Meiotic studies ofElymus nutans andE. jacquemontii (Poaceae,Triticeae) and their hybrids withPseudoroegneria spicata and seventeenElymus species

Hybridizations ofElymus nutans andE. jacquemontii were carried out with one species ofPseudoroegneria (S genome), and 20Elymus species, each containing either of the SH, SY, SYH, or SYW genomes. Chromosome configurations were analysed at metaphase I of the two target taxa and their interspecific hybrids. It is concluded that (i)E. nutans is an allohexaploid containing the SYH genomes, andE. jacquemontii is an allotetraploid having the SY genomes; (ii) the genomic affinity is associated with the geographic distance between the species studied; (iii) minor genomic structural rearrangements have occurred within the hexaploid taxon ofE. nutans.     

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.     

Genomic relationships of theElymus parviglumis,E. semicostatus,andE. tibeticus groups (Triticeae: Poaceae)

In order to investigate genomic relationships of tetraploidElymus species (all containing the SY genomes) among three morphological groups, i.e. theE. parviglumis, E. semicostatus, andE. tibeticus groups, interspecific hybridizations were carried out among representative species from the three groups. Chromosome associations at metaphase I were analysed in the interspecific hybrids, and genomic relationships of the species within and among the three groups were estimated. Hybridizations among species of the three groups were fairly easy to perform, but no general pattern for crossabilities among certain species or groups was obtained. All the hybrids were completely sterile. Meiotic pairing was moderately high, but ranges of chiasmata were observed in different hybrid combinations. There is no tendency for genomic affinities to be higher within groups than between the groups. Therefore, the meiotic data do not support the division of the SY-genome species into the three groups. However, there is a clear tendency that the differentiation of the SY genomes in the tetraploids is associated with the geographic distribution of the species.     

Production and cytogenetic analysis of intergeneric hybrids betweenElymus anthosachnoides andPsathyrostachys huashanica (Poaceae: Triticeae)

Intergeneric hybridization betweenElymus anthosachnoides (2n = 28,SSYY) andPsathyrostachys huashanica (2n = 14,NN) was performed. Three hybrid plants, obtained via embryo rescue, were intermediate between the parents in morphology and developed vigorously, but were completely sterile. The mean chromosome configuration was 19.48 I + 0.76 II per cell in the hybrids at meiotic metaphase I. This result indicates thatE. anthosachnoides andP. huashanica are distantly related and that there is little or no homoeology betweenN (P. huashanica) andS orY (E. anthosachnoides) genomes.     

Genomic relationships within theElymus parviglumis group (Triticeae: Poaceae)

Genomic relationships of 13 tetraploid species within the AsiaticElymus parviglumis group containing the SY genomes were assessed by analysing chromosome pairing at metaphase I of the parental species and their interspecific hybrids. Two major genomic subgroups among the tetraploids were identified from the cluster analysis of the averaged c-values, namely, theE. caucasicus subgroup (two species) and theE. parviglumis subgroup (11 species). The genomic affinity of theElymus species is associated with the interspecific geographic distance.After October 1, 1993     

Interspecific hybridizations among species of theElymus semicostatus andElymus tibeticus groups (Poaceae)

Interspecific hybridizations were made between species of theE. semicostatus group, viz.,E. semicostatus (Nees exSteud.)Meld.,E. validus (Meld.)B. Salomon,E. abolinii (Drob.)Tzvel., andE. fedtschenkoi Tzvel., and species of theE. tibeticus group, viz.,E. pendulinus (Nevski)Tzvel.,E. tibeticus (Meld.)Singh,E. shandongensis B. Salomon, andE. gmelinii (Ledeb.)Tzvel., as well as among species within theE. tibeticus group. All species are tetraploid (2n = 4x = 28) and possess SY genomes. Meiotic pairing data from 24 hybrids involving 17 interspecific combinations are presented. The average number of chiasmata per cell ranged from 17.91 to 26.20 in hybrids within theE. tibeticus group, compared with 7.26 to 22.04 in hybrids between the two species groups. Despite the extensive collection of cytological data, there was no definite evidence for confirming or disproving the separate existence of the two groups.     

