Genome analysis ofElymus angulatus andE. patagonicus (Poaceae) and their hybrids with N. and S. AmericanHordeum spp.

The results of genome analysis of five hybrids, viz.Elymus patagonicus ×Hordeum procerum, E. patagonicus ×H. tetraploidum, E. angulatus ×H. jubatum, E. angulatus ×H. lechleri, andE. angulatus ×H. parodii, are reported. The genomic constitution ofHordeum tetraploidum andH. jubatum is best given as H₁H₁H₂H₂, ofH. lechleri andH. parodii as H₁H₁H₂H₂H₄H₄, ofH. procerum as H₁H₁H₂H₂H₃H₃, and ofElymus patagonicus andE. angulatus as SSH₁H₁H₂H₂.     

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.     

Genome relationships between Hordeum procerum (6x) and H. lechleri (6x)

Investigations on the meiotic behaviour of chromosomes in interspecific hybrids (2n=6x=42) between Hordeum lechleri (6x) and H. procerum (6x) and in their component haploids have been utilized to assess the nature of pairing and the extent of genome homology between the two species. In the F₁ hybrids an average of 25 (60%) chromosomes associated at metaphase I, mostly as bivalents. A majority (60%) of the pollen mother cells (PMCs) in H. procerum haploids (2n=3x=21) displayed 21 univalents and even in the remainder, a maximum of two rod bivalents were formed resulting in an average of 0.52 bivalents per cell. In haploids of H. lechleri (2n=3x=21) however, 30% of chromosomes pair. The sum of the chromosomal associations in the component haploids represents only 17% of the complement, far below the observed frequency (60%) in the hybrids. Thus, the pairing displayed in hybrids between H. lechleri and H. procerum was mostly allosyndetic and suggestive of two genomes being common in these species.In haploid H. procerum 1/3 of the PMCs displayed a tripolar organisation of chromosomes leading to triad and hexad formation after divisions I and II respectively. The significance of hexad formation in the trihaploid H. procerum and a possible suppression of homoeologous pairing in H. procerum haploids are discussed.     

Elimination and duplication of particular Hordeum vulgare chromosomes in aneuploid interspecific Hordeum hybrids

Summary Seeds formed in crosses Hordeum lechleri (6x) x H. vulgare (2x and 4x), H. arizonicum (6x) x H. v. (2x), H. parodii (6x) x H. v. (2x), and H. tetraploidum (4x) x H. v. (2x) produced plants at high or rather high frequencies through embryo rescue. Giemsa C-banding patterns were used to analyze chromosomal constitutions and chromosomal locations on the methaphase plate. Among 100 plants obtained from H. vulgare (2x) crosses, 32 plants were aneuploid with 2n=29 (1), 28 (3), 27 (13), 26 (5), 25 (4), 24 (4), or 22 (2); 50 were euploid (12 analyzed), and 18 were polyhaploid (5 analyzed). Four plants had two sectors differing in chromosome number. Two of four hybrids with H. vulgare (4x) were euploid and two were aneuploid. Parental genomes were concentrically arranged with that of H. vulgare always found closest to the metaphase centre. Many plants showed a certain level of intraplant variation in chromosome numbers. Except for one H. vulgare (4x) hybrids, this variation was restricted to peripherally located non-H. vulgare genomes. This may reflect a less firm attachment of the chromosomes from these genomes to the spindle. Interplant variation in chromosome numbers was due to the permanent elimination or, far less common, duplication of the centrally located H. vulgare chromosomes in all 34 aneuploids, and in a few also to loss/gain of non-H, vulgare chromosomes. This selective elimination of chromosomes of the centrally located genome contrasts conditions found in diploid interspecific hybrids, which eliminate the peripherally located genome. The difference is attributed to changed genomic ratios. Derivatives of various H. vulgare lines were differently distributed among euploid hybrids, aneuploids, and polyhaploids. Chromosomal constitutions of hypoploid hybrids revealed a preferential elimination of H. vulgare chromosomes 1, 5, 6, and 7, but did not support the idea that H. vulgare chromosomes should be lost in a specific order. H. vulgare SAT-chromosomes 6 and 7 showed nucleolar dominance. Aneuploidy is ascribed to the same chromosome elimination mechanism that produces haploids in cross-combinations with H. vulgare (2x). The findings have implications for the utilization of interspecific Hordeum hybrids.     

