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 constitution and taxonomy of the Chinese hexaploids Elymus cylindricus and E. breviaristatus (Poaceae: Triticeae)

Elymus cylindricus (2n = 6x = 42) and E. breviaristatus (2n = 6x = 42) are distributed in grasslands and deserts of northern and north‐western China. Their genomic constitution and taxonomic status are unclear. Elymus cylindricus was crossed with E. wawawaiensis J.R.Carlson & Barkworth ( StH ), Roegneria grandis Keng ( StY ) and Campeiostachys dahurica (Turcz. ex Griseb.) B.R.Baum, J.L. Y ang & C. Y en var. dahurica ( StYH ). Meiotic pairing in the hybrids E. cylindricus × E. wawawaiensis ( StH ), E. cylindricus × R. grandis ( StY ) and E. cylindricus × C. dahurica var. dahurica ( StYH ) showed on average 10.00, 11.30 and 20.92 bivalents per cell, respectively. Elymus breviaristatus was crossed with C. dahurica var. dahurica ( StYH ) and E. cylindricus. Chromosome pairing in the hybrids of E. breviaristatus × C. dahurica var. dahurica and E. breviaristatus × E. cylindricus showed on average 19.60 and 19.27 bivalents, respectively. Genomic in situ hybridization (GI SH ) revealed the presence of St , Y and H genomes in E. cylindricus and E. breviaristatus. An intergenomic rearrangement was observed in E. cylindricus using GI SH . Meiotic pairing data and GI SH indicated that both E. cylindricus and E. breviaristatus are allohexaploids containing the StYH genomes. Elymus cylindricus and E. breviaristatus should be treated as Campeiostachys dahurica var. cylindrica and Campeiostachys breviaristata, respectively.     

10个四倍体披碱草属物种的核型（简报）

本文报道了来自北美和亚洲10个四倍体披碱草属物种Elymus(StStHH)的核型.其核型属于1A或2A型,核型公式如下:24m+4sm(2sat)(加拿大披碱草E. canadensis);20m+8sm(4sat)(E. elymoides);20m+8sm(4sat)(E. glaucus);22m+6sm(4sat)(E.lanceolatus);22m+6sm(2sat)(E. mutabilis);22m+6sm(2sat)(老芒麦.E.sibiricus);24m+4sm(E.trachycaulus);22m+4sm+2st(2sat)(E.trachycaulus ssp.subsecundus);24m+4sm(4sat)(E.virginicus);20m+8sm(4sat)(E.wawawaiensis).     

Genome discrimination by in situ hybridization in Icelandic species of Elymus and Elytrigia (Poaceae: Triticeae).

The genome constitution of Icelandic Elymus caninus, E. alaskanus, and Elytrigia repens was examined by fluorescence in situ hybridization using genomic DNA and selected cloned sequences as probes. Genomic in situ hybridization (GISH) of Hordeum brachyantherum ssp. californicum (diploid, H genome) probe confirmed the presence of an H genome in the two tetraploid Elymus species and identified its presence in the hexaploid Elytrigia repens. The H chromosomes were painted uniformly except for some chromosomes of Elytrigia repens which showed extended unlabelled pericentromeric and subterminal regions. A mixture of genomic DNA from H. marinum ssp. marinum (diploid, Xa genome) and H. murinum ssp. leporinum (tetraploid, Xu genome) did not hybridize to chromosomes of the Elymus species or Elytrigia repens, confirming that these genomes were different from the H genome. The St genomic probe from Pseudoroegneria spicata (diploid) did not discriminate between the genomes of the Elymus species, whereas it produced dispersed and spotty hybridization signals most likely on the two St genomes of Elytrigia repens. Chromosomes of the two genera Elymus and Elytrigia showed different patterns of hybridization with clones pTa71 and pAes41, while clones pTa1 and pSc119.2 hybridized only to Elytrigia chromosomes. Based on FISH with these genomic and cloned probes, the two Elymus species are genomically similar, but they are evidently different from Elytrigia repens. Therefore the genomes of Icelandic Elymus caninus and E. alaskanus remain as StH, whereas the genomes of Elytrigia repens are proposed as XXH.     

Cytogenetics of Elymus magellanicus and its intra- and inter-generic hybrids with Pseudoroegneria spicata, Hordeum violaceum, E. trachycaulus, E. lanceolatus, and E. glaucus (Poaceae: Triticeae).

