may be possible to select lines that combine higher GPC and acceptable yield level, but later in maturity and taller in plant height. This revised version was published online in July 2006 with corrections to the Cover Date.     

Association of RFLP markers with trait loci affecting clubroot resistance and morphological characters in Brassica oleracea L.

Summary Resistance to Plasmodiophora brassicae Wor. race 7, the causal agent of the disease clubroot, was examined in an F₂ population of a cross between a clubroot resistant broccoli (Brassica oleracea var. italica) and a susceptible cauliflower (B. oleracea var. botrytis). A genetic linkage map was constructed in the same population based on the segregation of 58 dispersed restriction fragment length polymorphism (RFLP) markers. Associations between the inheritance of RFLP marker genotypes and segregation for disease resistance, morphological and maturity characteristics were examined. For each triat examined, several chromosomal regions marked by RFLP probes appeared to contain trait loci, suggesting that each trait was under polygenic control. RFLP marker linkage to a major factor imparting dominance for clubroot resistance from the broccoli parent was observed in this population. Additionally, RFLP marker linkage to an independently segregating factor contributing clubroot resistance from the cauliflower parent was observed, indicating that it should be possible to use RFLP markers to facilitate selection of transgressive segregants having the combined resistance from both parental sources. In some instances, RFLP markers from the same or closely linked chromosomal regions were associated with both clubroot resistance and morphological traits. Analysis of RFLP marker genotypes at linked loci should facilitate the selection of desired disease resistant morphotypes.     

The introduction into bread wheat of a major gene for resistance to powdery mildew from wild emmer wheat

Summary A new source of resistance to wheat powdery mildew caused by Erysiphe graminis has been transferred to hexaploid bread wheat, Triticum aestivum, from the wild tetraploid wheat, Triticum dicoccoides. The donor was crossed to bread wheat and the pentaploid progeny was then self-pollinated. Plants having a near stable hexaploid chromosome complement were selected in the F₃ progeny and topcrossing and backcrossing of these to a second wheat cultivar to improve the phenotype was undertaken. Monosomic analysis of early backcross lines showed the transferred gene to be located on chromosome 4A. The gene has been designated Pm16.     

Responses of Israeli wild emmers to selected Australian pathotypes of Puccinia species

Summary Seedling responses to one Australian isolate of each of the stripe rust, stem rust and leaf rust pathogens were determined for 541 accessions of T. dicoccoides collected from 23 locations in Israel. Resistance to stripe rust was more frequent than resistance to stem rust. Stripe rust responses showed a wide range of variability indicative of a number of genes for resistance. Comparison of the present stem rust data and that reported for the same accessions tested in Israel indicated that different genes were operating in each country. Only moderately resistant responses to stem rust were obtained. This level of resistance is probably inadequate for transfer to commercial wheat cultivars. We found no potentially useful seedling resistance to leaf rust.     

Inheritance and chromosomal location of the homoeologous genes affecting phenol colour reaction of kernels in durum wheat

End-use quality of wheat for noodles is influenced by polyphenol oxidase activity and its corresponding substrates. This study investigated the chromosomal location of genes that determine phenol colour reaction of kernels in tetraploid wheat using aneuploid stocks. Polyphenol oxidase activity was estimated by the colour reaction of kernels to phenol solution. It was found that the genes located on homoeologous group 2 chromosomes have an important effect on the level of phenol colour reaction of kernels. The genes (Tc1 and Tc2) responsible for high phenol colour reaction of kernels were mapped to the long arms of chromosome 2A and chromosome 2B, respectively. The map distances were estimated to be 46.8 cM for Tc1 and 40.7 cM for Tc2 from the centromere using double-diltelosomics of durum wheat.     

Genetic analysis of resistance to bacterial leaf streak caused by Xanthomonas campestris pv. undulosa in bread wheat

Summary The inheritance of resistance to bacterial leaf streak or black chaff of wheat was studied under field conditions, with an artificial epidemic of Xanthomonas campestris pv. undulosa. A complete series of crosses between five parents, differing in reaction to X. c. pv. undulosa, was generated. Disease was recorded at two different stages of growth. No evidence of cytoplasmic effect was found from the comparison between reciprocal F₁ crosses. The study of the F₃ generations attested that five genes were involved in resistance to bacterial leaf streak. Separate analyses carried out for the two scoring dates were mutually consistent: genotypes, number of genes, and their action and relative importance were verified. The genes differed in strength of expression of resistance. One of the two strongest genes, Bls1, is present in all three superior parents, Pavon 76, Mochis T88 and Angostura F88. Resistance was not complete, and proved to be stable over the season.     