Dihaploids of Elymus from the interspecific crosses E. dolichatherus x E. tibeticus and E. brevipes x E. panormitanus

Summary Dihaploids (n=2x=14, SY) of two Elymus species, i.e., E. dolichatherus (Keng) Löve (2n=4x=28, SSYY) and E. brevipes (Keng) Löve (2n=4x=28, SSYY), were obtained from the interspecific hybrid combinations E. dolichatherus () x E. tibeticus (Meld.) G. Singh () and E. brevipes () x E. panormitanus (Parl.) Tzvelev (). The dihaploids were probably formed through selective elimination of male parental chromosomes in early embryo development. Meiotic chromosome behavior was studied in E. dolichatherus, E. brevipes, and their dihaploids. The two parental Elymus species had regular meioses with predominantly ring bivalent formation. A low frequency of homoeologous chromosome pairing was observed, with an average of 0.81 bivalents and 0.03 trivalents in the dihaploid of E. dolichatherus, and 0.26 bivalents in the dihaploid of E. brevipes. Up to two chromatid bridges accompanied by small fragments were present at anaphase I of the E. dolichatherus dihaploid. It is concluded from this study that: (i) both E. dolichatherus and E. brevipes are allotetraploid species; (ii) a low affinity exists between the S and Y genomes of the two Elymus species.     

Comparison of the Giemsa C-banded and N-banded karyotypes of twoElymus species,E. dentatus andE. glaucescens (Poaceae: Triticeae)

The karyotypes ofElymus dentatus from Kashmir andE. glaucescens from Tierra del Fuego, both carrying genomesS andH, were investigated by C- and N-banding. Both taxa had 2n = 4x = 28. The karyotype ofE. dentatus was symmetrical with large chromosomes. It had 18 metacentric, four submetacentric and six satellited chromosomes. The karyotype ofE. glaucescens resembled that ofE. dentatus, but a satellited chromosome pair was replaced by a morphologically similar, non-satellited pair. The C-banding patterns of both species had from one to five conspicuous and a few inconspicuous bands per chromosome. N-banding differentiated the chromosomes of the constituent genomes by producing bands in theH genome only. TheS genomes of both species were similar with five metacentric and two satellited chromosomes having most conspicuous C-bands at telomeric and distal positions. They resembled theS genome of the genusPseudoroegneria. TheH genomes had four similar metacentric and two submetacentric chromosomes. The seventhH genome chromosome ofE. dentatus was satellited, that ofE. glaucescens nonsatellited, but otherwise morphologically similar. The C-bands were distributed at no preferential positions. TheH genome ofE. dentatus resembles theH genomes of some diploidHordeum taxa.     

The cytogenetics of a Triticum turgidum x Psathyrostachys juncea hybrid and its backcross derivatives

Psathyrostachys juncea (2n = 2x = 14, NN), a source of barley yellow dwarf (BYDV) virus resistance with tolerance to drought and salinity, has been successfully hybridized in its autotetraploid form (2n = 4x = 28, NNNN) as the pollen parent to durum wheat (Triticum turgidum L.). The 2n = 4x = 28 (ABNN) F₁ hybrid has a mean meiotic metaphase-I configuration of 20.29 univalents + 0.29 ring bivalents + 3.36 rod bivalents + 0.14 trivalents. Spike length, internode length, glume awn length and lemma awn length, as well as the general spike morphology of the F₁ hybrid, are intermediate with those of the two parents. Pollinating the ABNN F₁ hybrid has given backcross (BC)-I derivatives of an amphiploid (AABBNN) that expresses limited self-fertility. BC-2 derivatives have been obtained from these plants. Direct transfers of useful genes from Ps. juncea to wheat would require substantial genetic manipulation strategies. Both conventional and novel methodologies, which may complement each other, and so facilitate reaching an agricultural objective end point, are addressed.     

