长芒猬草与华山新麦草属间杂种的形态学和细胞学研究

为研究长芒猬草Hystrix duthiei ssp．longearistata的染色体组成,将其与华山新麦草Psathyrostachys huashanica进行了人工杂交,获得杂种F₁。对亲本及杂种F₁,花粉母细胞减数分裂染色体配对行为、繁育 特性和形态特征进行了比较分析。结果表明：杂种F₁的许多形态特征介于父母本之间,花粉完全不育, 结实率为0;杂种F₁花粉母细胞减数分裂中期Ⅰ染色体配对较高,55.12％的细胞形成5个或6个二价 体,其构型为：9.83Ⅰ+5.46Ⅱ+0.07Ⅲ,C-值为0.57。以上结果表明H．duthiei ssp．Longearistata含有Ns染色体组。本文还讨论了Hystrix与Leymus的关系。     

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。     

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.     

Meiotic analysis ofElymus caucasicus,E. longearistatus,and their interspecific hybrids with twenty-threeElymus species (Triticeae,Poaceae)

Interspecific hybridizations were carried out between the two tetraploidsElymus caucasicus andE. longearistatus, and 23 tetraploids and hexaploids ofElymus containing SH, SY, SYH, and SYW genomes and representing various geographical regions. Meiotic pairing was studied in the two target species and their hybrids. It is concluded from this study that (i) interspecific hybridization is fairly easy to perform although strong reproductive barriers exist between the species; (ii)Elymus caucasicus andE. longearistatus are allotetraploids, and share the diverged SY genomes; (iii) the divergence of SY genomes is correlated with the geographic distance between theElymus spp. studied.     

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.     

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.     

Meiotic behaviour in three interspecific three-way hybrids between Brachiaria ruziziensis and B. brizantha (Poaceae: Paniceae)

The meiotic behaviour of three three-way interspecific promising hybrids (H17, H27, and H34) was evaluated. These hybrids resulted from the crosses between B. ruziziensis X B. brizantha and crossed to another B. brizantha. Two half-sib hybrids (H27 and H34) presented an aneuploid chromosome number (2n = 4x = 33), whereas hybrid H17 was a tetraploid (2n = 4x = 36), as expected. Chromosome paired predominantly as multivalents suggesting that genetic recombination and introgression of specific target genes from B. brizantha into B. ruziziensis can be expected. Arrangement of parental genomes in distinct metaphase plates was observed in H27 and H34, which have different male genitors. Hybrids H17 and H34 have the same male genitor, but did not display this abnormality. In H17, abnormalities were more frequent from anaphase II, when many laggard chromosomes appeared, suggesting that each genome presented a different genetic control for meiotic phase timing. Despite the phylogenetic proximity among these two species, these three hybrids presented a high frequency of meiotic abnormalities, mainly those related to irregular chromosome segregation typical of polyploids, H34, 69.1%; H27, 56.1% and H17, 44.9%. From the accumulated results obtained through cytological studies in Brachiaria hybrids, it is evident that cytogenetical analysis is of prime importance in determining which genotypes can continue in the process of cultivar development and which can be successfully used in the breeding. Hybrids with high frequency of meiotic abnormalities can seriously compromise seed production, a key trait in assuring adoption of a new apomictic cultivar of Brachiaria for pasture formation.     

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.     

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.     

Genome analysis of species in the genus Hystrix (Triticeae; Poaceae)

Genomic in situ hybridisation (GISH) and Southern genomic hybridisation were applied in order to gain further knowledge regarding generic delimitation of the genus Hystrix as well as to clarify the genomes of the Hystrix species H. patula, H. longearistata, H. coreana, H. duthiei and H. komarovii. The hybridisation intensity of different genomic probes was compared among the Hystrix species and with other Triticeae species. The Southern- and GISH results confirm that H. patula contains the StH genome and show that H. komarovii most likely has a variant of this StH genome. The other Hystrix species under study, i.e. H. longearistata, H. coreana and H. duthiei, contain an Ns basic genome, and most probably two variants of this basic genome, Ns ¹ Ns ² . The genus Hystrix is thus not a monophyletic group of species.     

Morphological,cytological, and molecular evidences for natural hybridization between Roegneria stricta and Roegneria turczaninovii (Triticeae: Poaceae)

Some plants with low fertility are morphologically intermediate between Roegneria stricta and Roegneria turczaninovii, and were suspected to be natural hybrids between these species. In this study, karyotype analysis showed that natural hybrids and their putative parents were tetraploids (2n = 4x = 28). Meiotic pairing in natural hybrids is more irregular than its putative parents. Results of genomic in situ hybridization and fluorescence in situ hybridization indicate that natural hybrids contain the same genome as their putative parents. The nuclear gene DNA meiotic recombinase 1 (DMC1) and the chloroplast gene rps16 of natural hybrids and their putative parents were analyzed for evidence of hybridization. The results from molecular data supported by morphology and cytology demonstrated that the plants represent natural hybrids between R. stricta and R. turczaninovii. The study is important for understanding species evolution in the genus since it demonstrates for the first time the existence of populations of natural homoploid hybrids in Roegneria. The study also reports for the first time that the composition of the genomic formula of R. turczaninovii is StY, confirming that the current taxonomic status is correct.     

