Elymus sibiricus, E. nutans 和 E. burchan-buddae 的形态学鉴定及其染色体组亲缘关系的研究

为了进一步研究Elymus sibiricus L.、E．nutans Griseb．和E．burchan-buddae(Neuski)Tzelev [＝Roegneria nutans(Keng)Keng]的外部形态差异及其系统学关系,本文对这三种植物的6个穗部形 态性状进行了观测和比较,并对这三个Elymus种进行了种间杂交及杂种F1的减数分裂染色体配对行 为的分析研究。结果表明：这三个Elymus种的穗长及颖长等性状均变异很大,而内稃的长、宽则变异 不大并具有明显的种间差异。E. nutans×E．barchan-buddae及E.nutans×E．sibiricus的杂种F1均完 全不育,减数分裂不规则。E．nutans×E．burchan-buddae杂种F1的减数分裂构型为：7.70I+13.40 Ⅱ+0.06Ⅲ+O.08 Ⅳ,而E．sibiricus×E．nutans杂种F1的构型为11.98 Ⅰ+9.61Ⅱ+O.64Ⅲ+0.39 Ⅳ+0.01V。由本实验的形态学和细胞学的研究结果得出以下结论：1．利用内稃形态性状结合穗部 其它性状的差异能对这三个物种进行较准确的鉴定;2．E．nutans与E.burchan-buddae的亲缘关系较 近,而E．nutans与E．sibiricus的亲缘关系则较远;3. E,burchan-buddae×E．nutans的杂种Fl中存在着染色体配对控制因子。     

Biosystematic studies of Kengyilia melanthera and K. kokonorica (Poaceae: Triticeae)

Kengyilia melanthera (Keng)J. L. Yang, Yen et Baum and K. kokonorica (Keng)J. L. Yang, Yen et Baum are two hexaploid perennial grasses of the tribe Triticeae native in west China. K. melanthera and K. kokonorica were hybridized with Roegneria kamoji Ohwi(2n=42,StStHHYY) and K. hirsuta (Keng)J. L. Yang, Yen et Baum (2n = 42, PPStStYY) respectively. Chromosome pairing behaviour at metaphase I in the parents and hybrids was studied. Meiotic configurations were 18.20 Ⅰ + 11.74 Ⅱ + 0.09 Ⅲ + 0.01V for R. kamoji×K. melanthera, 1.06Ⅰ + 20.47 Ⅱ for K. hirsuta×K. melanthera, 19.36Ⅰ + 11.26 Ⅱ + 0.04Ⅲ for R. kamoji×K. kokonorica, and2.46Ⅰ + 19.44Ⅱ + 0.14 Ⅲ + 0.06 Ⅳ for K. hirsuta×K. kokonorica. Considering chromosome pairing in the hybrids, as well as morphological characters, K. melanthera and K. kokonorica should be grouped in Kengyilia Yen et J. L. Yang instead of being keeped inRoegneria sect. Paragropyron Keng , or in Elymus L. or Elytrigia Desv.     

Genomic constitutions of four Chinese endemic Elymus species: E. brevipes, E. yangii, E. anthosachnoides, and E. altissimus (Triticeae, Poaceae).

