Intergeneric hybridizations with Eremopyrum (Poaceae)

Intergeneric crosses were made between the four species of Eremopyrum (2n = 14, 28) and 16 species (2n = 14, 28, 42) from 8 genera of the Triticeae. Seed set was obtained in 22 of 42 different combinations. Only 9 resulted in progeny and only one was diploid. The hybrids were morphologically intermediate between the parents and it has been confirmed that perenniality dominates over annuality. The very low pollen fertility of the hybrids was caused by very low chromosome pairing in meiosis and supports the uniqueness of the Eremopyrum genome. Autosyndesis between the chromosomes of E. orientale is proposed and for that reason reevaluation of the relationships within Eremopyrum is needed.     

Taxonomic studies in Dasypyrum (Poaceae)

Dasypyrum (tribe Triticeae) is revised. Two species are recognized: the annual D. villosum (2n = 14) and the perennial D. breviaristatum (2n = 28). The typification of D. villosum is discussed. The combination D. hordeaceum is demonstrated to be based on a later homonym and for that reason a new combination, D. breviaristatum , is made. A key and a map showing the distribution of the species are presented. Interspecific hybridization and the phylogenetic relationships of the genus are discussed.     

普通小麦×东方旱麦草属间杂种的产生及无性系的建立

本研究以普通小麦（ＴｒｉｔｉｃｕｍａｅｓｔｉｖｕｍＬ．;２ｎ＝６ｘ＝４２,ＡＡＢＢＤＤ）为母本,以东方旱麦草（Ｅｒｅｍｏｐｙｒｕｍｏｒｉｅｎｔａｌｅ（Ｌ．）Ｊａｕｂ．ｅｔＳｐａｃｈ;２ｎ＝４ｘ＝２８）为父本,首次成功地获得了属间远缘杂种Ｆ１,其平均结实率为０．０８％。利用植物细胞工程技术,对杂种幼胚愈伤组织的诱导、胚性无性系的建立、植株再生、壮苗培养等,最终获得了生长正常的杂种Ｆ１植株。同时,通过对杂种幼胚愈伤组织、根尖细胞的细胞学观察,结果表明该杂种为真杂种,即２ｎ＝５ｘ＝３５（预期染色体数）的杂种细胞占主体;另外,因组培过程中发生了染色体数目的变异,故也有少量２ｎ＝２８－３４染色体数的细胞。以上杂种的获得为将旱麦草优异基因向小麦的转移奠定了基础。     

Two new Tibetan species of Elymus (Poaceae: niticeae) and their genomic relationships

Two Elymus species, E. cacuminus (2n = 4x = 28) and E. retroflexus (2n = 2x = 28). native to western China are described as species new to science. In order to determine genomic constitutions of the two species, intergeneric and interspecific hybridizations were carried out between the two new species and Pseudoroegneria spicata (2n = 2x = 14, SS), and sixteen other Elymus species, fifteen tetraploids (2n = 4x = 28) and one hexaploid (2n = 6x = 42), containing the "SH", "SY", and "SYH" genomes. Chromosome pairing behaviour was studied in the two species and their hybrids. Generally low meiotic pairing was observed in the hybrids with S- and SH-genome species and high pairing in the hybrids with SY- and SYH-genome species. It is concluded from this study that (i) the two new species are strict allotetraploid (2n = 4x = 28) and self-pollinating taxa (inbreeders) in nature; (ii) they contain the "SY" genomes and; (iii) they have close genomic relationships with the SY-genome Elymus species from the same or geographically adjacent areas.     

新疆多年生小麦族植物染色体数的观察

本文对1987年采集于新疆的多年生小麦族(Triticeae Dum. )属种进行了细胞学观察。该地区多年生小麦族各属种的染色体数目变化范围是从2n=14到2n=84,前者主要存在于大麦属(Hordeum)、新麦草属(Psathyrostachys),而后者全部集中于赖草属(Leymus).其中染色体数目为2n=28和2n=42的频率最高,主要存在于鹅冠草属(Roegneria)和披碱草属(Elymus)。Roegneria gobicola, Roegneria kuqaensis, Roegneria tahelacona, Roegneria zhoasuensis的染色体数为首次报道。     