Biosystematic study ofRoegneria tenuispica, R. ciliaris andR. pendulina (Poaceae:Triticeae)

Morphological and cytological studies of three tetraploidRoegneria species,R. tenuispica, R. pendulina andR. ciliaris, and their artificial hybrids were carried out.Roegneria tenuispica was morphologically similar toR. pendulina. The general appearance of the interspecific hybrids was intermediate between the parents. The hybrids showed comparatively high chromosome pairing at meiosis, but were completely or almost completely sterile. The results indicate that the three independent species share two basic genomes (S_tY) and thatR. tenuispica is more closely related toR. pendulina than toR. ciliaris. The genomes ofR. tenuispica could be designated as S _t ^t Y^t.     

The genome constitution ofRoegneria grandis (Poaceae, Triticeae)

Roegneria grandis was hybridized withR. ciliaris var.japonensis (2n = 28, SSYY),Elymus caninus (2n = 28, SSHH), andPseudoroegneria spicata (2n = 28, SSSS). Chromosome pairing was studied in parents and hybrids. It is concluded from this study that: (i)R. grandis is an allotetraploid species and contains the basic genomes S and Y: (ii) a certain degree of homoeology exists between the S and Y genomes of the species studied.     

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.     

Genomic relationships between species of theElymus semicostatus group andElymus sensu lato (Poaceae)

Meiotic pairing behaviour in 19 interspecificElymus hybrids is reported and discussed. The hybrids were made between four species belonging to theE. semicostatus group of sect.Goulardia, viz.,E. semicostatus, E. abolinii, E. fedtschenkoi, andE. panormitanus (all 2n = 28), andElymus species of seven different sections, viz., sect.Clinelymiopsis:E. caucasicus (2n = 28); sect.Elymus:E. sibiricus (2n = 28); sect.Goulardia:E. caninus (2n = 28),E. trachycaulus (2n = 28), andE. tsukushiensis (2n = 42); sect.Hyalolepis:E. batalinii (2n = 42); sect.Hystrix:E. hystrix (2n = 28); sect.Macrolepis:E. canadensis (2n = 28); and sect.Turczaninovia:E. dahuricus (2n = 42). Chromosomal pairing at meiotic metaphase I indicated that the species of theE. semicostatus group are genomically closer to the tetraploidE. caucasicus and the hexaploid species, regardless of sectional origin, than to the other tetraploid species of sectionGoulardia. Highest meiotic pairing was found in hybrids involvingE. caucasicus, E. tsukushiensis, andE. dahuricus. The presence of pairing regulating genes inE. abolinii is suspected.     

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.     

Genomic groups,morphology, and sectional delimitation in EurasianElymus (Poaceae,Triticeae)

Seven tetraploid species ofElymus, viz.E. sibiricus, E. caninus, E. gmelinii, E. semicostatus, E. caucasicus, E. parviglume, andE. longearistatus subsp.canaliculatus, representing five sections were studied morphologically and used in interspecific hybridizations. The aim was to investigate whether the present sectional delimitation of the genus was in agreement with genomic data and if there was a correlation between genome constitution and morphology. The study revealed: (i) further information on the genomic affinities between the different species, (ii) that there is no congruence between genome constitution of the species and current sectional delimitation, and (iii) that there is a correlation between genome constitution and morphology in the palea apex shape and in the size of cilia of the palea.     

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.     

