Genomic analysis in the genus Aegilops.

Summary Hybrids between different Aegilops species and Secale cereale were studied at metaphase I by means of a C-banding technique. On the basis of differences in the C-banding patterns of some of the chromosomes of these hybrids it was possible to carry out an accurate analysis of several types of Aegilops-Aegilops and Aegilops-Secale chromosome associations and, consequently, to establish intraspecific and intergeneric genome relationships. Genomes present in the majority of polyploid Aegilops species are shown to maintain similar patterns of evolutionary affinity to those reported for their proposed diploid parents although in some species there are differences indicating either that differentiations occurred during the evolution of the polyploid species or, on the contrary, that the diploid donors proposed are not the correct ones. On the other hand, differences in the relationships not only between the R genome and different Aegilops genomes but also among different homoeologous groups have been found.     

A chromosome-specific sequence common to the B genome of polyploid wheat and Aegilops searsii

A low-copy, non-coding chromosome-specific DNA sequence, isolated from common wheat, was physically mapped to the distal 19% region of the long arm of chromosome 3B (3BL) of common wheat. This sequence, designated WPG118, was then characterized by Southern hybridization, PCR amplification and sequence comparison using a large collection of polyploid wheats and diploid Triticum and Aegilops species. The data show that the sequence exists in all polyploid wheats containing the B genome and absent from those containing the G genome. At the diploid level, it exists only in Ae. searsii, a diploid species of section Sitopsis, and not in other diploids including Ae. speltoides, the closest extant relative to the donor of the B genome of polyploid wheat. This finding may support the hypothesis that the B-genome of polyploid wheat is of a polyphyletic origin, i.e. it is a recombined genome derived from two or more diploid Aegilops species.     

Restriction Fragment Length Polymorphism (RFLP) for protein disulfide isomerase (PDI) gene sequences in Triticum and Aegilops species

RFLP variation revealed by protein disulfide isomerase (PDI) coding gene sequences was assessed in 170 accessions belonging to 23 species of Triticum and Aegilops. PDI restriction fragments were highly conserved within each species and confirmed that plant PDI is encoded either by single-copy sequences or by small gene families. The wheat PDI probe hybridized to single EcoRI or HindIII fragments in different diploid species and to one or two fragments per genome in polyploids. Four Aegilops species in the Sitopsis section showed complex patterns and high levels of intraspecific variation, whereas Ae. searsii possessed single monomorphic fragments. T. urartu and Ae. squarrosa showed fragments with the same mobility as those in the A and D genomes of Triticum polyploid species, respectively, whereas differences were observed between the hybridization patterns of T. monococcum and T. boeoticum and that of the A genome. The single fragment detected in Ae. squarrosa was also conserved in most accessions of polyploid Aegilops species carrying the D genome. The five species of the Sitopsis section showed variation for the PDI hybridization fragments and differed from those of the B and G genomes of emmer and timopheevi groups of wheat, although one of the Ae. speltoides EcoRI fragments was similar to those located on the 4B and 4G chromosomes. The similarity between the EcoRI fragment located on the 1B chromosome of common and emmer wheats and one with a lower hybridization intensity in Ae. longissima, Ae. bicornis and Ae. sharonensis support the hypothesis of a polyphyletic origin of the B genome. Received: 25 June 1999 / Accepted: 14 September 1999     

RAPD analysis of the intraspecific and interspecific variation and phylogenetic relationships of <Emphasis Type="Italic">Aegilops</Emphasis> L. species with the U genome