准噶尔荒漠3种短命植物气体交换特征的日变化

采用LI-6400便携式光合测定仪,在晴天条件下对准噶尔荒漠3种典型短命植物东方旱麦草(Eremopyrum orientale)、卷果涩芥(Malcolmia scorpioides)和四齿芥(Tetracme quadricormis)的气体交换特征的日变化规律进行了研究.结果表明:①东方旱麦草和卷果涩芥的净光合速率(简称Pn,后同)的日变化呈"双峰"型,14:00(采用时间均为北京时间,后同)左右存在明显的光合"午休"现象,四齿芥Pn的日变化呈"单峰"型,峰值出现在12:00与前两者的第一峰值出现时刻相同.3种植物蒸腾速率(简称E,后同)的日变化均呈"单峰"型,但不同植物的峰谷值出现时刻不同.水分利用效率(WUE)日变化,四齿芥呈"单峰"型,东方旱麦草和卷果涩芥呈"双峰"型,峰值分别出现在8:00～10:00之间,后两者第二峰值分别出现在16:00和18:00.②根据Pn、胞间CO2浓度(Ci)和气孔限制值(Ls)的变化方向,推测3种短命植物的光合"午休",东方旱麦草和卷果涩芥主要受非气孔因素限制,而四齿芥主要受气孔因素限制.③卷果涩芥和四齿芥两种十字花科草本日平均Pn、E 和WUE 均高于禾本科东方旱麦草,尤其卷果涩芥是一种高光合、高蒸腾、高水分利用率的物种.④相关分析结果表明,对Pn影响最显著的环境因子是光合有效辐射(PAR),对E影响最显著的因子3种植物各不相同.     

Karyotypes of Eleven Species of Triticeae in Northeast China

In this paper the karyotypes of eleven species of Triticeae from Northeast China are reported. The karyotype formulae are as follows: Agropyron cristatum, 2n=4x=28=20m+8sm; Elytrigia repens, 2n=6x=42=34m(2SAT) + 8sm; Hordeum brevisubulatum, 2n = 4x = 28 = 20m + 8sm( 4SAT ); Roegneria nakaii, 2n = 4x = 28 = 20m + 8sm( 4SAT ); R. turczaninovii var. macrathera, 2n = 4x = 28 = 20m(2SAT ) + 8sm(2SAT ); Elymus sibiricus, 2n = 4x = 28 = 20m + 8sm ( 4SAT); E. dahuricus, 2n=6x=42=32m+10sm( 6SAT); E. excelsus, 2n=6x=42=32m+10sm( 6SAT); Leymus chinensis, 2n=4x=28=20m(4SAT) + 8sm; Roegneria ciliaris, 2n = 4x = 28 = 22m( 2SAT ) + 6sm( 2SAT ); R. kamoji, 2n= 6x = 42= 30m+ 12sm(4SAT). The karyotypes of the first five species are re-ported for the first time.     

Chromosome Number and Development of Gametophytes in Euptelea pleiospermum (Eupteleaceae)

Euptelea Sieb. et Zucc. is a genus of 2 species, endemic to East Asia and disjunctly distributed in China and Japan. The present paper deals with cytology and embryology of the Chinese species E.pleiospermum Hook. f. et Thoms., providing materials for discussing the classificatory rank and relationships of the genus. This work reports the chromosome number, n = 14 and 2n = 28 (Plate 1: 1, 2), for E.pleiospermum, which is consistent with that of the Japanese species, E. plyandra Sieb. et Zucc.,reported by Sugiure (1931) and Whitaker (1933). This number is different from those of the related genera, i. e. n=20 or 2n=40 in Trochodendron and 2n = 48(46)in Tetracentron (Cronquist, 1981), and, therefore, the cytological evidence supports the treatmemt of the genus as a separate family by Smith (1945, 1946), Chrlotte and Bailey (1946), Cronquist(1981), Thorne (1983), Dahlgren(1983), Wang (1984). The ovule of E. pleiospermum is anatropous, bitegminous and crassinucellate (Plate 1: 10), the characters which have been already reported by Davis (1966). In addition, the present work shows that in E. pleiospermum the tapetum is glandular, consisting of 2-or 4-nucleate tapetal cells; cytokinesus at meiosis of pollen mother cells is simultaneous; microspore tetrads are mainly tetrahedral; pollen grains are 2-celled (Plate 1: 4, 5, 6, 9), the third megaspore from the micropyle is functional and develops into the Polygonum type of embryo sac, and synergids are of a wide-openly dichotomous filiform apparatus (Plate 2: 14, 18;Fig. 1:4,6). These are the characters reported here for the first time. Due to the lack of embryological da-ta for the related families, it is impossible to make a systematic comparison of embryology.     

Complex genome rearrangements reveal evolutionary dynamics of pericentromeric regions in the Triticeae.

