The Deletion Stocks of Common Wheat

The construction of comparative genetic maps of chromosomes 4Am and 5Am of Triticum monococcum and chromosomes of homoeologous groups 4, 5 and 7 of T. aestivum has provided insight into the evolution of these chromosomes. The structures of chromosomes 4A, 5A and 7B of modern-day hexaploid bread wheat can be explained by a 4AL/5AL translocation that occurred at the diploid level and is present both in T. monococcum and T. aestivum. Three further rearrangements, a 4AL/7BS translocation, a pericentric inversion and a paracentric inversion, have taken place in the tetraploid progenitor of hexaploid wheat. These structural rearrangements and the evolution of chromosomes 4A, 5A and 7B of bread wheat are discussed. The presence of the 4AL/5AL translocation in several Triticeae genomes raises two questions — which state is the more primitive, and is the translocation of mono- or poly-phylogenetic origin?

/pdf/structural-evolution-of-wheat-chromosomes-4a-5a-and-7b-and-1hh34sgbe5.pdf

Structural evolution of wheat chromosomes 4A, 5A, and 7B and its impact on recombination

Several studies have indicated a noncorrespondence between genetic and physical distances in wheat chromosomes. To study the physical distribution of recombination, polymorphism for C-banding patterns was used to monitor recombination in 67 segments in 11 B-genome chromosome arms of Triticum turgidum. Recombination was absent in proximal regions of all chromosome arms; its frequency increased exponentially with distance from the centromere. A significant difference was observed between the distribution of recombination in physically short and physically long arms. In physically short arms, recombination was almost exclusively concentrated in distal segments and only those regions were represented in their genetic maps. In physically long arms, while a majority of the genetic distance was again based upon recombination in distal chromosome segments, some interstitial recombination was observed. Consequently, these regions also contributed to the genetic maps. Such a pattern of recombination, skewed toward terminal segments of chromosomes, is probably a result of telomeric pairing initiation and strong positive chiasma interference. Interference averaged 0.81 in 35 pairs of adjacent segments and 0.57 across the entire recombining portions of chromosome arms. The total genetic map lengths of the arms corresponded closely to those expected on the basis of their metaphase-I chiasma frequencies. As a consequence of this uneven distribution of recombination there can be a 153-fold difference (or more) in the number of DNA base pairs per unit (centiMorgan) of genetic length.

Physical distribution of recombination in B-genome chromosomes of tetraploid wheat.

Understanding the distribution of crossovers along chromosomes is crucial to evolutionary genomics because the crossover rate determines how strongly a genome region is influenced by natural selection on linked sites. Nevertheless, generalities in the chromosome-scale distribution of crossovers have not been investigated formally. We fill this gap by synthesizing joint information on genetic and physical maps across 62 animal, plant and fungal species. Our quantitative analysis reveals a strong and taxonomically widespread reduction of the crossover rate in the centre of chromosomes relative to their peripheries. We demonstrate that this pattern is poorly explained by the position of the centromere, but find that the magnitude of the relative reduction in the crossover rate in chromosome centres increases with chromosome length. That is, long chromosomes often display a dramatically low crossover rate in their centre, whereas short chromosomes exhibit a relatively homogeneous crossover rate. This observation is compatible with a model in which crossover is initiated from the chromosome tips, an idea with preliminary support from mechanistic investigations of meiotic recombination. Consequently, we show that organisms achieve a higher genome-wide crossover rate by evolving smaller chromosomes. Summarizing theory and providing empirical examples, we finally highlight that taxonomically widespread and systematic heterogeneity in crossover rate along chromosomes generates predictable broad-scale trends in genetic diversity and population differentiation by modifying the impact of natural selection among regions within a genome. We conclude by emphasizing that chromosome-scale heterogeneity in crossover rate should urgently be incorporated into analytical tools in evolutionary genomics, and in the interpretation of resulting patterns.

https://onlinelibrary.wiley.com/doi/pdf/10.1111/mec.14699

Meta-analysis of chromosome-scale crossover rate variation in eukaryotes and its significance to evolutionary genomics.

During meiosis, homologous chromosomes are sorted into pairs and are then intimately aligned, or synapsed, along their lengths while a proteinaceous structure, the synaptonemal complex, is assembled between them. However, little is known about how chromosomes first recognise each other. Here we show, by comparing the behaviour of wild-type wheat and wheat mutant for Ph1 (a suppressor of homeologous chromosome pairing), that when chromosomes recognise a partner to pair with, a conformational change to the chromatin is triggered in both partners that is followed by their intimate alignment [corrected]. Thus, a conformational change in the chromosomes at the onset of meiosis can be correlated directly with recognition.

/pdf/homologue-recognition-during-meiosis-is-associated-with-a-7g5r9bb2h5.pdf

Homologue recognition during meiosis is associated with a change in chromatin conformation.

Metaphase I pairing of deficient chromosomes was analyzed in a set of 'Chinese Spring' (CS) wheat (Triticum aestivum L. em. Thell.) plants with varying lengths of deficiencies in the long arm of chromosome 4A (6, 8, 11, 17, 23, 34, 36, 39, and 50% missing), the long arm of chromosome 5B (49% missing), and the long arm of chromosome 2B (33% missing). Pairing in homologous chromosomes between deficient and complete arms was greatly reduced even by small differences in arm length. In deficiency homozygotes and in an isochromosome derived from a deficient 4AL arm, pairing of the two deficient arms was high and approached that of two complete arms. In plants where deficient and complete arms competed for pairing partners, pairing was exclusively between arms of the same length. These results suggest that in wheat, pairing initiation sites are distributed throughout at least the distal halves of the arms and that the alignment of telomeres may be critical for pairing success. Genetic mapping of the deficiency b...

M. Chrzastek

Papers

Metaphase I pairing of deficient chromosomes and genetic mapping of deficiency breakpoints in common wheat