RAPD analysis was used to study the genetic variation and phylogenetic relationships of polyploid Aegilops species with the U genome. In total, 115 DNA samples of eight polyploid species containing the U genome and the diploid species Ae. umbellulata (U) were examined. Substantial interspecific polymorphism was observed for the majority of the polyploid species with the U genome (interspecific differences, 0.01–0,2; proportion of polymorphic loci, 56.6–88.2%). Aegilops triuncialis was identified as the only alloploid species with low interspecific polymorphism (interspecific differences, 0–0.01, P = 50%) in the U-genome group. The U-genome Aegilops species proved to be separated from other species of the genus. The phylogenetic relationships were established for the U-genome species. The greatest separation within the U-genome group was observed for the US-genome species Ae. kotschyi and Ae. variabilis. The tetraploid species Ae. triaristata and Ae. columnaris, which had the UX genome, and the hexaploid species Ae. recta (UXN) were found to be related to each other and separate from the UM-genome species. A similarity was observed between the UM-genome species Ae. ovata and Ae. biuncialis, which had the UM genome, and the ancestral diploid U-genome species Ae. umbellulata. The UC-genome species Ae. triuncialis was rather separate and slightly similar to the UX-genome species.     

Syntenic Relationships between the U and M Genomes of Aegilops,Wheat and the Model Species Brachypodium and Rice as Revealed by COS Markers

Diploid Aegilops umbellulata and Ae. comosa and their natural allotetraploid hybrids Ae. biuncialis and Ae. geniculata are important wild gene sources for wheat. With the aim of assisting in alien gene transfer, this study provides gene-based conserved orthologous set (COS) markers for the U and M genome chromosomes. Out of the 140 markers tested on a series of wheat-Aegilops chromosome introgression lines and flow-sorted subgenomic chromosome fractions, 100 were assigned to Aegilops chromosomes and six and seven duplications were identified in the U and M genomes, respectively. The marker-specific EST sequences were BLAST-ed to Brachypodium and rice genomic sequences to investigate macrosyntenic relationships between the U and M genomes of Aegilops, wheat and the model species. Five syntenic regions of Brachypodium identified genome rearrangements differentiating the U genome from the M genome and from the D genome of wheat. All of them seem to have evolved at the diploid level and to have been modified differentially in the polyploid species Ae. biuncialis and Ae. geniculata. A certain level of wheat–Aegilops homology was detected for group 1, 2, 3 and 5 chromosomes, while a clearly rearranged structure was showed for the group 4, 6 and 7 Aegilops chromosomes relative to wheat. The conserved orthologous set markers assigned to Aegilops chromosomes promise to accelerate gene introgression by facilitating the identification of alien chromatin. The syntenic relationships between the Aegilops species, wheat and model species will facilitate the targeted development of new markers specific for U and M genomic regions and will contribute to the understanding of molecular processes related to allopolyploidization.     

The molecular basis of genetic diversity among cytoplasms of Triticum and Aegilops

Summary Many related species and strains of common wheat were compared by matching differences among their mitochondrial genomes with their parent nuclear genomes. We examined three species of Aegilops, section Sitopsis (Ae. bicornis, Ae. sharonensis, and Ae. speltoides), emmer wheat (Triticum dicoccoides, T. dicoccum, and T. durum), common wheat (T. spelta, T. aestivum, and T. compaction), and timopheevi wheat (T. araraticum, T. timopheevi, and T. zhukovskyi). A single source of the cytoplasm was used in all the species, except Ae. speltoides (two sources), T. araraticum (two), and T. aestivum (three). Following restriction endonuclease analyses, the mitochondrial genomes were found to comprise seven types, and a dendrogram showing their genetic relatedness was constructed, based upon the percentage of common restriction fragments. MtDNAs from T. dicoccum, T. durum, T. aestivum, and T. compactum yielded identical restriction fragment patterns; these differed from T. dicoccoides and T. spelta mtDNAs in only 2.3% of their fragments. The fragment patterns of T. timopheevi and T. zhukovskyi were identical, and these differed from T. araraticum mtDNA by only one fragment. In both the emmer-dinkel and timopheevi groups, mitochondrial genome differentiation is evident, suggesting a diphyletic origin of each group. MtDNAs from four accessions of the Sitopsis species of Aegilops differ greatly from one another, but those of Ae. bicornis, Ae. sharonensis, and Ae. searsii, belonging to the same subsection Emarginata, are relatively similar. MtDNAs of timopheevi species are identical, or nearly so, to those of Ae. speltoides accession (09), suggesting that the latter was the cytoplasm donor to the former, polyploid group. The origin of this polyploid group seems to be rather recent in that the diploid and polyploid species possess nearly identical mitochondrial genomes. We cannot determine, with precision, the cytoplasm donor to the emmer-dinkel group. However, our results do suggest that mitochondrial DNAs show larger evolutionary divergence than do the ctDNAs from these same strains.Contribution no. 507 from the Laboratory of Genetics, Faculty of Agriculture, Kyoto University, Japan     

