Morphology and cytology of intergeneric hybrids involvingLeymus multicaulis (Poaceae)

The morphology and meiotic behaviour of pollen mother cells were studied in hybrids involvingLeymus multicaulis (2n = 28) ×Psathyrostachys huashanica (2n = 14),L. multicaulis ×P. juncea (2n = 14), andL. secalinus (2n = 28) ×L. multicaulis. Chromosome pairing was almost identical in theL. multicaulis ×P. huashanica, andL. multicaulis ×P. juncea hybrids, in which it averaged 7.30 univalents + 6.69 bivalents + 0.096 trivalents and 7.48 univalents + 6.75 bivalents, respectively. The meiotic pairing in the two hybrids indicated that oneL. multicaulis genome was closely homologous with theP. huashanica andP. juncea genomes. BothP. huashanica andP. juncea are possibly donors of oneL. multicaulis genome. Chromosome pairing in theL. secalinus ×L. multicaulis hybrid averaged 4.49 univalents + 11.71 bivalents + 0.02 trivalents, indicating that the genomes ofL. multicaulis andL. secalinus are to some degree homologous. However, they are sufficiently differentiated to insure species distinctness.     

Relationships between species ofLeymus,Psathyrostachys, andHordeum (Poaceae,Triticeae) inferred from Southern hybridization of genomic and cloned DNA probes

We have used total genomic DNA as a probe to size-fractionated restriction enzyme digests of genomic DNA from a range ofTriticeae species from the generaLeymus Hochst.,Psathyrostachys Nevski, andHordeum L., and hybrids betweenHordeum andLeymus to investigate their taxonomic relationships. Genomic Southern hybridization was found to be an effective and simple way to assess the distribution and diversity of essentially species-specific and common, repetitive DNA sequences, and is hence especially useful in evolutionary studies. The DNA sequences ofH. vulgare seem to diverge substantially from those ofH. brachyantherum, H. lechleri, H. procerum, andH. depressum. The genome ofThinopyron bessarabicum shows little homology to those of theLeymus species investigated, confirming thatT. bessarabicum is not an ancestral genome inLeymus. Although the genomes ofLeymus andPsathyrostachys share substantial proportions of DNA sequences, they include divergent repeated sequences as well. Hybridization with a ribosomal DNA probe (pTa 71) showed that the coding regions containing structural genes encoding the 18 S, 5.8 S, and 26 S ribosomal RNA were conserved among the species investigated, whereas the intergenic spacer region was more variable, presenting different sizes of restriction fragments and enabling a classification of the species. The rye heterochromatin probe pSc 119.2 hybridized to DNA fromH. lechleri andT. bessarabicum, but not to DNA from the other species investigated.     

Investigations of Genome Relationships Between Leymus, Psathyrostachys and Hordeum Inferred by Genomic DNA:DNA in situ Hybridization

Genome relationships between the genera Leymus Hochst., PsathyrostachysNevski and Hordeum L. (Poaceae, Triticeae) were investigatedby fluorescent in situ hybridization using both total genomicDNA and cloned DNA sequences as probes. In hybrids between speciesof Hordeum and Leymus there was a clear differentiation betweenthe H genomes of Hordeum species and the genomes of Leymus speciesafter probing with genomic Hordeum or Leymus DNA. Chromosomesof species of Leymus and Psathyrostachys were also differentiatedby subtelomeric heterochromatic segments or by negative bandsalong their length. The number and location of 18S-5·8S-26SrRNA genes varied between the investigated genera. Unusually,L. angustus and P. stoloniformis rDNA sites were localized onboth ends of some chromosomes. Interphase nuclei of the Hordeumx Leymus hybrids had groups of chromosomes from both parentalgenomes in discrete, non-intermixed domains.Copyright 1994,1999 Academic Press Taxonomy, evolution, molecular evolution, repetitive DNA, rDNA sites, in situ hybridization, Triticeae, Leymus, Hordeum, Psathyrostachys     

Phylogenetic relationships between Leymus (Poaceae,Triticeae) and related diploid Triticeae species based on isozyme and genome-specific random amplified polymorphic DNA (RAPD) markers