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.     

Discovery of the genus Pseudoroegneria (Triticeae,Poaceae) in the Western Mediterranean on exploring the generic boundaries of Elymus

In this study, we review the classification of two species, Elymus hispanicus and E. marginatus, which are restricted to highly valuable and sensitive Mediterranean ecosystems. The genomic composition of the two species is analysed by in situ hybridization. In addition, lodicule morphology and foliar anatomy of both species are compared with those of E. caninus, E. repens, E. sibiricus (i.e., the type species of Elymus s.s.) and Pseudoroegneria strigosa (i.e., the type species of Pseudoroegneria). The genomic formula 2n = 8x = 56; HStStSt is proposed for E. hispanicus and 2n = 4x = 28; StSt for E. marginatus. In this latter species, the absence of the ribosomal genes in one of the two St genomes suggests that diploidization may have occurred during the evolution the species. Regarding foliar anatomy, E. hispanicus, E. caninus, E. repens, and E. sibiricus shared several characteristics, but the leaf blades of E. marginatus proved anatomically more similar to those of Ps. strigosa. The data compiled support the contention that: (1) E. hispanicus belongs to Elymus s.s.; (2) E. marginatus should be transferred to Pseudoroegneria; and (3) the morphology of the lodicules should be carefully reconsidered for appropriately describing the boundaries between Elymus s.s. and Pseudoroegneria. The new combination Ps. marginata is proposed and a detailed iconography of the plant is provided.     

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.     

Morphology,fertility and cytogenetics of intergeneric hybrid between Roegneria kamoji Ohwi and Dasypyrum villosum (L.) Candargy (Poaceae: Triticeae)

The intergeneric hybrids between Roegneria kamoji Ohwi and Dasypyrum villosum (L.)Candargy were successfully obtained by means of embryo culture in vitro. Studies on morphology, fertility and chromosome pairing behavior in meiosis of the parents and their hybrid F_l were carried out in the present work. The results showed that: (1) there were ob vious morphological differences between R. kamoji and D. villosum, and spikes of F_l plants were morphologically intermediate between the two parental species; (2) the seed set of the cross was 11.63%; the hybrid plant was infertile, which indicated that strong repro ductive isolation existed between the parents and R. kamoji and D. villosum were inde pendent biological species; (3) The somatic chromosome number in root-tips of F_l hybrids was 28. Chromosome pairing at MI of PMCs in F_l hybrids was quite low. The meiotic con figuration was 26.72 Ⅰ + 0.62 Ⅱ + 0.02 Ⅲ, which indicated that very low homoeology was detected between the St, H, Y genomes of R. kamoji and the V genome of D. villo- sum, and the relationship between the parental species was remote.     

鹅观草和簇毛麦属间杂种的形态学和细胞遗传学研究

通过幼胚离体拯救培养,成功地获得鹅观草Roegneria kamoji Ohwi(2n=6x=42,StStHHYY)和 簇毛麦Dasypyrum villosum(L．)Candargy(2n=2x=14,VV)属间杂种,对这两个种及其杂种F_l的形 态学、繁育学和减数分裂配对行为进行了研究。结果表明：(1)鹅观草和簇毛麦形态差异极大,杂种F1 的形态特征介于父母本之间;(2)杂交结实率低,为11．63％;经幼胚组织培养,获2株杂种苗;杂种F_l高 度不育,表明亲本间存在极强的生殖隔离,是独立的生物学种;(3)杂种F_l体细胞染色体数目为稳定的 28条,减数分裂中期I染色体配对很低,其构型为：26．72Ⅰ+0．62Ⅱ+0．02Ⅲ。表明鹅观草的St、H、Y染色体组与簇毛麦的V染色体组部分同源性很低,它们的亲缘关系较远。     

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.     

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.     

Genome analysis of intergeneric hybrids of apomictic and sexual Australian Elymus species with wheat,barley and rye: implication for the transfer of apomixis to cereals

Summary Four hybrids were obtained between three Australian Elymus taxa and three cereal grains: wheat, rye, and barley. Mean meiotic metaphase-I configurations were 41.14 I, 0.42 rod II, 0.003 ring II, and 0.01 III for E. scabrus var plurinervis x Triticum aestivum (1 hybrid plant), 22.27 I, 2.63 rod II, 0.06 ring II, and 0.12 III for E. scabrus var scabrus x Secale cereale (4 hybrid plants), and 26.65 I, 0.66 rod II, 0.00 ring II, and 0.01 III for E. scabrus var plurinervis x Hordeum vulgare (13 hybrid plants). The I genome of barley also paired very little in a B_III hybrid of apomictic E. rectisetus x H. vulgare (2 hybrid plants). Megasporogenesis in this B_III hybrid was at least facultatively apomeiotic, with the same sort of nuclear elongation, apomeiotic division, and dyad formation seen previously in E. rectisetus itself. All four hybrid combinations were sterile. While spike morphology in the E. scabrus x T. aestivum and E. scabrus x H. vulgare hybrids were intermediate to their parents, E. scabrus x S. cereale and E. rectisetus x H. vulgare looked like their maternal parents.