Most pericentromeric regions of eukaryotic chromosomes are heterochromatic and are the most rapidly evolving regions of complex genomes. The closely related genomes within hexaploid wheat (Triticum aestivum L., 2n=6x=42, AABBDD), as well as in the related Triticeae taxa, share large conserved chromosome segments and provide a good model for the study of the evolution of pericentromeric regions. Here we report on the comparative analysis of pericentric inversions in the Triticeae, including Triticum aestivum, Aegilops speltoides, Ae. longissima, Ae. searsii, Hordeum vulgare, Secale cereale, and Agropyron elongatum. Previously, 4 pericentric inversions were identified in the hexaploid wheat cultivar 'Chinese Spring' ('CS') involving chromosomes 2B, 4A, 4B, and 5A. In the present study, 2 additional pericentric inversions were detected in chromosomes 3B and 6B of 'CS' wheat. Only the 3B inversion pre-existed in chromosome 3S, 3Sl, and 3Ss of Aegilops species of the Sitopsis section, the remaining inversions occurring after wheat polyploidization. The translocation T2BS/6BS previously reported in 'CS' was detected in the hexaploid variety 'Wichita' but not in other species of the Triticeae. It appears that the B genome is more prone to genome rearrangements than are the A and D genomes. Five different pericentric inversions were detected in rye chromosomes 3R and 4R, 4Sl of Ae. longissima, 4H of barley, and 6E of Ag. elongatum. This indicates that pericentric regions in the Triticeae, especially those of group 4 chromosomes, are undergoing rapid and recurrent rearrangements.     

Studies on Morphological Evolution,Classification and Distribution of the Triticeae in China

_{The classification and the relationships among the genera of Chinese Triticeae were studied based on morphological characters with reference to geographical distribution and habitat conditions. The spike of Triticeae might have been derived from a panicled inflorescence like that in the Bromeae through a racemose inflorescence like the one in the Brachypodieae. There might be three evolutionary lines in the tribe. 1. Pedicels of the panicled inflorescence have become short and bracts decreased in size, which has resulted in a panicled spike with indefinite spikelets or false solitary spikelets at each node of rachis. The middle ribes of both glumes and lemmas and rachilla are not in a single plane. 2. A simple spike with usual solitary spikelets at each node of rachis has been derived from the raceme. The middle ribe of both glumes and lemmas and rachilla are in a single plane. 3. A cymose spike with 3-spikelets at each node of rachis has evolved from the cymose panicle. The glume on the central spikelet is behind the lemma, while those on the lateral spikelets are on lateral sides of the lemmas. From what we have described above Triticeae may be divided into three subtribes: Elyminae, Triticinae and Hordeinae. Then according to the morphological characters of glume, lemma and other organs as well as the habitats and distribution, the native and introduced triticeous plants are classified into 13 genera (Leymus, Elymus, Roegneria, Elytrigia, Aegilops, Triticum, Agropyron, Eremopyrum, Secale, Haynaldia, Psathyrostachys, Hordeum and Hystrix) and their relationships are also discussed meanwhile.}     

山西产9种野生植物的染色体观察

本文报道了产于山西的4科5属9种野生植物的染色体观察结果。其中有2种植物作了核型分析,5种植物的染色体为首次报道。     

Molecular phylogeny revealed complex evolutionary process in Elymus species

Recent molecular phylogenetic studies on Elymus have added to our understanding of the origination of Elymus species. However, evolutionary dynamics and speciation of most species in Elymus are unclear. Molecular phylogeny has demonstrated that reticulate evolution has occurred extensively in the genus, as an example, the largest subunit of RNA polymerase II (rpb2) and phosphoenolpyruvate carboxylase (pepC) data revealed two versions of the St genome, St1 and St2contributing to speciation of E. caninus. Phylogenetic analyses of E. pendulinus uncovered additional genome-level complexity. Our data indicated that both chloroplast and nuclear gene introgression have occurred in the evolutionary process of E. pendulinus. Non-donor species genomes have been detected in severalElymus species, such as in allohexaploid E. repens (StStStStHH), a Taeniatherum-like (Ta genome in Triticeae) GBSSI sequence, Bromus- (Bromeae) and Panicum-like (Paniceae) ITS sequences have been detected. The chloroplast DNA data indicated that Pseudoroegneria is the maternal genome donor to Elymus species, but whether different Elymus species originated from different St donors remains an open question. The origin of the Y genome in Elymus is puzzling. It is clear that the Ygenome is distinct from the St genome, but unclear on the relationships of Y to other genomes in Triticeae. Introgressive hybridization may be an important factor complicating the evolutionary history of the species in Elymus. The extent of introgression and its role in creating diversity in Elymus species should be the objective of further investigations.     