Abstract

To investigate the phylogenetic relationships between Leymus and related diploid species of the Triticeae tribe, the esterase isozyme (EST), superoxide dismutase (SOD) isozymes, and genome-specific random amplified polymorphic DNA (RAPD) markers were used to analyze for 14 Leymus species, together with two Psathyrostachys species (Ns), three Pseudoroegneria species (St), two Hordeum species (H), Lophopyrum elongatum (E^e), Australopyrum retrofractum (W), and Agropyron cristatum (P). The data were used to construct dendrograms by means of UPGMA in the NTSYS-pc computer program. The results suggested that (1) isozyme analysis can be used in the systematic studies of these perennial Triticeae; (2) there is a close relationship between Leymus, Psathyrostachys juncea, three Pseudoroegneria species, and Lophopyrum elongatum; (3) the Ns genome-specific RAPD marker was present in all 14 polyploid species of Leymus, while the E^e and P genome-specific RAPD markers were absent in 14 polyploid species of Leymus; the St, W and H genome-specific RAPD markers were present in some species of Leymus; (4) Leymus species have multiple origins, and different Leymus species derived their genomes from different donors.     

Intergeneric Hybridization between Species of Leymus, Psathyrostachys and Hordeum (Poaceae, Triticeae)

A total of 132 intergeneric crossing attempts (49 combinations)involving species of Leymus Hochst., Psathyrostachys Nevskiand Hordeum L. were performed, of which 103 were between Hordeumand Leymus. Embryo rescue was used throughout the experiment.Hybrids between Leymus and Psathyrostachys were difficult toobtain. Hybrid progeny were relatively easily obtained whencrossing Hordeum and Leymus. Plants from 20 different combinationswere obtained. Nineteen of these have not previously been reported.Meiotic analysis of three hybrid combinations of Hordeum x Leymusis reported. The high frequency of univalents in meiotic interphase(MI) indicates that allosyndetic chromosome pairing did notoccur, supporting the assumption that the genomes of Leymusare non-homologous to the H genomes of Hordeum.Copyright 1994,1999 Academic Press Taxonomy, Triticeae, Leymus, Psathyrostachys, Hordeum, intergeneric hybridization     

Hybridisation,genomic constitution and generic delimitation inElymus s. l. (Poaceae: Triticeae)

Intergeneric crosses were made between representatives of the genomically-defined generaElymus, Agropyron, Elytrigia, Pseudoroegneria, andThinopyrum. The genomic constitution ofElytrigia repens, the type species ofElytrigia, is shown to be SSH, a genomic combination otherwise found only inElymus. The S genome ofPseudoroegneria has almost always a dominant influence on the morphology of the taxa of which it is a component.Wang (1989) showed that the J genome inThinopyrum and the S genome have considerable homoeology, with a mean c-value of 0.35 in diploid SJ hybrids. A genetic coherence from S to SJ^e, J^e, J^eJ^b, and J^b can be expected, agreeing with the continuous morphologic variation pattern observed. Because of the absence of morphological discontinuities between the taxa,Pseudoroegneria (S),Elymus (SH, SY, sometimes with additional genomes),Elytrigia (SSH, SSHX), andThinopyrum (SJ, SJJ, J) are best treated as a single genus,Elymus, following the generic concept ofMelderis in Flora Europaea and Flora of Turkey. The basic genomic constituents ofElymus will then be the S and/or J genomes.Agropyron, with diploids, tetraploids, and hexaploids based on the P genome is morphologically distinct from other genera inTriticeae. In a few species ofElymus andPseudoroegneria, a P genome is an additional constituent. In these cases the P genome has a negligible morphological influence. Therefore, it seems reasonable to maintainAgropyron as a separate genus.     

Synthesis and evaluation of Triticum durum — T. monococcum amphiploids

Summary Nine Triticum durum — T. monococcum amphiploids (AABBA^mA^m) were synthesized by chromosome doubling of sterile triploid F₁ hybrids involving nine T. durum (AABB) cultivars and a T. monococcum (A^mA^m) line. The triploid F₁ hybrids had a range of 4–7 bivalents and 7–13 univalents per PMC. The synthetic amphiploids, however, showed a high degree of preferential pairing of chromosomes of the A genomes of diploid and tetraploid wheats. The amphiploids were meiotically stable and fully fertile. Superiority of four amphiploids for tiller number per plant, 100-grain weight, protein content and resistance to Karnal bunt demonstrated that these could either be commercially exploited as such after overcoming certain inherent defects or used to introgress desirable genes into durum and bread wheat cultivars. Methods for improvement of these amphiploids are discussed.     