Tissue-culture-facilitated production of aneupolyhaploid Thinopyrum ponticum and amphidiploid Hordeum violaceum x H. bogdanii and their uses in phylogenetic studies

Summary An aneupolyhaploid (2n = 36) of the decaploid Thinopyrum ponticum and an amphidiploid (2n = 28) of Hordeum violaceum x Hordeum bogdanii were produced through anther and inflorescence culture, respectively. Meiotic associations in pollen mother cells at metaphase I of these plants were analyzed. The aneupolyhaploid arose by direct embryogenesis from a microspore without passing through a callus phase. The mean pairing frequencies of 2.67 univalents ⁺ 0.54 rod bivalents ⁺ 8.85 ring bivalents ⁺ 2.75 trivalents ⁺ 0.17 chain quadrivalents ⁺ 0.56 ring quadrivalents ⁺ 0.65 pentavalents in the aneupolyhaploid (2n = 36) best fit the 221 model. However, the frequent multivalents (up to five trivalents, or three quadrivalents, or four pentavalents in a cell) indicated that decaploid T. ponticum has five sets of closely related genomes representable by the genome formula J₁ J₁ J₁ J₂ J₂. Colchicine treatment of inflorescence-derived H. violaceum x H. bogdanii regenerants greatly enhanced the rate of chromosome doubling, and completely doubled regenerants could be isolated. The H. violaceum x H. bogdanii amphidiploid had a mean pairing pattern of 12.53 univalents ⁺ 5.57 rod bivalents ⁺ 1.97 ring bivalents ⁺ 0.07 trivalents ⁺ 0.03 hexavalents, indicating the presence of desynaptic gene(s) in the original diploiid hybrid. Therefore, the pairing frequency in that diploid hybrid was an under-estimate of chromosome homology between the parental genomes, and additional diploid hybrids are needed to assess the genome homology between H. violaceum and H. bogdanii. These two contrasting experiments demonstrated that tissue culture techniques are useful in altering the ploidy level to produce plant materials suitable for genome analysis and phylogenetic studies.Cooperative investigation of the USDA-ARS, Forage and Range Research Laboratory, Logan, UT 84322-6300, and the Utah Agricultural Experiment Station, Utah State University, Logan, UT 84322-4810. Approved as journal paper No. 3991     

A novel intergeneric hybrid in the Triticeae: Triticum aestivum x Psathyrost achys juncea

Summary Two hybrid embryos of intergeneric origin between Triticum aestivum cv Fukuho (2n=6x=42, AABBDD) and Psathyrostachys juncea (2n=2x=14, NN) were successfully rescued. One hybrid plant had the expected chromosome number of 28 (ABDN), whereas the second plant had 35 chromosomes. The average meiotic chromosome pairing in the 35-chromosome hybrid was 21.87 univalents + 6.38 bivalents + 0.11 trivalents + 0.009 quadrivalents, which indicates that two copies of the N genome were present. Chromosome pairing in the 28-chromosome hybrid was low (1.35 bivalents), and pointed out the lack of homology between the wheat genomes and the P. juncea genome. These new hybrids showed some necrosis and chlorosis, which caused severe floral abortion in the plant that had 35 chromosomes. These problems became gradually less severe after 18 months.Contrib. no. 372     

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     

Characterisation of progeny from backcrosses of triploid hybrids between Hordeum vulgare L. (2x) and H. bulbosum L (4x) to H. vulgare

Interspecific hybridisations between Hordeum vulgare L. (cultivated barley) and H. bulbosum L. (bulbous barley grass) have been carried out to transfer desirable traits, such as disease resistance, from the wild species into barley. In this paper we report the results of an extensive backcrossing programme of triploid hybrids (H. vulgare 2x x H. bulbosum 4x) to two cultivars of H. vulgare. Progenies were characterised cytologically and by restriction fragment length polymorphism analysis and comprised (1) haploid and diploid H. vulgare plants, (2) hybrids and aneuploids, (3) single and double monosomic substitutions of H. bulbosum chromosomes into H. vulgare and (4) chromosomal rearrangements and recombinants. Five out of the seven possible single monosomic chromosome substitutions have now been identified amongst backcross progeny and will be valuable for directed gene introgression and genome homoeology studies. The presence amongst progeny of 1 plant with an H. vulgare-H. bulbosum translocated chromosome and one recombinant indicates the value of fertile triploid hybrids for interspecific gene introgression.     

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.     

