MAT heterozygosity and the second sterility barrier in the reproductive isolation of Saccharomyces species

The genetic analysis of large numbers of Saccharomyces cerevisiae × S. uvarum (“cevarum”) and S. kudriavzevii × S. uvarum (“kudvarum”) hybrids in our previous studies revealed that these species are isolated by a postzygotic double-sterility barrier. We proposed a model in which the first barrier is due to the abruption of the meiotic process by the failure of the chromosomes of the subgenomes to pair (and recombine) in meiosis and the second barrier is assumed to be the result of the suppression of mating by allospecific MAT heterozygosity. While the former is analogous to the major mechanism of postzygotic reproductive isolation in plants and animals, the latter seems to be Saccharomyces specific. To bolster the assumed involvement of MAT in the second sterility barrier, we produced synthetic alloploid two-species cevarum and kudvarum hybrids with homo- and heterothallic backgrounds as well as three-species S. cerevisiae × S. kudvarum × S. uvarum (“cekudvarum”) hybrids by mass-mating and examined their MAT loci using species- and cassette-specific primer pairs. We found that the allospecific MAT heterozygosity repressed MAT switching and mating in the hybrids and in the viable but sterile spores produced by the cevarum hybrids that had increased (allotetraploid) genomes. The loss of heterozygosity by meiotic malsegregation of MAT-carrying chromosomes in the latter hybrids broke down the sterility barrier. The resulting spores nullisomic for the S. uvarum chromosome produced vegetative cells capable of MAT switching and conjugation, opening the way for GARMe (Genome Autoreduction in Meiosis), the process that leads to chimeric genomes.

Frequency and fidelity of alien chromosome transmission in Gossypium hexaploid bridging populations

The Australian diploid Gossypium species possess traits of potential agronomical value, such as gossypol-free seeds and Fusarium wilt resistance. However, they belong to the tertiary germplasm pool, which is the most difficult group of species from which to introgress genes into G. barbadense L. and G. hirsutum L. Interspecific triploid hybrids can be generated but they are sterile. The sterility barrier can be overcome using synthetic polyploids as introgression bridges, but whether there is sufficient homoeologous chromosome interaction at meiosis to allow recombination is still an open question. To ascertain, genetically, observable levels of homoeologous introgression, 2 synthetic hexaploid lines (2x G. hirsutum × G. australe and 2x G. hirsutum × G. sturtianum) were crossed to G. hirsutum to generate pentaploid F1 plants that, in turn, were backcrossed to G. hirsutum to generate BC1 and BC2 multiple alien chromosome addition lines (MACALs). Gossypium australe F. Muell. and G. sturtianum Willis chromosome-specific markers were used to track the frequency and fidelity of chromosome transmission to the BC1 and BC2 MACALs. The chromosomal location of the AFLP markers was determined by their distribution among the MACALs and confirmed in parental F2 families. Roughly half the available chromosomes were transmitted to the G. hirsutum × G. australe (54%) and G. hirsutum × G. sturtianum (52%) BC1 MACALs. The BC2 MACAL families again inherited about half of the available chromosomes. There were, however, notable exceptions for specific chromosomes. Some chromosomes were preferentially eliminated, while others were preferentially transmitted. Consistent with the genomic stability of Gossypium synthetic polyploids, the de novo loss or gain of AFLP fragments was rarely observed. While restructuring of the donor G. australe and G. sturtianum chromosomes was observed, this is more likely the result of chromatin loss, and no clear cases of introgression of donor chromatin into the recipient G. hirsutum genome were observed.

Chromosomal and genic barriers to introgression in Helianthus.

The sexual transfer of genes between taxa possessing different structural karyotypes must involve the passage of genes through a chromosomal sterility barrier. Yet little is known about the effects of structural differences on gene introgression within or adjacent to the rearranged chromosomal fragments or about the patterns of introgression in collinear regions. Here, we employ 197 mapped molecular markers to study the effects of chromosomal structural differences on introgression in backcrossed progeny of the domesticated sunflower, Helianthus annuus, and its karyotypically divergent wild relative, H. petiolaris. Forty percent of the genome from the seven collinear linkages introgressed, whereas only 2.4% of the genome from the 10 rearranged linkages was transferred. Thus, chromosomal rearrangements appear to provide an effective mechanism for reducing or eliminating introgression in rearranged chromosomal segments. On the other hand, observations that 60% of the markers from within the collinear portion of the genome did not introgress suggests that genic factors also resist introgression in Helianthus. That is, selection against H. petiolaris genes in concert with linkage may have reduced or eliminated parts of the genome not protected by structural changes. Thus, barriers to introgression in Helianthus appear to include both chromosomal structural and genic factors.

Mating biology in Peniophora cinerea (Basidiomycetes)

Mating tests were performed to analyze the genetic relationship between two intersterile sibling species in Peniophora cinerea (Fr.) Cooke in Europe. Two newly collected specimens from North Europe were found to be compatible with both sibling species, which strongly suggests a close genetic relationship and a sterility barrier of simple genetic origin. The two sibling species, which differ in their substrate selectivity, are accepted as subspecies. One subspecies is restricted to decorticated wood of Fagus, and occasionally the fruit bodies are associated with insect galls. Intersterility was also found in some combinations with two other specimens from Canada and Turkey, but no linkage was found with a particular substrate. Specimens from Taiwan were found to be partially compatible with specimens from Europe, Turkey, and Canada. Distinct differences between the subspecies were found in banding patterns from isoelectric focusing of buffer-soluble mycelial proteins. It is proposed that the kind of intersterility found here is intraspecific and should be looked upon as part of a propagation strategy. Key words: speciation, evolution, Basidiomycetes, isoelectric focusing, insect gall, mating test.

CYTOTAXONOMIC STUDIES ON EREMOPYRUM

Five species are included in the genus Eremopyrum, namely, E. triticeum, E. hirsutum, E. buonapartis, E. orientale, and E. distans. Of these, the first two are diploids with 2n = 14, while the rest are tetraploids with 2n = 28. The karyotypes of the two diploid species are different from those of any other member, of the Triticineae in that they have chromosomes with extremely subterminal centromeres. The karyotype of each of the three tetraploid species shows a mixture of two types of chromosomes: seven pairs with extreme subterminal centromeres and the other seven pairs with median or submedian centromeres. This indicates that these are amphidiploids between triticeum–hirsutum-like taxa and perhaps some diploid species of crested Agropyron. In view of the karyotypic peculiarity, annual habit, and reported sterility barrier, the decision to give Eremopyrum generic status is supported.