Multidisciplinary approach to genome analysis in the diploid species,Thinopyrum bessarabicum and Th. elongatum (Lophopyrum elongatum), of the Triticeae

Summary The J and E genome species of the Triticeae are invaluable sources of salt tolerance. The evidence concerning the phyletic relatedness of the J genome of diploid Thinopyrum bessarabicum and the E genome of diploid Th. elongatum (=Lophopyrum elongatum) is discussed. Low level of chromosome pairing between J and E at different ploidy levels, suppression of J-E pairing by the Ph1 pairing regulator that inhibits homoeologous pairing, complete sterility of the diploid hybrids (JE), karyotypic divergence of the two genomes, differences in total content and distribution of heterochromatin along their chromosomes, and marked differences in gliadin proteins, isozymes, 5S DNA, and rDNA indicate that J and E are distinct genomes. Well-defined biochemical markers have been identified in the two genomes and may be useful in plant breeding. The level of distinction between J and E is comparable to that among the universally accepted homoeologous genomes A, B, and D of wheat. Therefore, the J and E genomes are homoeologous and not homologous, although some workers continue to call them homologous. The previous workers' data on chromosome pairing in diploid hybrids and/ or karyotypic differences in the conventionally stained chromosomes do not provide sufficient evidence for the proposed merger of J and E genomes (and, hence, of the genera Thinopyrum and Lophopyrum) specifically and for establishing genome relationships generally. Extra precautions should be exercised before changing the designation of an established genome and before merging two genera. A uniform, standardized system of genomic nomenclature for the entire Triticeae is proposed, which should benefit cytogeneticists, plant breeders, taxonomists, and evolutionists.Cooperative investigations of the USDA-Agricultural Research Service and the Utah Agricultural Experiment Station, Logan, UT 84322, USA. Approved as Journal Paper no. 3832     

Genetic relationships among <Emphasis Type="Italic">Hystrix patula,H. duthiei</Emphasis> and <Emphasis Type="Italic">H. longearistata</Emphasis> according to meiotic studies and genome-specific RAPD assay

Hybrids including Hystrix patula, H. duthiei and H. longearistata were obtained and genetic relationships among them were studied. Meiotic pairing in hybrids of H. duthiei × Psathyrostachys juncea (Ns), H. longearistata × Psa. juncea (Ns), Leymus multicaulis (NsXm) × H. duthiei, L. multicaulis (NsXm) × H. longearistata, Elymus sibiricus (StH) × H. patula, Roegneria ciliaris (StY) × H. patula, R. ciliaris (StY) × H. duthiei and R. ciliaris (StY) × H. longearistata averaged 5.76, 5.44, 11.94, 10.88, 10.08, 3.57, 0.46 and 0.90 bivalents per cell, respectively. The results indicated that H. duthiei and H. longearistata had the NsXm genomes of Leymus, while H. patula contained the StH genomes and had a low genome affinity with the StY genomes of Roegneria. Results of genome-specific RAPD assay were comparable with the chromosome pairing data. According to the genomic system of classification in Triticeae, H. patula should be considered as Elymus hystrix L., while H. duthiei and H. longearistata as Leymus duthiei and Leymus duthiei ssp. longearistata, respectively.     