Cytogenetic analyses inKengyilia laxiflora (Poaceae, Triticeae)

Kengyilia laxiflora (2n = 42) was cytogenetically studied with testersK. hirsuta (2n = 42, PPStStYY) andRoegneria kamoji (2n = 42, HHStStYY). Our data suggested thatK. laxiflora may possess a modified P genome, the P¹. Its St and Y genomes may also contain some structural changes and are more closely related to the St and Y genomes ofK. hirsuta than to those ofR. kamoji. Kengyilia laxiflora is reproductively separated from bothK. hirsuta andR. kamoji. These results indicated thatK. laxiflora is a good species in the genusKengyilia.     

Kengyilia habahenensis (Poaceae: Triticeae) — a new species from the Altai mountains,China

Kengyilia habahenensis, spec. nova, from the Altai mountains, China, is described morphologically and cytologically. It has 2n = 42 chromosomes, and the genome formula PYS.     

Meiotic analysis of the interspecific and intergeneric hybrids between Hystrix patula Moench and H. duthiei ssp. longearistata, Pseudoroegneria, Elymus, Roegneria, and Psathyrostachys species (Poaceae, Triticeae)

Interspecific and intergeneric hybridizations were carried out in an investigation of genome homology between Hystrix patula and other species of Hystrix , as well as the generic relationships between H. patula and its related species. Meiotic pairing in the hybrids H. patula  ×  H. duthiei ssp. longearistata (Ns–), H. patula  ×  Pseudoroegneria spicata (St), H. patula  ×  Pse. libanotica (St), Elymus sibiricus (StH) ×  H. patula , H. patula  ×  E. wawawaiensis (StH), Roegneria ciliaris (StY) ×  H. patula , H. patula  ×  R. grandis (StY), and H. patula  ×  Psathyrostachys huashanica (Ns^h) averaged 1.32, 6.53, 5.62, 10.08, 12.83, 3.57, 3.98, and 0.29 bivalents per cell, respectively. The results indicate that: (1) H. patula has no genome homology with H. duthiei ssp. longearistata or the Ns genome from Psathyrostachys ; (2) H. patula contains the same StH genomes as the Elymus species, and the St genome is homologous to the genome of Pse. spicata and Pse. libanotica ; and (3) H. patula has a low genome affinity with the StY genomes of Roegneria . Therefore, it is reasonable to treat H. patula Moench as E. hystrix L.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 213–219.     

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.     

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.     

Genome constitution of Elymus tangutorum (Poaceae: Triticeae) inferred from meiotic pairing behavior and genomic in situ hybridization

Elymus tangutorum (Nevski) Hand.-Mazz (2n = 6x = 42) is a perennial species in the tribe Triticeae, which distributes in Nepal and north and northwest China. However, the genome constitution of E. tangutorum is controversial and its taxonomic status is not clear. Hybridizations of E. tangutorum were carried out with E. wawawaiensis J. R. Carlson & Barkworth (StH), Roegneria grandis Keng (StY), and E. dahuricus Turcz. ex Griseb. (StYH). Meiotic pairing in the hybrids E. tangutorum × E. wawawaiensis (StH), E. tangutorum × R. grandis (StY), and E. tangutorum × E. dahuricus (StYH) averaged 10.48, 11.12, and 20.92 bivalents per cell, respectively. The results suggested that E. tangutorum is an allohexaploid and contains the StYH genomes. Results of genomic in situ hybridization analysis strongly supported the chromosome pairing data. Therefore, E. tangutorum should be treated as Campeiostachys dahurica var. tangutorum (Nevski) B. R. Baum, J. L. Yang & C. Yen. Intergenomic rearrangements of E. tangutorum may be affected by environmental factors.     

Hystrix patula与Pseudoroegneria libanotica属间杂种的细胞学研究

为研究猬草Hystrixpatula的染色体组组成,进行了H.patula与Pseudoroegnerialibanotica的人工杂交,获得杂种F1,观察了亲本和杂种F1花粉母细胞减数分裂染色体配对行为。杂种F1染色体配对较高,84%的细胞形成7个或7个以上二价体,其构型为6.08Ⅰ+7.48Ⅱ,C-值为0.69。结果表明,H.patula含有St染色体组。