The effects of homoeologous group 3 chromosomes on grain colour dependent seed dormancy and brittle rachis in tetraploid wheat

`Langdon' (LDN), a durum wheat (Triticum turgidum L. var. durum) cultivar, and a set of chromosome substitution lines of Langdon, where A or B genome chromosome were replaced with a homologous chromosome of wild emmer wheat, T. turgidum ssp. dicoccoides (DIC), were used to assess the effect of the specific chromosome on seed dormancy in tetraploid wheat. The LDN(DIC 3A) and LDN (DIC 313) lines showed significantly lower seed germination than Langdon. It appears that LDN(DIC 3A) and LDN(DIC 3B) have red grain whose allele were designated as R-A1b and R-B1b, respectively and the rachises of LDN(DIC 3A) and LDN(DIC 3B) were fragile. The alleles for brittle rachis were designated as Br ₂ for LDN(DIC 3A) and Br ₃ for LDN(DIC 3B). From the F₂ of the crosses, Langdon/LDN(DIC 3A) and Langdon/LDN(DIC 3B), Br ₂ was located approximately 44.2 cM from the R-A1b locus and Br ₃ approximately 47.0 cM from the R-B1b locus, respectively. Recombinant inbred chromosomal lines for 3A and 3B were used to assess (1) the linkage relationship between grain colour and fragile rachis, and (2) the effect of grain colour on germination. Estimated distance between R-B1b – Br ₂ was 39.6 cM. For the 3A population, germination percentage of both colour groups was 12.4% for the red grain group and 68.6% for the amber group, respectively. For the 3B population, germination percentage of the red group was 7.3% and that of the amber group was 82.1%. For both populations, differences were statistical significant by t-tests. We considered that seed dormancy of T. turgidum ssp. dicoccoides was dependent on grain colour. It raised the possibility that brittle rachis is due to a paralogous gene set on homoeologous group 3 chromosomes. This revised version was published online in July 2006 with corrections to the Cover Date.     

Origin,diversity and utilization of the Cuban germplasm of common bean (Phaseolus vulgaris L.)

Summary Wheat root characters which influence vital plant processes have scarcely been explored for their genetic control. This study was conducted to i) examine the diversity of root traits and associated shoot traits in spring wheat (Triticum aestivum L.) cultivars; ii) study the nature of genetic control of selected traits; and iii) examine associations among root and shoot traits. Three experiments were conducted in the greenhouse with plants grown in the vermiculite medium in clear plastic tubes. In the first experiment, 42 spring wheat cultivars were grown for three weeks and measurements were taken on root length, leaf length, root number, leaf number, root dry weight, and top dry weight. In the second study, 15F₁'s originating from a partial diallel mating of six cultivars along with the parents were evaluated for 4 weeks. The data on root length, leaf length, and root number were subjected to diallel analysis according to Griffing's method 4, fixed model. A third experiment consisted of studying 2 F₂ populations with 141 plants per population. In the first study the 42 cultivars exhibited a wide range of variation for all six traits. Diallel analysis revealed significant effects of both general combining ability (GCA) and specific combining ability (SCA) for root length, leaf length, and root number. Parents with high GCA estimates were identified. Significant positive correlation coefficients were found among root and shoot traits. Analysis of F₂'s for root length indicated quantitative nature of inheritance of root length.     

Plant nitrogen distribution in wild tetraploid (Triticum turgidum dicoccoides) and hexaploid wheat (Triticum aestivum)

Summary Differences were found in total nitrogen uptake and its pattern of distribution in the main tiller amongst five lines of wild tetraploid wheat (Triticum turgidum dicoccoides) and between it and two hexaploid wheats (Triticum aestivum) under low (48 ppm) and higher (240 ppm) levels of soil nitrogen.Under the low soil nitrogen level the hexaploids had higher amounts of total nitrogen in the main tiller than the dicoccoides lines, but under the higher soil nitrogen level, three of the dicoccoides lines had significantly (P<0.01) higher, and the other two lines, similar amounts as the hexaploids. The total amount of grain nitrogen in the hexaploids was significantly (P<0.01) higher than the five dicoccoides under the low nitrogen soil level but under the higher level, two of the dicoccoides lines had similar amounts as one of the hexaploids (cv. Bencubbin) but significantly (P<0.01) lower than the other (cv. Argentine IX).The efficiency of nitrogen translocation to the grain was significantly (P<0.01) lower in a primitive, compared with four cereal forms of dicoccoides under both low and high levels of soil nitrogen. The cereal forms of dicoccoides, while similar in nitrogen translocation efficiency under low soil nitrogen as the lower translocation efficiency hexaploid (cv. Bencubbin), were significantly (P<0.01) and substantially lower than it under the higher soil nitrogen level.     

Origin and variation of gliadin proteins associated with pasta quality in durum wheat

Summary Studies were made of the presence and frequency of occurrence of gliadin bands 42 and 45 in three samples of Aegilops sharonensis Eig and 59 samples of wild tetraploid wheat (Triticum dicoccoides Korn.) from natural distributions of these species in Israel.Two samples of Ae. sharonensis possessed a band in position 45 and one possessed no bands corresponding to either band 45 or band 42. In T. dicoccoides, band 45 was either present or not and 42 was always absent. In its grassy and intermediate growth habit forms, (believed to be more primitieve than the cercal forms) band 45 appeared to be more frequent than in the cereal form.The presence of band 45 in the Ae. sharonensis, and its relatively high frequency in T. dicoccoides, populations from Mt. Hermon (likely to be relatively free from introgression from cultivated tetraploid wheat) indicate the likelihood of a primary origin of the allele coding for band 45. The absence of band 42 from all Ae. sharonensis and T. dicoccoides populations in this study, indicates a more recent evolutionary origin of the allele coding for this band, possibly arising as a mutation during the domestication of tetraploid wheat.The results have implications for breeding programmes in tetraploid wheat.