Chromosome pairing relationships among the A,B, and D genomes of bread wheat

Summary Chromosome pairing and chiasma frequency were studied in bread wheat euhaploids (2n = 3x = 21; ABD genomes) with and without the major pairing regulatorPh1. This constitutes the first report of chromosome pairing relationships among the A, B, and D genomes of wheat without the influence of an alien genome. AllPh1 euhaploids had very little pairing, with 0.62–1.05 rod bivalents per cell; ring bivalents were virtually absent and mean arm-binding frequency (c) values ranged from 0.050 to 0.086. In contrast, theph1b euhaploids had extensive homoeologous pairing, with chiasma frequency 7.5–11.6 times higher than that in thePh1 euhaploids. They had 0.53–1.16 trivalents, 1.53–1.74 ring bivalents, and 2.90–3.57 rod bivalents, withc from 0.580 to 0.629. N-banding of meiotic chromosomes showed strongly preferential pairing between chromosomes of the A and D genomes; 80% of the pairing was between these genomes, especially in the presence of theph1b allele. The application of mathematical models to unmarked chromosomes also supported a 21 genomic structure of theph1b euhaploids. Numerical modeling suggested that about 80% of the metaphase I association was between the two most related genomes in the presence ofph1b, but that pairing under Ph1 was considerably more random. The data demonstrate that the A and D genomes are much more closely related to each other than either is to B. These results may have phylogenetic significance and hence breeding implications.This paper is dedicated to the memory of the late Ernest R. SearsCooperative investigations of the USDA-Agricultural Research Service and the Utah Agricultural Experiment Station, Logan, UT 84322, USA. Approved as Journal Paper No. 3986     

Synthesis and cytological characterization of trigeneric hybrids involving durum wheat,Thinopyrum bessarabicum,and Lophopyrum elongatum

Summary In an attempt to transfer genes for salt tolerance and other desirable traits from the diploid wheatgrasses, Thinopyrum bessarabicum (2n=2x=14; JJ genome) and Lophopyrum elongatum (2n=2x=14; EE genome), into durum wheat cv Langdon (2n=4x=28; AABB genomes), trigeneric hybrids with the genomic constitution ABJE were synthesized and cytologically characterized. C-banding analysis of somatic chromosomes of the A, B, J, and E genomes in the same cellular environment revealed distinct banding patterns; each of the 28 chromosomes could be identified. They differed in the total amount of constitutive heterochromatin. Total surface area and C-banded area of each chromosome were calculated. The B genome was the largest in size, followed by the J, A, and E genomes, and its chromosomes were also the most heavily banded. Only 25.8% of the total chromosome complement in 10 ABJE hybrids showed association, with mean arm-pairing frequency (c) values from 0.123 to 0.180 and chiasma frequencies from 3.36 to 5.02 per cell. The overall mean pairing was 0.004 ring IV + 0.046 chain IV + 0.236 III + 0.21 ring II + 2.95 rod II + 20.771. This is total pairing between chromosomes of different genomes, possibly between A and B, A and J, A and E, B and J, B and E, and J and E, in the presence of apparently functional pairing regulator Ph1. Because chromosome pairing in the presence of Ph1 seldom occurs between A and B, or between J and E, it was inferred that pairing between the wheat chromosomes and alien chromosomes occurred. The trigeneric hybrids with two genomes of wheat and one each of Thinopyrum and Lophopyrum should be useful in the production of cytogenetic stocks to facilitate the transfer of alien genes into wheat.     

Molecular cytogenetic evidence for a high level of chromosome pairing among different genomes in Triticum aestivum–Thinopyrum intermedium hybrids

Intergeneric hybrids (ABDJJ^sS genomes) were made between Triticum aestivum cv. Chinese Spring (CS) and Thinopyrum intermedium. Genomic in situ hybridization (GISH) using genomic DNA probes from Pseudoroegneria libanotica (Hackel) D.R. Dewey (genome S, 2n = 14) was used to study chromosome pairing among J, J^s, S and wheat ABD genomes in the hybrids. It was shown that in the hexaploid (ABDJJ^sS) hybrids, high pairing occurred among wheat chromosomes and among Thinopyrum chromosomes. A closer relationship was observed among the three genomes of Th. intermedium than among the three genomes of T. aestivum. It was further discerned that S genome chromosomes paired with J- and J^s-genome chromosomes at a high frequency. The frequency of heterologous pairing between S and J or S and J^s chromosomes was higher than those between J and J^s chromosomes, indicating that the S-genome was more closely related with these two genomes. Our results provided direct molecular cytogenetic evidence for the hypothesis that S-genome chromosomes are genetically similar to the J-genome chromosomes and, therefore, genetic exchange between these genomes is possible. The discovery of a close relationship among S, J and J^s genomes provides valuable markers for molecular cytogenetic analyses using S-genomic DNA probes in monitoring the transfer of useful traits from Thinopyrum species into wheat. Received: 23 August 2000 / Accepted: 5 September 2000     