This paper describes the cytogenetics of the South American grass Elymus magellanicus (2n = 4x = 28) and its inter- and intra-generic hybrids with North American Pseudoroegneria spicata (2n = 2x = 14; SS), Iranian Hordeum violaceum (2n = 2x = 14; HH), and North American E. trachycaulus (2n = 4x = 28; SSHH), E. lanceolatus (2n = 4x = 28; SSHH), and E. glaucus (2n = 4x = 28; SSHH). Chromosome pairing in the F1 hybrid of E. magellanicus x P. spicata (2n = 3x = 21) averaged 4.09 bivalents per cell. Bivalent frequencies in the E. magellanicus x H. violaceum (2n = 3x = 21) averaged 4.86 bivalents per cell. Bivalent formation (frequencies) in hybrids between North and South American tetraploids averaged 11.42 in E. magellanicus x E. trachycaulus, 13.02 in E. magellanicus x E. lanceolatus, and 10.91 in E. magellanicus x E. glaucus. Meiotic data demonstrated that E. magellanicus is an allotetraploid having the same basic genomes (SSHH) as North American E. trachycaulus, E. lanceolatus, and E. glaucus.     

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.     

利用RAPD分析披碱草属、鹅观草属和猬草属模式种的亲缘关系

利用随机扩增多态性DNA（RAPD）技术对小麦族披碱草属、鹅观草属和猬草属3个属的模式种进行了基因组DNA多态性分析。42个引物产物的290条谱带中,257条（88.62%)表现出多态性,说明披碱草属、鹅观草属和猬草属3个属的模式种间具有丰富的遗传多样性。利用290个RAPD标记,计算材料间Nei氏遗传相似性系和遗传距离,在NTSYS程序中利用UPGMA进行聚类。结果表明,Elymus sibiricus种不同居群间的遗传差异较小,遗传距离在0.097-0.180之间。E.sibiricus,Roegneria caucasica和Hystrix patula的种间遗传差异明显,遗传距离在0.458-0.605之间。H.patula与E.sibiricus的亲缘关系较近。R.caucasica与E.sibiricus的亲缘关系较远。     

Meiotic configuration and chromosome number in some Iranian species of Elymus L. and Agropyron Gaertner (Poaceae: Triticeae)

Chromosome numbers and analyses of meiotic metaphase I are reported for the following taxa: Agropyron cristatum subsp. incanum (2n= 42), A. cristatum subsp. pecttnatum (2n=28 – 33), Elymus elongatus subsp. ponticus (2n= 69, 70), E. hispidus var. hispidus (2n= 41 43), var. podperae (2n= 42) and var. villosus (2n= 41, 42), E. libanoticus (2n= 14), E. pertenuis (2n= 28, 28+1B), E. repens (2n= 42), E. transhyrcanus (2n= 40–42), E. hispidus var. villosus x E. cf. repens (2n= 42). Chromosome numbers only are reported for the following taxa: E. gentri (2n= 41, 42), E. nodosus subsp. dorudicus (2n= 28), and E. elongatiformis (2n= 56, 57). The haploid genomic constitution SP is reported for Elymus pertenuis. Variable chromosome numbers (2n= 28–32) were observed in the meiotic metaphase I within single anthers of Agropyron cristatum subsp. pectinatum, and the supernumerary chromosomes in this taxon are assumed to have originated from crosses with hexaploids. Partial elimination of these supernumerary chromosomes probably occurs during archesporial mitotic divisions or at an early stage in the meiotic cycle. A hybrid, morphologically intermediate between E. hispidus and E. repens, was obtained from a seed of E. hispidus collected in the field. The meiotic metaphase I configuration in this E. hispidus hybrid suggests that the pollen parent may itself be a hybrid or hybrid derivative of E. repens x E. hispidus.     