Monosomic and double monosomic substitutions of Hordeum bulbosum L. chromosomes into H. vulgare L.

Summary One of the aims of the interspecific crossing programme between Hordeum vulgare L. and H. bulbosum L. has been to introgress desirable genes into barley from the wild species. However, despite their close taxonomic relationship there have been few reports of achieving this objective using amphidiploid hybrids. In order to broaden the range of available hybrids, partially fertile triploids between H. vulgare (2n = 2x = 14) and H. bulbosum (2n = 4x = 28) were developed and crossed with H. vulgare as female parent. From 580 progeny which were screened, eight putative single monosomic chromosome substitution lines and one double monosomic substitution were identified by cytological analysis. These involved the substitution of H. vulgare chromosome 1 (two lines), 6 (four lines), 6L (one line), 7 (one line) and 1 + 4 (one line) by their respective H. bulbosum homoeologues. The H. bulbosum chromosome was frequently eliminated during plant development, but it was observed regularly in pollen mother cells of two lines. However, pairing between the H. bulbosum chromosome and its H. vulgare homoeologue was low. Several of the lines were more resistant than their H. vulgare parents to powdery mildew (Erysiphe graminis DC. f.sp. hordei Em. Marchai), net blotch (Drechslera teres Sacc.) and scald (Rhynchosporium secalis (Oudem.) Davis). Apart from their use in studying genome relationships, their value to plant breeders will depend on the ease of inducing translocations between the parental chromosomes.     

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.     

Genome relationships between Hordeum vulgare L. and H. bulbosum L.

Interrelationships between H. vulgare (2x=14) and H. bulbosum (2x=14; 4x=28) were estimated on the basis of the karyotypes and the pairing behaviour of the chromosomes in diploid, triploid and tetraploid hybrids obtained with the aid of embryo culture. — A comparison of the karyotypes of the two species revealed similarities as well as differences. It was concluded that at least 4 or more of the chromosomes were similar in morphology and probably closely related. — Diploid and tetraploid hybrids are rarely obtained and their chromosome numbers tend to be unstable whereas triploid hybrids (1 vulgare + 2 bulbosum genomes) were stable and relatively easy to produce. In the diploid hybrid only 40% of the meiotic cells contained 14 chromosomes while the numbers ranged from 7 to 16 in other cells. All hybrids exhibited pairing between the chromosomes of the two species. Diploid hybrids had a mean of 5.0 and a maximum of 7 bivalents per cell in those cells having 14 chromosomes. Triploid hybrids from crosses between 2x H. vulgare and 4x H. bulbosum exhibited a mean of 1.5 and a maximum of 5 trivalents per cell. In a hexaploid sector found following colchicine treatment of a triploid the mean frequencies of chromosome associations per cell were: 5.5_I+8.0_II+0.7_III+3.7_IV+0.3_V+0.4_VI. One unstable 27 chromosome hybrid obtained from crosses between the autotetraploid forms had a mean of 1.1 and a maximum of 4 quadrivalents per cell. The chromosome associations observed in these hybrids are consistent and are taken as evidence of homoeologous pairing between the chromosomes of the two species. Interspecific hybridization between these two species also reveals that chromosome stable hybrids are only obtained when the genomes are present in a ratio of 1 vulgare2 bulbosum. Based upon the results obtained, the possibility of transferring genetic characters from H. bulbosum into cultivated barley is discussed.     

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.     

C-banding variation in the Moroccan oat species Avena agadiriana (2n=4x=28)

The C-banding technique was used to describe the chromosomes of a relatively recently-discovered Moroccan oat species, Avena agadiriana (2n=4x=28). A substantial amount of polymorphism for arm ratios and C-banding patterns was observed among five accessions of this species. However a common set of ten putatively homologous chromosomes was identifiable among the five accessions. The chromosomes of A. Agadiriana do not closely match those of any of the previously described diploid or tetraploid oat species in terms of their arm ratios and C-banding patterns. However, their overall C-banded appearance generally resembles the A/B/D groups of chromosomes of Avena species, rather than the more hetrochromatic C genomes. Implications of these findings in terms of chromosome evolution in the genus Avena are discussed.Contribution no. 95-490-J of the Kansas Agricultural Experiment Station, Kansas State University, Manhattan, KS, USA     