Production and cytogenetics of hybrids of Triticum aestivum x Leymus innovatus

Summary Hybrid plants were obtained between Triticum aestivum (2n=6x=42, AABBDD) and Leymus innovatus (2n=4x=28, JJNN) at a frequency varying from 0.4% to 1.2% of the pollinated florets. Improvement of the embryo culture medium resulted in a higher frequency of embryo rescue. Eight of ten hybrids had the expected chromosome number of 35 (ABDJN). Meiotic analysis indicated that there was no homology between the genomes of the two species. Two hybrids had only 28 chromosomes. Comparison of chromosome pairing between the two types of hybrids suggested that Leymus innovatus carries genes that affect chromosome pairing and behavior. The relatively high occurrence of spontaneous doubling in the meiocytes of these hybrids may indicate that backcrossing of the hybrids to wheat should be possible, although frequent chromosome irregularities observed in the meiocytes of the hybrids may decrease the probability of success of this step, which is essential to the process of gene transfer from L. innovatus to wheat.Contrib. no. 366     

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.     

The effect of ph mutations on homoeolgous pairing in hybrids of wheat with Triticum longissimum

Homoeologous pairing at metaphase-I was analyzed in wild-type, ph2b, and ph1b hybrids of wheat and a low-pairing type of T. longissimum in order to study the effect of ph mutations on the pairing of T. longissimum chromosomes with wheat chromosomes. Chromosomes of both species, and their arms, were identified by C-banding. The three types of hybrids, with low-, intermediate-, and high-pairing levels, respectively, exhibited a very similar pairing pattern which was characterized by the existence of two types, A-D and B-S¹, of preferential pairing. These results confirm that the S¹ genome of T. longissimum is closely related to the B genome of wheat. The possible use of ph1b and ph2b mutations in the transfer to wheat of genes from related species is discussed.     

Intergeneric hybridization and C-banding patterns inHordelymus (Triticeae,Poaceae)

Crosses ofHordelymus europaeus (2n = 4x = 28) with four genera in theTriticeae were attempted. Adult hybrids were obtained in combinations withHordeum bogdanii (2x),Hordeum depressum (4x), andSecale cereale (2x). The meiotic pairing was very low in the hybrids withH. bogdanii andSecale cereale (0.12 and 0.30 chiasmata/cell, respectively), whereas high pairing (9.90 chiasmata/cell) was found in hybrids withH. depressum due to autosyndetic pairing ofH. depressum chromosomes. The chromosome complement ofHordelymus europaeus comprised 16 metacentrics, 8 submetacentrics, and 4 SAT-chromosomes. The Giemsa C-banding patterns of the chromosomes were characterized by small to minute bands at no preferential positions. It is hypothesized thatHordelymus europaeus may genomically be closest related toTaeniatherum andPsathyrostachys spp.     

Cytogenetics of a Hordeum vulgare x Elymus patagonicus hybrid (n=4x=28)

Summary Hordeum vulgare L. (2n=2x=14) was hybridized with Elymus patagonicus Speg. (2n=6x=42). The hybrid had 28 chromosomes, genomically represented as HSH₁H₂, and was perennial with a codominant phenotype. The chromosomes were meiotically associated as 19.6 univalents + 0.004 ring bivalents + 2.6 rod bivalents + 0.8 trivalents + 0.14 quadrivalents in 1,129 meiocytes, with a chiasma frequency of 4.77 per cell. The bivalent pairing presumably is an autosyndetic but modified expression of the H₁H₂ genomes of E. patagonicus, since ring bivalents were rare. This does not preclude the association of the H. vulgare H genome chromosomes with either H₁ and/or H₂ genomes of E. patagonicus to form bivalent or multivalent associations. A further evaluation of the genome homologies of H. vulgare, H. bogdanii, E. canadensis and E. patagonicus is proposed.     