The pattern of zygotene and pachytene pairing in allotetraploid Aegilops species sharing the D genome

Chromosome pairing behaviour of the allotetraploid Aegilops species sharing the D genome, Ae. crassa (DDMM), Ae. cylindrica (DDCC) and Ae. ventricosa (DDNN), was analyzed by electron microscopy in surfacespread prophase-I nuclei. Synaptonemal-complex analysis at zygotene and pachytene revealed that synapsis in the allotetraploids was mostly between homologous chromosomes, although a few multivalents were also formed. Only homologous bivalents were observed at metaphase-I. It is concluded that the mechanism controlling bivalent formation in these species acts mainly at zygotene by restricting pairing to homologous chromosomes, but also acts at pachytene by preventing chiasma formation in homoeologous associations. These observations are discussed in relation to mechanisms of diploidization of polyploid meiosis.     

NADP-dependent aromatic alcohol dehydrogenase in polyploid wheats and their diploid relatives. On the origin and phylogeny of polyploid wheats

Summary The three major isoenzymes of the NADP-dependent aromatic alcohol dehydrogenase (ADH-B), distinguished in polyploid wheats by means of polyacrylamide gel electrophoresis, are shown to be coded by homoeoalleles of the locus Adh-2 on short arms of chromosomes of the fifth homoeologous group. Essentially codominant expression of the Adh-2 homoeolleles of composite genomes was observed in young seedlings of hexaploid wheats (T. aestivum s.l.) and tetraploid wheats of the emmer group (T. turgidum s.l.), whereas only the isoenzyme characteristic of the A genome is present in the seedlings of the timopheevii-group tetraploids (T. timopheevii s.str. and T. araraticum).The slowest-moving B³ isoenzyme of polyploid wheats, coded by the homoeoallele of the B genome, is characteristic of the diploid species Aegilops speltoides S.l., including both its awned and awnless forms, but was not encountered in Ae. bicornis, Ae. sharonensis and Ae. longissima. The last two diploids, as well as Ae. tauschii, Ae. caudata, Triticum monococcum s.str., T. boeoticum s.l. (incl. T. thaoudar) and T. urartu all shared a common isoenzyme coinciding electrophoretically with the band B² controlled by the A and D genome homoeoalleles in polyploid wheats. Ae. bicomis is characterized by the slowest isoenzyme, B⁴, not found in wheats and in the other diploid Aegilops species studied.Two electrophoretic variants of ADH-B, B¹ and B², considered to be alloenzymes of the A genome homoeoallele, were observed in T. dicoccoides, T. dicoccon, T. turgidum. s.str. and T. spelta, whereas B² was characteristic of T. timopheevii s.l. and only B¹ was found in the remaining taxa of polyploid wheats. The isoenzyme B¹, not encountered among diploid species, is considered to be a mutational derivative which arose on the tetraploid level from its more ancestral form B² characteristic of diploid wheats.The implication of the ADH-B isoenzyme data to the problems of wheat phylogeny and gene evolution is discussed.     

Standard karyotypes ofAegilops uniaristata,Ae. mutica,Ae. comosa subspeciescomosa andheldreichii (Poaceae)

C-banding patterns and polymorphisms were analyzed in several accessions of the diploidAegilops speciesAe. uniaristata, Ae. mutica, andAe. comosa subsp.comosa and subsp.heldreichii, and standard karyotypes of these species were established. Variation in C-band size and location was observed between different accessions, but did not prevent chromosome identification. One accession ofAe. uniaristata was homozygous for whole-arm translocations involving chromosomes 1N and 5N. The homoeologous relationships of these chromosomes were established by comparison of chromosome morphologies and C-banding patterns to other diploidAegilops species with known chromosome homoeology. In addition, in situ hybridization analysis with a 5S rDNA probe was used to identify homoeologous groups 1 and 5 chromosomes. The present analysis permitted the assignment of allAe. mutica, comosa subsp.comosa, andAe. comosa subsp.heldreichii chromosomes, and three of the sevenAe. uniaristata chromosomes according to their homoeologous groups. The data presented will be useful analyzing genome differentiation in polyploidAegilops species.     