Genome analysis of Elytrigia pycnantha and Thinopyrum junceiforme and of their putative natural hybrid using the GISH technique.

Genomic in situ hybridization (GISH), using genomic DNA probes from Thinopyrum elongatum (Host) D.R. Dewey (E genome, 2n = 14), Th. bessarabicum (Savul. & Rayss) A. Love (J genome, 2n = 14), Pseudoroegneria stipifolia (Czern. ex Nevski) Love (S genome, 2n = 14), and Agropyron cristatum (L.) Gaertner (P genome, 2n = 14), was used to characterize the genome constitution of the polyploid species Elytrigia pycnantha (2n = 6x = 42) and Thinopyrum junceiforme (2n = 4x = 28) and of one hybrid population (2n = 5x = 35). GISH results indicated that E. pycnantha contains S, E, and P genomes; the first of these was closely related to the S genome of Ps. stipifolia, the second was closely related to to the E genome of Th. elongatum, and the third was specifically related to A. cristatum. The E and P genomes included 2 and 10 chromosomes, respectively, with S genome DNA sequences in the centromeric region. GISH analysis of Th. junceiforme showed the presence of two sets of the E genome, except for fewer than 10 chromosomes for which the telomeric regions were not identified. Based on these results, the genome formula SSPsPsEsEs is proposed for E. pycnantha and that of EEEE is proposed for Th. junceiforme. The genomic constitution of the pentaploid hybrid comprised one S genome (seven chromosomes), one P genome (seven chromosomes), and three E genomes (21 chromosomes). The E and P genomes both included mosaic chromosomes (chromosomes 1 and 5, respectively) with the centromere region closely related to S-genome DNA. On the basis of these data, the genome formula SPSESEE is suggested for this hybrid and it is also suggested that the two species E. pycnantha and Th. junceiforme are the parents of the pentaploid hybrid.     

Further cytogenetical studies on diploid and polyploid species of Eryngium L. (Saniculoideae, Apiaceae) from Argentina

Meiotic studies are carried out in 7 species of Eryngium L. (Saniculoideae, Apiaceae), belonging to both sections Foetida and Panniculata. The chromosome number of E. dorae Norm. (n=8) (Foetida) is reported for the first time, while the gametic chromosome number of E. nudicaule Lam. (n=7) (Foetida) and E. eburneum Decne. (n=8), E. horridum Malme (n=8), E. megapotamicum Malme (n=16), E. mesopotamicum Pedersen (n=24), and E. pandanifolium Cham. et Schlechtd. (n=24) (all belonging to Panniculata) is confirmed in several natural populations. Whereas in section Foetida all species are diploids and two basic chromosome numbers are present (x=8 and x=7), in section Panniculata all species are x=8 but there are three different ploidy levels (diploid, tetraploid, hexaploid). This study reveals that meiosis in all species is normal, with regular bivalent formation in all studied cells. Furthermore, the pollen stainability is above 80% in all cases. These data, together with the previous karyotype analyses, will contribute to the clarification of the relationships between members of both sections, where different mechanisms of speciation have been postulated.     

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.     

新疆、青海和四川等地区小麦族植物的细胞学观察

本文对采集于新疆、青海和四川等地的小麦族(Triticeae Dumortier)10属、52种、370份种子材料进行了细胞学观察。该地区小麦族各属种的染色体数目变化范围是从2n=14到2n=84,前者主要存在于大麦属(Hordeum)、新麦草属(Psathyrostachys)和黑麦属(Secale),而后者全部集中于赖草属(Leymus)。其中染色体数目为2n=28和2n=42的类型出现的频率很高,大多存在于鹅观草属(Roegneria)和披碱草属(Elymus)。除个别种内存在不同倍性的细胞型外,绝大多数种的染色体数目非常稳定。在所有的样本中均没有观察到具非整倍体和B-染色体的材料。     

Biosystematics and evolutionary relationships of perennial Triticeae species revealed by genomic analyses