Origin and Reticulate Evolutionary Process of Wheatgrass Elymus trachycaulus (Triticeae: Poaceae)

To study origin and evolutionary dynamics of tetraploid Elymus trachycaulus that has been cytologically defined as containing StH genomes, thirteen accessions of E. trachycaulus were analyzed using two low-copy nuclear gene Pepc (phosphoenolpyruvate carboxylase) and Rpb2 (the second largest subunit of RNA polymerase II), and one chloroplast region trnL–trnF (spacer between the tRNA Leu (UAA) gene and the tRNA-Phe (GAA) gene). Our chloroplast data indicated that Pseudoroegneria (St genome) was the maternal donor of E. trachycaulus. Rpb2 data indicated that the St genome in E. trachycaulus was originated from either P. strigosa, P. stipifolia, P. spicata or P. geniculate. The Hordeum (H genome)-like sequences of E. trachycaulus are polyphyletic in the Pepc tree, suggesting that the H genome in E. trachycaulus was contributed by multiple sources, whether due to multiple origins or introgression resulting from subsequent hybridization. Failure to recovering St copy of Pepc sequence in most accessions of E. trachycaulus might be caused by genome convergent evolution in allopolyploids. Multiple copies of H-like Pepc sequence from each accession with relative large deletions and insertions might be caused by either instability of Pepc sequence in H- genome or incomplete concerted evolution. Our results highlighted complex evolutionary history of E. trachycaulus.     

Characterization and analysis of microsatellite loci in Elymus caninus (Triticeae: Poaceae)

 Microsatellites are highly variable DNA sequences that can be used as markers for the genetic analysis of plants. The potential of microsatellite markers for use in a genetic diversity study in Elymus species was evaluated. Genomic libraries of Elymus caninus were constructed. The libraries were screened with two dinucleotide, (GA)n and (GT)n, and two trinucleotide repeats, (TCT)n and (CAC)n. A total of 19 positive clones were found for the two dinucleotide repeats; no positive clone was found for the trinucleotide repeats. Positive clones were sequenced to confirm the presence of microsatellites and to generate polymerase chain reaction (PCR) primers based on the sequences flanking the microsatellite. All sequenced (GA)n clones have repeats of n>10; over half of the (GT)n microsatellites have n<10 repeats. Primer pairs were designed and evaluated for 8 selected microsatellites. PCR products were amplified from 15 Elymus caninus accessions. The number of alleles found for the eight loci varied from 1 for ECGA89 and ECGT35 to 13 for ECGA22, as determined by non-denaturing polyacrylamide electrophoresis. Six microsatellite loci were found to be polymorphic in E. caninus. The eight primer pairs were tested on three other species; seven were successful in amplifying DNA from Elymus alaskanus and E. mutabilis, and four amplified DNA from E. caucasicus. Based on these results, microsatellites appear to be useful markers in detecting variation in E. caninus. Received: 8 September 1997/Accepted: 6 October 1997     

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     

Molecular diversity and relationships among Elymus trachycaulus, E. subsecundus, E. virescens, E. violaceus, and E. hyperarcticus (Poaceae: Triticeae) as determined by amplified fragment length polymorphism.

Morphological similarity among E. trachycaulus, E. virescens, E. violaceus, and E. hyperarcticus has often been noted. Taxonomists have tried to discriminate among these taxa using morphological characters and a number of different relationships among them have been suggested. However, the genetic relationships among these taxa are still unknown. AFLP analysis was used to characterize the molecular diversity of these taxa and to examine genetic relationships among them. A high degree of genetic identity was apparent among 7 accessions of E. virescens. The similarity values ranged from 0.90 to 0.99 with an average of 0.94. The mean similarity values among 3 E. hyperarcticus and among 5 E. violaceus accessions were 0.84 (0.81-0.87) and 0.77 (0.66-0.90), respectively. The similarity values among 17 E. trachycaulus accessions ranged from 0.49 to 0.92 with an average of 0.75. The 5 accessions of E. subsecundus displayed high variation, with similarity values between 0.52 and 0.68 and a mean value of 0.59. Both maximum-parsimony (MP) and neighbor-joining (NJ) analyses showed that all 7 accessions of E. virescens formed a clade, indicating a monophyletic origin. On the other hand, Elymus trachycaulus, E. subsecundus, and E. violaceus were each paraphyletic and separated into different genetically distinct groups. Among these 5 taxa, E. virescens was genetically similar to E. trachycaulus, and E. violaceus was genetically similar to E. hyperarcticus.     

Genomic relationships among diploid and hexaploid species of Andropogon (Poaceae).

