A population of single chromosome recombinant lines was developed from the cross between a frost-sensitive, vernalization-insensitive substitution line, 'Chinese Spring' (Triticum spelta 5A) and a frost-tolerant, vernalization-sensitive line, 'Chinese Spring' ('Cheyenne' 5A), and used to map the genes Vrn1 and Fr1 controlling vernalization requirement and frost tolerance, respectively, relative to RFLP markers located on this chromosome. The Vrn1 and Fr1 loci were located closely linked on the distal portion of the long arm of 5AL, but contrary to previous observations, recombination between them was found. Three RFLP markers, Xpsr426, Xcdo504 and Xwg644 were tightly linked to both. The location of Vrn1 suggests that it is homoeologous to other spring habit genes in related species, particularly the Sh2 locus on chromosome 7 (5H) of barley and the Sp1 locus on chromosome 5R of rye.
Two populations of single chromosome recombinant lines were used to map genes controlling flowering time on chromosome 5B of wheat, and one of the populations was also used to map a new frost resistance gene. Genetic maps were developed, mainly using microsatellite markers, and QTL analysis was applied to phenotypic data on the performance of each population collected from growth-room tests of flowering time and frost tolerance. Using a recombinant substitution-line mapping population derived from a cross between the substitution-line 'Chinese Spring' ('Cheyenne' 5B) and 'Chinese Spring' (CS), the gene Vrn-B1, affecting vernalization response, an earliness per se locus, Eps-5BL1, and a gene, Fr-B1, affecting frost resistance, were mapped. Using a 'Hobbit Sib' ('Chinese Spring' 5BL) x 'Hobbit Sib' recombinant substitution line mapping population, an earliness per se locus, Eps-5BL2 was mapped. The Vrn-B1 locus was mapped on the distal portion of the long arm of chromosome 5B, to a region syntenous with the segments of chromosomes 5A and 5D containing Vrn-A1 and Vrn-D1 loci, respectively. The two Eps-5BL loci were mapped close to the centromere with a 16-cM distance from each other, one in agreement with the position of a homoeologous locus previously mapped on chromosome 5H of barley, and suggested by the response of 'Chinese Spring' deletion lines. The Fr-B1 gene was mapped on the long arm of chromosome 5B, 40 cM from the centromeric marker. Previous comparative mapping data with rice chromosome 9 would suggest that this gene could be orthologous to the other Fr genes mapped previously by us on chromosomes 5A or 5D of wheat, although in a more proximal position. This study completes the mapping of these homoeoallelic series of vernalization requirement genes and frost resistance genes on the chromosomes of the homoeologous group 5 in wheat.
SUMMARYUsing "alien" chromosome substitution lines of the wheat variety Chinese Spring (CS), it was found that the chromosomes of homoeologous group 2 were important in determining the time to ear-emergence under different day lengths. In terms of their ability to produce plants insensitive to different day lengths, chromosome 2B of CS was the most potent, followed by 2A of CS, 2M of Aegilops comosa, 2C of Ae. umbellulata, 2D of CS and finally 2Rm of Secale montanum. Dominant epistasis towards insensitivity was observed.
Genetic studies of frost resistance were performed on various wheat varieties using diallel, F2 monosomic and substitution analysis.A six-parental cross including reciprocals was carried out, and F1 hybrids and their parents were used for the freezing tests under controlled conditions. Both the general combining ability (GCA) and the specific combining ability (SCA) were significant, indicating additive and non-additive gene action in the inheritance of frost resistance. The high GCA∶SCA ratio revealed a preponderance of additive genetic variance. No significant reciprocal differences were found between the reciprocal crosses. The variance/covariance graphical analysis indicated the partial dominance of frost sensitivity. Frost sensitive varieties had the largest number of dominant genes, while frost resistant varieties had the highest proportion of recessive genes. The magnitude of the additive component of variation was higher than that of the dominance component, and the overall measure of the degree of dominance was smaller than one, so average dominance is incomplete. The increasing and decreasing alleles are not equally frequent at all loci. In this set of wheat varieties the values of narrow and broad heritability are relatively high.F2 monosomic analysis of the winter wheat variety 'Arthur' crossed with the monosomics of 'Chinese Spring' revealed that the average frost resistance of all the 21 monosomics was lower than that of the disomic. F2 monosomic hybrids 5A, 2B, 4B and 5D proved to be relatively frost resistant, while monosomics 3A, 3B and 6D were the most sensitive.The control of frost resistance in the set of chromosome substitution lines of the variety 'Cheyenne' into 'Chinese Spring' (with the exception of 2B) indicated that the genes responsible for the frost resistance of 'Cheyenne' are localised in chromosomes 5A, 7A, 4B, 5B, 4D and 5D.The genetic basis of frost resistance and problems of analysis are discussed.
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