The objectives of this study were to determine the genomic constitution and to explore the genomic variation within four Chinese endemic Elymus species, i.e., E. brevipes (Keng) L?ve (2n = 4x = 28) and E. yangii B.R. Lu (2n = 4x = 28), E. anthosachnoides (Keng) L?ve (2n = 4x = 28), and E. altissimus (Keng) L?ve (2n = 4x = 28). Intraspecific crosses between different populations of the four Elymus species, as well as interspecific hybridizations among the four target species, and with six analyzer species containing well-known genomes, i.e., E. caninus (L.) L. (2n = 4x = 28, SH), E. sibiricus L. (2n = 4x = 28, SH), E. semicostatus (Lees ex Steud.) Melderis (2n = 4x = 28, SY), E. parviglumis (Keng) L?ve (2n = 4x = 28, SY), E. tsukushiensis Honda (2n = 6x = 42, SHY), and E. himalayanus (Nevski) Tzvelev (2n = 6x = 42, SHY), were achieved through the aid of embryo rescue. Chromosome pairing behaviors were studied in the parental species and their hybrids. Numerical analysis on chromosome pairing was made on the interspecific hybrids. With one exception, each meiotic configuration at metaphase I in the hybrids involving the target taxa and the analyzer species containing the "SH" genomes fit a 2:1:1 model with x-values ranging between 0.91 and 1.00; chromosome pairing in the hybrids involving analyzer parents with the "SY" genomes match a 2:2 model, with x-values between 0.97 and 0.99. All pentaploid hybrids with a genomic formula "SSYYH," except for two crosses having unexpected low c-values, had pairing patterns fitting the 2:2:1 model with x-values varying between 0.96 and 1.00. It is concluded based on hybridization, fertility, and chromosome pairing data that (i) the four target Elymus species are strictly allotetraploid taxa, (ii) they are closely related species, all comprised of the "SY" genomes, (iii) minor genomic structural rearrangements have occurred within the four Elymus species, and (iv) meiotic pairing regulator(s) exists in some of the Elymus taxa studied.     

利用SSR标记对垂穗披碱草和老芒麦进行物种鉴定和遗传变异分析

垂穗披碱草(Elymus nutans)和老芒麦(Elymus sibiricus)皆为花序下垂类披碱草属物种,高海拔地区的E.sibiricus的部分变异类型也具有小穗紧密排列等特征,与E.nutans在形态学性状上具有较多的交叉,造成野外种质资源采集时进行直接的田间鉴定存在困难。本研究利用12对小麦族SSR引物对8份垂穗披碱草和10份老芒麦种质进行遗传变异和物种鉴定分析,UPGMA聚类分析表明供试材料明显可依据物种差异划分成两大类,主向量分析(PCo A)与聚类分析的结果保持一致。种质间遗传相似系数分析和分子方差分析(AMOVA)也表明种间变异远高于种内。另外,本研究筛选出3对引物,ESGS79和ESGS155能够在垂穗披碱草材料中扩增出特异性条带,Xgwm311能够在老芒麦材料中扩增出特异性条带,这3对引物能够作为区分垂穗披碱草和老芒麦的依据,为野外种质资源的收集以及田间育种工作提供指导。     

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.     

老芒麦和垂穗披碱草rDNA-ITS序列分析

分析了老芒麦和垂穗披碱草rDNA-ITS序列,为这两个种的鉴别提供分子指纹图谱,为系统发育提供分子生物学依据。结果表明,22份材料的ITS、ITS1、5.8S、ITS2序列长度均相同,依次为604bp、222bp、164bp、218bp,GC含量依次为62.25%～63.08%,62.16%～63.06%,59.76%,64.22%～65.60%。除5.8S外,碱基位点都有不同程度的变化,一些变异位点有明显的种性变异规律,可作为老芒麦和垂穗披碱草种质DNA指纹特异鉴别位点。7个ITS序列的同源性98.7%～99.8%,遗传分歧0.2～1.3,具有保守性,为非近期分化类群。系统发育分析表明不同来源地的同种材料,差异主要表现在DNA变化快慢及碱基替换数,但相同的种归为一类,进一步反映了种内的遗传稳定性。     

根据形态和叶片微形态特征讨论无芒披碱草的归并

通过形态学观测和叶片解剖特征分析,比较了无芒披碱草、短芒披碱草及老芒麦3个近缘种的主要性状差异,以探讨无芒披碱草的系统分类归属.结果表明,在外部形态上无芒披碱草与短芒披碱草差异甚小,难以进行区分,但与老芒麦差异明显,是典型的种间关系;在叶片解剖上,无芒披碱草的绝大多数特征与短芒披碱草的一致或类同,可与老芒麦的却存在明显间断.故研究认为:无芒披碱草与短芒披碱草是同一个种,无芒披碱草应作为短芒披碱草的异名.     

Interspecific hybridizations in the Elymus semicostatus group (Poaceae).