Understanding the classification and biosystematics of species in Triticeae Dumort., an economically important tribe in the grass family (Poaceae), is not an easy task, particularly for some perennial species. Does genomic analysis facilitate the understanding of evolutionary relationships of these Triticeae species? We reviewed literature published after 1984 to address questions concerning: (1) genome relationships among the monogenomic diploid species; (2) progenitors of the unknown Y genome in Elymus polyploids, X genome in Thinopyrum intermedium, and Xm genome in Leymus; and (3) genome constitutions of some perennial Triticeae species that were unknown or misidentified. A majority of publications have substantiated the close affinity of the E^b and E^e genomes in Th. bessarabicumand Th. elongatum, supporting the use of a common basic genome symbol. The E genome is close to the St genome of Pseudoroegneria and ABD genomes ofTriticum/Aegilops complex, providing an explanation for transferring genes from the E to ABD genomes with relative ease. Although the solid proof is still lacking, theW, P, and especially Xp genomes are possible origins for the Y genome of polyploid Elymus. The absence of the E genome and the allopolyploidy nature of tetraploidLeymus species have been unequivocally confirmed by both cytogenetic and molecular studies. However, the donor of the Xm genomes of Leymus was only speculated to be related to the P genome of Agropyron and F genome of Eremopyrum. Intermediate wheatgrass (Th. intermedium) has been extensively studied. The presence of the St (as the previously designated X) genome in Th. intermedium is now unequivocal. Its two more closely related E₁ and E₂ genomes are shown to be older versions of the E genome rather than the current E^b and E^e genomes. Speciation of Th. intermedium was similar to that of Triticum aestivum, in which the J^s/E^s(like B) genomes had the greatest differentiation from the current J (E^b) genome owning to repetitive sequences of the V genome, whereas its St (like D) had the least differentiation from the current St genome. Species with unknown or misidentified genomes have been correctly designated, including those with the ESt, StP, StPY,StWY, EStP, HW, StYHW, and NsXm genomes. Some of those species have been transferred to and renamed in appropriate genera.     

新疆独尾草属植物的核型分析

对新疆独尾草属（Eremurus）植物的核型进行了研究。核型公式如下：阿尔泰独尾草[E.altaicus（Pall.）Stev.]2n=2x=14=4m＋8sm＋2st;异翅独尾草[E.anisopterus（Kar.et Kit）Regel]2n=4x=28=4m＋4sm＋20st;粗柄独尾草[E.inderiensis （M.Bieb）Regel]2n=2x=14=10sm＋4st,首次发现古尔班通古特沙漠南缘所产的异翅独尾草2n=4x=28,与前人报道其为二倍体2n=2x=14的结果不一致。     

Genetic Relationships Among Five Basic Genomes St, E, A, B and D in Triticeae Revealed by Genomic Southern and in situ Hybridization

The St and E are two important basic genomes in the perennial tribe Triticeae （Poaceae）. They exist in many perennial species and are very closely related to the A, B and D genomes of bread wheat （Triticum aestivum L.）. Genomic Southern hybridization and genomic in situ hybridization （GISH） were used to analyze the genomic relationships between the two genomes （St and E） and the three basic genomes （A, B and D） of T. aestivum. The semi-quantitative analysis of the Southern hybridization suggested that both St and E genomes are most closely related to the D genome, then the A genome, and relatively distant to the B genome. GISH analysis using St and E genomic DNA as probes further confirmed the conclusion. St and E are the two basic genomes of Thinopyrum ponticum （StStE^eE^bE^x） and Th. intermedium （StE^eE^b）, two perennial species successfully used in wheat improvement. Therefore, this paper provides a possible answer as to why most of the spontaneous wheat-Thinopyrum translocations and substitutions usually happen in the D genome, some in the A genome and rarely in the B genome. This would develop further use of alien species for wheat improvement, especially those containing St or E in their genome components.     

Production,morphology, and cytogenetics of Triticum aestivum (L.) Thell × Elymus scabrus (R. Br.) Love intergeneric hybrids obtained by in ovulo embryo culture

Summary Intergeneric hybrids were produced between common wheat, Triticum aestivum (2n=6x=42, AABBDD), and an apomictic Triticeae species, Elymus scabrus (syn. Agropyron scabrum) (2n=6x=42, HHSSSS), the first successful report of this cross. Nine tiny, underdeveloped, and structureless embryos were obtained in vitro only by in ovulo embryo culture at 4 days after pollination, which gave rise to five mature hybrid plants. All the hybrid plants were vigorous and possessed a phenotype intermediate to the two parents. There were 2n=6x=42 (ABDHSS) somatic chromosomes in the hybrids. There was little or no homology between the parental genomes, as shown by an overall meiotic chromosome association of 32.83 I + 4.08 rod II + 0.21 ring II + 0.18 III + 0.02 IV. The hybrids were completely sterile and so far backcrosses to wheat parent have not been successful. Alternate approaches to induce gene transfer(s) from E. scabrus to wheat are being attempted.Agriculture Canada Contribution No. 398.