Molecular cytogenetic study on the plants of Elymus nutans with varying fertility on the Qinghai-Tibet Plateau

A molecular cytogenetic investigation was conducted on plants of the allohexaploid species Elymus nutans with varying fertility on the Qinghai-Tibet Plateau. Molecular karyotyping revealed that chromosome variants were distributed unevenly among genomes and among different homologue chromosomes in each genome. The plants with varying fertility exhibited significantly higher numbers of chromosome variants than did the normal fertility samples, although both kinds of plants showed the same pattern of high-to-low polymorphism from the Y to St and H genomes. Heterozygosis and karyotype heterozygosity in the plants with varying fertility were 3- and 13-fold higher than those in normal samples, respectively. Significant negative correlations were found not only between seed setting rates and total genome heterozygosity but also between seed setting rates and heterozygosity of each genome in the plants of varying fertility. Chromosome pairing analysis was performed using genomic in situ hybridization in selected plants of different fertility levels. The pairing of chromosomes at meiotic metaphase I was mostly bivalent, although univalent, trivalent, quadrivalent, and other polyvalents also occurred; in addition, chromosome configuration forms and frequencies varied among the studied samples. ANOVA results showed that the average number of ring bivalents in the Y genome was significantly higher than those in the St and H genomes. Significant positive correlations between pollen grain fertility and ring bivalent number were found in the St and H genomes but not in the Y genome. Furthermore, chromosome configuration parameters (total bivalents, numbers of ring and rod bivalents) were found to be significantly correlated with heterozygosity and seed setting rates in the St and H genomes, respectively, but not in the Y genome. It was inferred that the seed setting rate and pollen grain fertility in E. nutans are strongly influenced by the heterozygosity of each genome, but the Y genome differs from the St and H genomes due to chromosome pair alterations. The St and H genomes may contain more chromosome structural variations than the Y genome in E. nutans.     

Cytogenetic and genomic relationships ofThinopyrum elongatum with twoPsathyrostachys species and withLeymus secalinus (Poaceae)

Intergeneric hybridizations were made betweenT. elongatum, and twoPsathyrostachys and fiveLeymus species. The seed set obtained onT. elongatum ×Leymus hybrids ranged from 5.65% to 20.00%, depending onLeymus species. The seed set obtained onT. elongatum ×Psathyrostachys hybrids ranged from 16.07% to 19.70%. Meiotic pairing at metaphase-I in JN diploid hybrids ofT. elongatum ×Psathyrostachys species revealed a very low level homology between the basic J and N genomes, and further demonstrated that the two genomes are quite diverged. Chromosome pairing in theT. elongatum ×Leymus secalinus hybrid averaged 15.19 univalents + 2.62 rod bivalents + 0.26 ring bivalents + 0.02 trivalents, suggesting that the partial J^e chromosomes ofT. elongatum has homology withLeymus secalinus genomes.L. secalinus might have 3–4 chromosomes originating from J^e genome.     

Cytogenetic studies of the intergeneric hybrids between Secale cereale and Elymus caninus,E. brevipes,and E. tsukushiensis (Triticeae: Poaceae)

Summary Integeneric hybridizations were carried out between Secale cereale L. (2n = 14, RR) and three Elymus species, namely, E. caninus (L.) L. (2n = 28, SSHH), E. brevipes (Keng) Löve (2n = 28, SSYY) and E. tsukushiensis Honda (2n = 42, SSHHYY). Chromosome pairing was studied at metaphase I in the parental species and the hybrids. Meiotic configurations of the hybrids were 20.74 1+0.14 II for E. caninus x S. cereale (SHR), 16.35 I+2.17 II+0.09 III for E. brevipes x S. cereale (SYR) and 25.84 I+1.10 II+0.02 III for E. tsukushiensis x S. cereale (SHYR), in addition to some secondary associations in the different hybrids. It is concluded from the study that (1) a certain, different homoeologous relationship exists among S, H and Y genomes in the investigated Elymus species; (2) low homoeology is present between genomes of Elymus (S or H or Y) and rye (R); (3) the Secale genome affects homoeologous chromosome pairing between different genomes in E. brevipes and E. tsukushiensis.