BAC libraries of Triticum urartu, Aegilops speltoides and Ae. tauschii, the diploid ancestors of polyploid wheat

Triticum urartu, Aegilops speltoides and Ae. tauschii are respectively the immediate diploid sources, or their closest relatives, of the A, B and D genomes of polyploid wheats. Here we report the construction and characterization of arrayed large-insert libraries in a bacterial artificial chromosome (BAC) vector, one for each of these diploid species. The libraries are equivalent to 3.7, 5.4 and 4.1 of the T. urartu, Ae. speltoides, Ae. tauschii genomes, respectively. The predicted levels of genome coverage were confirmed by library hybridization with single-copy genes. The libraries were used to estimate the proportion of known repeated nucleotide sequences and gene content in each genome by BAC-end sequencing. Repeated sequence families previously detected in Triticeae accounted for 57, 61 and 57% of the T. urartu, Ae. speltoides and Ae. tauschii genomes, and coding regions accounted for 5.8, 4.5 and 4.8%, respectively.     

Ecological distribution and species diversity of Aegilops L. genus in Bulgaria

The genus Aegilops has an important potential utilization in wheat improvement because of its resistance to different biotic and abiotic stresses and close relation with the cultivated wheat. Therefore, a better knowledge of the eco-geographical distribution of Aegilops species and their collection and conservation are required. A total of 297 Aegilops accessions representing nine (five tetraploid and four diploid) species were collected in different regions of Bulgaria, and the ecological characteristics of the 154 explored sites were recorded. The distribution of the diploid species (Ae. caudata L., Ae. speltoides Tausch, Ae. umbellulata Zhuk. and Ae. comosa Sibth. and Sm.) was limited to specific environments in south-central Bulgaria. Tetraploid species were present in harsher environments than diploid species and showed wider adaptation and distribution. Species–environment relationships were analysed by considering the worldwide distribution of the species and their physiological resistance to abiotic stress. Aegilops cylindrica Host was more frequently found in northern Bulgaria and at high altitudes. Its distribution was closely related to its tolerance to low temperatures. Aegilops geniculata Roth and Ae. neglecta Req. ex Bertol. were absent in the north of Bulgaria, but widely distributed in low rainfall areas. Aegilops neglecta, more frost resistant than Ae. geniculata, was present at higher altitude. Aegilops biuncialis Vis. and Ae. triuncialis L. showed adaptation to a wide range of climatic conditions. The study of Aegilops species ecology and distribution in Bulgaria provided useful information for the future collection and for the genetic resource management in this region.     

Insight into the genetic variability analysis and relationships among some Aegilops and Triticum species,as genome progenitors of bread wheat,using SCoT markers

We assessed the molecular genetic diversity and relationships among some Aegilops and Triticum species using 15 start codon-targeted (SCoT) polymorphism markers. A total of 166 bands amplified, of which 164 (98.79%) were polymorphic. Analysis of molecular variance and inter-population differentiation (Gst) indicated high genetic variation within the studied populations. Our analyses revealed high genetic diversity in T. boeoticum, Ae. cylindrica, T. durum and Ae. umbellulata, low diversity in Ae. crassa, Ae. caudata and Ae. speltoides, and a close relationship among Ae. tauschii, T. aestivum, T. durum, T. urartu, and T. boeoticum. Cluster analysis indicated 180 individuals divided into 8 genome homogeneous clades and 11 sub-groups. T. aestivum and T. durum accessions were grouped together, and accessions with the C and U genomes were grouped into the same clade. Our results support the hypothesis that T. urartu and Ae. tauschii are two diploid ancestors of T. aestivum, and also that Ae. caudata and Ae. umbellulata are putative donors of C and U genomes for other Aegilops species that possess these genomes. Our results also revealed that the SCoT technique is informative and can be used to assess genetic relationships among wheat germplasm.     