Meiotic pairing in 16 interspecific hybrids in the genus Elymus is reported. The hybrids were made among seven species in the Elymus semicostatus group, viz., E. semicostatus, E. validus (subgroup I), E. abolinii (subgroup II), E. fedtschenkoi, E. nevskii, E. praeruptus (subgroup III), and E. panormitanus (subgroup IV). All species are tetraploid (2n = 4x = 28) and possess the SY genomes. Meiotic pairing was distinctly higher in hybrids made within subgroups than between subgroups, but the genomes in E. panormitanus have differentiated from those in the other species. These results generally support the subdivision of the E. semicostatus group based on morphological data but also indicate that the subgroups are more distantly related than previously believed, and that the group may be nonmonophyletic.     

Physical mapping of repetitive sequences and genome analysis in six Elymus species by in situ hybridization

Genome constitution and genetic relationships between six Elymus species were assessed by physical mapping of different repetitive sequences using a technique of sequential fluorescence in situ hybridization and genomic in situ hybridization.The six Elymus species are all naturally growing species in northwest China,namely,E.sibiricus,E.nutans,E.barystachyus,E.xiningensis,E.excelsus,and E.dahuricus.An StStHH genome constitution was revealed for E.sibiricus and StStHHYY for the remainder species.Each chromosome could be clearly characterized by physical mapping with 18S-26S rDNA,5S rDNA,Afa-family,and AAG repeats,and be allocated to a certain genome by genomic in situ hybridization.Two 5S rDNA sites,each in the H and St genomes,and three 18S-26S rDNA sites,two in the St genome and one in the Y genome,were uncovered in most of the species.The strong Afa-family hybridization signals discriminated the H genome from the St and Y genomes.The H and Y genome carried more AAG repeats than St.A common non-Robertsonian reciprocal translocation between the H and Y genomes was revealed in E.barystachyus,E.xiningensis,E.excelsus and E.dahuricus.Comparison of molecular karyotypes strongly suggests that they can be classified into three groups,namely,E.sibiricus,E.nutans,and others.     

Biosystematic study of hexaploids Elymus tschimganicus and E. glaucissimus. II. Interspecific hybridization and genomic relationship.

To investigate genomic relationships of Elymus tschimganicus (Drobov) Tzvelev (2n = 6x = 42, S1S2Y genomes) and E. glaucissimus (M. Pop.) Tzvelev (2n = 6x = 42, S1S2Y genomes), interspecific hybridizations of the two target species were carried out with 27 other Elymus species containing the SH, SY, SYH, SYP, SYW, and SH1H2 genomes, respectively, collected from different geographic regions. Chromosome pairing behavior was analyzed at metaphase I in 27 hybrids representing 23 hybrid combinations, and overall genomic relationships of the two target species with the other Elymus taxa were estimated. The study concluded that (i) interspecific hybridization was principally easy to perform between the Elymus species, but no general pattern of crossability was obtained, and all hybrids were completely sterile, (ii) the two species have a similar meiotic pattern in their hybrids with the other Elymus species, and (iii) species containing the SY, SYP, and SYH genomes have a generally higher level of genomic homology to the target species than those possessing the SH genomes, and the South American hexaploid with the SH1H2 genomes has the lowest level of genomic homology to the two target taxa.     