Meiotic pairing behaviour reveals differences in genomic constitution between Hystrix patula and other species of the genus Hystrix Moench (Poaceae, Triticeae)

Interspecific and intergeneric hybridizations were carried out to evaluate the genomic relationships among species of Hystrix Moench and to study the relationships between Hystrix species and Psathyrostachys huashanica Keng (2n=2x=14, Ns^h). Meiotic pairing in hybrids of Hystrix duthiei ssp. duthiei × P. huashanica (2n=3x=21), Hystrix duthiei ssp. longearistata × P. huashanica (2n=3x=21) and H. patula × P. huashanica (2n=3x=21) averaged 5.18, 5.11 and 0.29 bivalents per cell, while H. patula × H. duthiei ssp. longearistata (2n=4x=28) averaged 25.36 univalents and 1.32 bivalents per cell, respectively. The results indicate that (i) H. duthiei ssp. duthiei and H. duthiei ssp. longearistata have one set of Ns genome from Psathyrostachys; (ii) H. patula has no Ns genome; (iii) genomes of H. duthiei ssp. duthiei and H. duthiei ssp. longearistata are non-homologous to those of H. patula. The genomic relationships between H. patula and other Hystrix species are also discussed.     

Diploidization and chromosomal pairing affinities in the tetraploid wheats and their putative amphiploid progenitor

Summary The genomes of the diploid wheats Triticum boeoticum and T. urartu are closely related, giving 7II in the f₁ hybrid (T^bT^u) and 8.4 (0–14) II + 2.5 (0–7) IV in the derived amphiploid (T^bT^bT^uT^u). The genomes of the tetraploid wheats are also closely related, giving up to 7II at the polyhaploid level (AB) in the absence of the gene Ph but 14II at the tetraploid level (AABB) in the normal presence of Ph. If the amphiploid is the progenitor of the tetraploids, one or the other homoeologue (T^b or T^u) in each of the 7 homoeologous groups (the 7 potential IV) must have differentiated with respect to pairing affinity in order to account for 14II in the tetraploid. Consequently, in tetraploid X amphiploid hybrids (T^bT^uAB) carrying the Ph gene from the tetraploid, the seven differentiated chromosomes (B) would be expected to give 7I while, on the basis of their observed chiasma frequency, T^b, T^u and the less differentiated A would be expected to give 4.17I + 3.57II + 3.23III), assuming homoeologous pairing. The expected chromosomal configuration freqencies at MI (11.17I + 3.57II + 3.23III) closely fit the observed values (11.22I + 3.45II + 3.19III + 0.071IV) for such hybrids (X² = 0.0046; P>0.99). Thus diploidization of the boeoticum-urartu amphiploid clearly could account for the origin of the tetraploid wheats. Furthermore, T. aestivum X amphiploid hybrids (T^bT^uABD) with and without Ph indicated that B as well as A chomosomes tended to pair with their presumed T^bT^u homologues in the absence of Ph. Other tests showed that the tetraploid wheats could not plausibly have originated from any postulated Triticum-Sitopsis (TTSS) parental combinations with or without such chromosomal differentiation.     

Cytogenetical studies on artificial hybrids amongElymus sibiricus,E. dahuricus andAgropyron tsukushiense in the tribe Triticeae,Gramineae

To explore the cytogenetical relationships ofElymus andAgropyron of the tribe Triticeae, Gramineae, two species of AsiaticElymus, E. sibiricus (2n=28) andE. dahuricus (2n=42), and a JapaneseAgropyron, A. tsukushiense (2n=42) were crossed. Pentaploid and hexaploid F₁ hybrids were vigorous. All pollen grains were aborted and none of the hybrids produced seed. For the crossE. sibiricus × A. tsukushiense, the average chromosome pairing per cell at the MI of the PMCs in the F₁ was 16.38 univalents, 8.93 bivalents, 0.25 trivalents and 0.01 quadrivalents; for the crossE. dahuricus × A. tsukushiense, it was 4.41 univalents, 17.67 bivalents, 0.32 trivalents, 0.28 quadrivalents and 0.04 quinquevalents; and for the crossE. dahuricus × E. sibiricus, it was 17.11 univalents, 8.74 bivalents, 0.04 trivalents and 0.07 quadrivalents. From the present results, it is concluded thatE. sibiricus contains one genome andE. dahuricus contains two genomes, which are homologous to those ofA. tsukushiense, and that the third genome ofE. dahuricus might be partially homologous to the remaining genome ofA. tsukushiense. This conclusion is also supported by the cytogenetical analysis ofE. dahuricus × E. sibiricus. Contribution No. 27 from the Plant Germ-plasm Institute, Faculty of Agriculture, Kyoto University, Kyoto, Japan.