Amylase protein inhibitors and the role of Aegilops species in polyploid wheat speciation

Protein inhibitors extracted with water from seeds of Triticum and genetically related species were characterized according to their apparent molecular weights, electrophoretic mobilities and their specificities in inhibiting α-amylases from human saliva and Tenebrio molitor L. larvae. No detectable amylase inhibition activity was found in extracts from diploid wheats, whereas in all tetraploid and hexaploid wheats as well as in the Aegilops species tested we found several amylase inhibitor groups of different molecular weights. In each group, several inhibitor components slightly different in their electrophoretic mobilities, but identical in their inhibition behaviour toward amylases from different origins have been shown. Both from the qualitative and quantitative standpoints, amylase protein inhibitors from hexaploid wheats were the summation of those from tetraploid wheats plus the ones from Aegilops squarrosa. Amylase inhibitors from Aegilops speltoides largely differed from those extracted from tetraploid wheats as well as from all the amylase inhibitors described in plant seeds up to now. These results indicate a relevant homology between the amylase inhibitor coding genes of the D wheat genome and those of the D Aegilops genome and confirm that Ae. squarrosa is the donor of the whole D genome to hexaploid wheats. They also suggest that Ae. speltoides is not the donor of the B genome to polyploid wheats, although a not yet identified Aegilops species might be such a donor.     

On the diploidization mechanism of the genus Aegilops: meiotic behaviour of interspecific hybrids

Chromosomal pairing of the three diploid hybrids Aegilops uniaristata × Ae. tauschii (ND), Ae. umbellulata × Ae. tauschii (UD) and Ae. comosa × Ae. uniaristata (MN), and a triploid hybrid Ae. cylindrica × Ae. caudata (DCC), was analyzed by electron microscopy in surface-spread-prophase-I nuclei and compared with light-microscopic observations of metaphase-I cells after C-banding and fluorescence in situhybridization. All hybrids showed extensive synapsis and complex multivalents in which up to 14 chromosomes were involved. In the diploid hybrids most metaphase-I chromosomal associations were between homoeologs, their frequencies being dependent on the relationship between the donor genomes. Despite the different overall bound-arm frequencies displayed by ND and MN hybrids at metaphase-I, chromosomes bearing rDNA sequences showed similar mean cell chromosomal association frequencies. In the triploid hybrid preferential associations involving C genomes were predominant. These observations are discussed in relation to the mechanism of diploidization showed by allotetraploid Aegilops species. Received: 13 January 1999 / Accepted: 12 March 1999     

Phylogenetic Relationships and Intraspecific Variation of D-Genome Aegilops L. as Revealed by RAPD Analysis

RAPD analysis was carried out to study the genetic variation and phylogenetic relationships of polyploid Aegilops species, which contain the D genome as a component of the alloploid genome, and diploid Aegilops tauschii, which is a putative donor of the D genome for common wheat. In total, 74 accessions of six D-genome Aegilops species were examined. The highest intraspecific variation (0.03–0.21) was observed for Ae. tauschii. Intraspecific distances between accessions ranged 0.007–0.067 in Ae. cylindrica, 0.017–0.047 in Ae. vavilovii, and 0–0.053 inAe. juvenalis.Likewise, Ae. ventricosaand Ae. crassa showed low intraspecific polymorphism. The among-accession difference in alloploidAe. ventricosa (genome D^vN^v) was similar to that of one parental species, Ae. uniaristata (N), and substantially lower than in the other parent, Ae. tauschii (D). The among-accession difference in Ae. cylindrica(C^cD^c) was considerably lower than in either parent, Ae. tauschii (D) orAe. caudata (C). With the exception of Ae. cylindrica, all D-genome species—Ae. tauschii (D),Ae. ventricosa (D^vN^v), Ae. crassa (X^crD^cr1 and X^crD^cr1D^cr2), Ae. juvenalis (X^jD^jU^j), andAe. vavilovii (X^vaD^vaS^va)—formed a single polymorphic cluster, which was distinct from clusters of other species. The only exception, Ae. cylindrica(C^cD^c), did not group with the other D-genome species, but clustered withAe. caudata (C), a donor of the C genome. The cluster of these two species was clearly distinct from the cluster of the other D-genome species and close to a cluster of Ae. umbellulata (genome U) and Ae. ovata (genome U^gM^g). Thus, RAPD analysis for the first time was used to estimate and to compare the interpopulation polymorphism and to establish the phylogenetic relationships of all diploid and alloploid D-genome Aegilops species.     