A Study on the Spore Morphology of Bryum Hedw. in China

Spore morphology of thirteen species of the genus Bryum Hedw. were observed by LM and SEM. The results show that the ornamentation of spore exine could be divided into three types: Type I , blunt at the top of baculate processes, to which four species belong: Bryum argenteum, B. lonchocaulon , B. uliginosum and B. arcticum. Type Ⅱ, sharp or with small processes at the top of baculate processes, represented by seven species: B. pallescens, B. caespiticium, B. pallens, B. pseudotriquetrum, B. paradoxum, B. alpinum and B. thomsoii. Type Ⅲ, expanded into hemispherical-shaped at the top of baculate processes, represented by two species: B.coronatum and B.sauteri. The Bryum species may also be divided into three groups according to the variation of spore diameter. Group I , with spore diameter under 10 μm, including one species, B. uliginosum. Group Ⅱ, with spore diameter 11～20 μm, including seven species: B. argenteum, B. alpinum, B. coronatum, B. pallens, B.paradoxum, B.sauteri, B.thomsonii. Group Ⅲ, spore diameter 21～30 μm, with five species: B. pallescens, B. caespiticium, B. pseudotriquetrum, B. lonchocaulon, B. arcticum. There are resemblances of spore morphology and exine ornamentation among the thirteen species. In the view of palynology, the genus Bryum is a natural taxon which is more advanced than the genus Pohlia Hedw. But spores of thirteen species are different at some characters such as diameter, shape ofproximal leptoma, etc., which indicates the genetic differentiation in the genus Bryum.     

曲穗草属天然杂种的鉴定及形成研究

本研究通过形态学、细胞学、系统发育分析等方法对小麦族曲穗草属(Campeiostachys)的两株疑似天然杂种进行鉴定,探究其可能的亲本及杂种形成机制.结果显示:两株天然杂种在形态上与短芒曲穗草(Cam-peiostachys breviaristata(Keng)Y.H.Zhou,H.Q.Zhang et C.R.Y...     

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。     

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.     

Cytological Studies of Hybrids of Roegneria ciliaris with R.alashanica and R.magnicaespes(Poaceae:Triticeae)

In order to investigate the relationships between Roegneria.ciliaris(Trin.)Nevski and R.alashanica Keng,and between R.ciliaris and R.magnicaespes D.F.Cui.R.ciliaris(Trin.)Nevski was crossed with the other two species,through the aid of embryo rescue. Hybrids were obtained from these two combinations.Chromosome pairing behavior of parents an dhybrids was observed at metaphase I of pollen mother cells. Meiotic configuratons were 10.62Ⅰ＋8.17Ⅱ＋0.32Ⅲ＋0.02Ⅳ for R.ciliaris×R.magnicaespes and 18.00Ⅰ＋4.76Ⅱ＋0.16Ⅲ for R.ciliaris ×R.magnicaespes;c-value of the two hybrids were 0.44 and 0.21,respectively.     

加拿大披碱草、老芒麦及其杂种F1代的RAPD分析

加拿大披碱草和老芒麦是披碱草属地理性远缘的2个种。采用RAPD技术,筛选了16个随机引物对这2个种及其F1代进行遗传多态性检测。在获得的186个位点中,114个为多念性位点,占61．29％。从种群内的遗传多态性来看,加拿大披碱草的多态性位点百分率最高(242．04%),其次是老芒麦(9．14%),杂种F1(1．61％)。从种群间的遗传关系来看,加拿大披碱草和老芒麦的遗传距离为0．4979,杂种F1与母本加拿大披碱草的遗传距离(0．2720)较近,而与父本老芒麦的遗传距离(0.4074)相对较远。     

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.     

Morphological characteristics of fruits of Fagopyrum (Polygonaceae) from China

Macro- and micro-morphological characteristics of fruits in eight species and one variety of the genus Fagopyrum Mill. (Polygonaceae) from China were observed under stereoscope and scanning electron microscope(SEM). Based on the results, the fruits of the species studied are divided into type Ⅰ , Ⅱ and Ⅲ. The fruits of type Ⅰ are triangular-pyramidal; their surface are rugosely reticulate, neither smooth nor shiny. Two species, F. tataricum and F. dibotrys have this fruit type. Those of type Ⅱ are ovoid-triangular-pyramidal; their surface are smooth and shiny, and striately reticulate. Three species, F. esculentum, F. statice and F. lineare, have this fruit type. In type Ⅲ, the fruits are ovoid-triangular-pyramidal; their surface are smooth and shiny, and covered with many warty grains and sparsely finely striate. F. urophyllum, F. gracilipes, F. leptopodum var. leptopodum, and F. leptopodum var. grossii have this fruit type. Judging from the morphological characteristics of fruits, F. dibotrys might be more closely related to F. tataricumthan to F. esculentum.     

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.     

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.