Andropogon is a pantropical grass genus comprising 100-120 species and found mainly in the grasslands of Africa and the Americas. While the genomic relationships between many Andropogon species have been resolved by studying chromosome behavior in interspecific hybrids, relationships between the North and South American diploids have remained elusive. Further, the genome composition of two hexaploid species (including the important forage grass Andropogon lateralis Nees) has been unclear because of the strong hybridization barriers that exist between species. Consequently, genomic in situ hybridization was applied to shed light on these issues. The results confirmed that (i) both the South American (Andropogon selloanus (Hack.) Hack., Andropogon macrothrix Trin.) and North American (Andropogon gyrans Michx.) diploid species shared a common S genome and (ii) the S genome comprises just one of the three genomes in the hexaploids A. lateralis Nees and Andropogon bicornis L. The evolutionary and taxonomic implications of these findings are discussed.     

Biosystematic studies of the Elymus hispidus (Poaceae: Triticeae) group in Iran

Biosystematic studies on the Elymus hispidus group included investigations on morphological characters and variability in cultivation, crossing experiments, hybrid fertility and meiotic configuration in hybrids. Elymus gentry! is a distinct species with a different altitudinal distribution from that of E. hispidus. Morphological characters which have been used to split E. hispidus into distinct species or subspecies are not constant in cultivation. The majority of the cultivated plants were morphologically similar to the maternal individuals from which seed was collected in the field. However, in some accessions the offspring represented different morphological variants when cultivated. Hybrids between different variants were fertile and showed regular meiosis. Different variants within the E. hispidus group tend to grow in different habitats and as characters to some extent show constancy in cultivation, three varieties, var. hispidus , var. podperae and var. villosus are recognized for E. hispidus in Iran.     

Intergeneric crosses of Psathyrostachys huashanica with Elymus spp. and cytogenetic studies of the hybrids with E. tsukushiensis (Poaceae, Triticeae)

Intergeneric crosses have been made between Psathyrostachys huashanica (2x= 14, NN) and three Elymus species, namely, E. pendulinus (2n = 4x = 28, SSYY), E. ciliaris (2n = 4x = 28, SSYY), and E. tsukushiensis (2n = 6x = 42, SSHHYY). Three accessions of E. tsukushiensis from different localities crossed with P. huashanica produced adult hybrid plants. Although completely sterile, the hybrid plants developed rather vigorously, and were morphologically intermediate between the two parents. Chromosome configuration of 24.62I + 1.51II + 0.03III per cell characterized meioses of the hybrids, but there existed slight variations among different combinations. These results indicate that there is little or no chromosome homoeology between "N" genome of P. huashanica and "S", "H" or "Y" genomes of E. tsukushiensis.     

The origin of the H, St, W, and Y genomes in allotetraploid species of Elymus L. and Stenostachys Turcz. (Poaceae: Triticeae)

Based on sequences from two single-copy nuclear genes (DMC1 and EF-G), four plastid genes (rbcL, rpoA, matK, and ndhF), and one mitochondrial gene (coxII), we investigate the origin of the H, St, W, and Y genomes in four allotetraploid species of Elymus and two allotetraploid species of Stenostachys. Despite significant incongruence between the two nuclear genes and between the nuclear and organelle data partitions, individual and combined analyses of the data partitions unequivocally show that the St and H genomes of the tetraploid American species of Elymus are derived from Pseudoroegneria and Hordeum, respectively, with Pseudoroegneria serving as the female parent, and that the H and W genomes of Stenostachys are derived from Hordeum and Australopyrum, respectively, with Hordeum serving as the female parent. The analyses equally clearly demonstrate that the St genome of the tetraploid Asiatic Elymus species is derived from Pseudoroegneria, with the latter serving as the female parent, but the relationship of the Y genome is less clear. Individual analyses of the nuclear genes provide conflicting results, but combined analysis of all data suggests a sister group relationship to Heteranthelium, albeit without any jackknife support.     

披碱草属(Elymus L.)两个物种之订正

披碱草属Elymus L．是小麦族Triticeae Dumort．中最大的属,全世界共有150多个种,广泛分布于 温带地区。我国约有80余种(包括鹅观草属Roegneria C．Koch),是披碱草属的重要分布区和多样性分 化中心。由于披碱草属植物种类繁多、分布广泛、生境多样以及形态变异较为复杂,导致该属在分类上 存在许多问题。同时,由于地域的局限和地区间交流的缺乏,致使同一种披碱草属植物被不同的学者多 次发表及同物异名现象的出现。本文对E．antiquus和E．burchan-buddae这两个曾多次被订名的披碱草 属物种进行了分类订正。