The pattern of zygotene and pachytene pairing in allotetraploid Aegilops species sharing the U genome

Allotetraploid Aegilops species sharing the U genome, Ae. columnaris (UUMM), Ae. ovata (UUMM), Ae. triaristata (UUMM), Ae. triuncialis (UUCC) and Ae. variabilis (UUSS), regularly form bivalents at metaphase I of meiosis. The pattern of zygotene and pachytene pairing was analyzed by whole-mount surface-spreading of synaptonemal complexes under the electron microscope. The data indicated that at the zygotene stage the chromosomes were almost exclusively associated as bivalents; only a few multivalents (7%) were observed. These observations are discussed in relation to mechanisms of diploidization of polyploid meiosis.     

Variation in the distribution of a genome-specific DNA sequence on chromosomes reveals evolutionary relationships in the Triticum and Aegilops complex

 The present study analyzed the distribution pattern of the Ae. speltoides–derived repetitive clone pGc1R-1 in the Triticum/Aegilops complex. Fluorescence in situ hybridization analysis showed that clone pGc1R-1 is a S-genome-specific repetitive sequence that hybridized to the S-genome of three species in the section Sitopsis, Aegilops speltoides (S), Ae. longissima (S^l), and Ae. sharonensis (S^sh), but not to Ae. bicornis (S^b) and Ae. searsii (S^s), nor to any other diploid Aegilops species. This clone also hybridized to the very closely related G-genome of T. timopheevii subsp. armeniacum and T. timopheevii ssp. timopheevii, but not to the B-genome of T. turgidum and T. aestivum. Hybridization also was observed in the polyploid Aegilops species, Ae. kotschyi (U^kS^k), Ae. peregrina (U^pS^p), and Ae. vavilovii (X^vaD^vaS^va). Large inter- and intraspecific variations were observed. Our results confirm that the S genome is related more to the S^l and S^sh genomes than to the S^b and S^s genomes; there is a greater affinity between the G and S genomes than between the B and S genomes. Mechanisms to account for the variation in the FISH pattern with different genomes include sequence amplification and deletion. Variation in the distribution of this genome-specific DNA sequence, pGc1R-1, on chromosomes can be used to reveal evolutionary relationships in the Triticum and Aegilops complex. Received April 10, 2002; accepted July 12, 2002 Published online: November 28, 2002 Address of the authors: Peng Zhang, Bernd Friebe (e-mail: friebe@ksu.edu), Bikram S. Gill, Wheat Genetics Resource Center, Department of Plant Pathology, 4024 Throckmorton, Plant Sciences Center, Kansas State University, Manhattan, KS 66506-5502, USA.     

Molecular characterization of novel low-molecular-weight glutenin genes inAegilops longissima

Extensive genetic variations of low-molecular-weight glutenin subunits (LMW-GS) and their coding genes were found in the wild diploid A- and D-genome donors of common wheat. In this study, we reported the isolation and characterization of 8 novel LMW-GS genes fromAe.longissima Schweinf. & Muschl., a species of the sectionSitopsis of the genusAegilops, which is closely related to the B genome of common wheat. Based on the N-terminal domain sequences, the 8 genes were divided into 3 groups. A consensus alignment of the extremely conserved domains with known gene groups and the subsequent cluster analysis showed that 2 out of the 3 groups of LMW-GS genes were closely related to those from the B genome, and the remaining was related to those from A and D genomes of wheat andAe. tauschii. Using 3 sets of gene-group-specific primers, PCRs in diploid, tetraploid and hexaploid wheats andAe. tauschii failed to obtain the expected products, indicating that the 3 groups of LMW-GS genes obtained in this study were new members of LMW-GS multi-gene families. These results suggested that theSitopsis species of the genusAegilops with novel gene variations could be used as valuable gene resources of LMW-GS. The 3 sets of group-specific primers could be utilized as molecular markers to investigate the introgression of novel alien LMW-GS genes fromAe. longissima into wheat.     

Metaphase I-bound arms frequency and genome analysis in wheat-Aegilops hybrids. 3. Similar relationships between the B genome of wheat and S or S l genomes of Ae. speltoides,Ae. longissima and Ae. sharonensis

The meiotic behaviour of Triticum aestivum × Aegilops speltoides, T. aestivum × Ae. sharonensis and T. aestivum × Ae. longissima tetraploid hybrids (genome constitution ABDS, ABDS ^l , and ABDS ^l , respectively) has been analysed by the C-banding technique. Of the six types of pairing normally occurring, at metaphase I three were recognized: A-D, AD-BS/AD-BS ^l and B-S/B-S ^l . The relative order observed in the low pairing hybrid, A-D> B-S ^l >AD-BS ^l , as well as that found in high-pairing Chinese Spring × Ae. speltoides hybrids, A-D>AD-BS>ß-S, revealed the existence of preferential pairing patterns among the different genomes that are in competition. In all of the hybrids analysed the mean number of bound arms per cell for the A-D type was significantly higher than the mean number of associations between the B and S/S ^l genomes. Usually the relative contribution of each type of pairing is maintained among hybrids with different Aegilops species. These results indicate that the genomes of Ae. speltoides, Ae. sharonensis and Ae. longissima show a similar affinity with the genomes of hexaploid wheat; therefore none of these species can be considered to be a distinct donor of the B genome of wheats.     

Genome differentiation in Aegilops. 3. Evolution of the D-genome cluster

 Six polyploid Aegilops species containing the D genome were studied by C-banding and fluorescence in situ hybridization (FISH) using clones pTa71 (18S-5.8S-26S rDNA), pTa794 (5S rDNA), and pAs1 (non-coding repetitive DNA sequence) as probes. The C-banding and pAs1-FISH patterns of Ae. cylindrica chromosomes were identical to those of the parental species. However, inactivation of the NOR on chromosome 5D with a simultaneous decrease in the size of the pTa71-FISH site was observed. The N^v and D^v genomes of Ae. ventricosa were somewhat modified as compared with the N genome of Ae. uniaristata and the D genome of Ae. tauschii. Modifications included minor changes in the C-banding and pAs1-FISH patterns, complete deletion of the NOR on chromosome 5D^v, and the loss of several minor 18S-5.8S-26S rDNA loci on N^v genome chromosomes. According to C-banding and FISH analyses, the D^cr1 genome of Ae. crassa is more similar to the D^v genome of Ae. ventricosa than to the D genome of Ae. tauschii. Mapping of the 18S-5.8S-26S rDNA and 5S rDNA loci by multicolor FISH suggests that the second (X^cr) genome of tetraploid Ae. crassa is a derivative of the S genome (section Emarginata of the Sitopsis group). Both genomes of Ae. crassa were significantly modified as the result of chromosomal rearrangements and redistribution of highly repetitive DNA sequences. Hexaploid Ae. crassa and Ae. vavilovii arose from the hybridization of chromosomal type N of tetraploid Ae. crassa with Ae. tauschii and Ae. searsii, respectively. Chromosomal type T1 of tetraploid Ae. crassa and Ae. umbellulata were the ancestral forms of Ae. juvenalis. The high level of genome modification in Ae. juvenalis indicates that it is the oldest hexaploid species in this group. The occurrence of hexaploid Ae. crassa was accompanied by a species-specific translocation between chromosomes 4D^cr1 and 7X^cr. No chromosome changes relative to the parental species were detected in Ae. vavilovii, however, its intraspecific diversity was accompanied by a translocation between chromosomes 3X^cr and 3D^cr1. Received July 24, 2001 Accepted October 1, 2001