Genome constitutions of Thinopyrum curvifolium, T. scirpeum, T. distichum, and T. junceum (Triticeae: Gramineae).

The objective of this study is to elucidate genome constitutions of Thinopyrum curvifolium (Lange) D.R. Dewey, T. scirpeum (K. Presl) D.R. Dewey, T. distichum (Thunb.) A. L?ve, and T. junceum (L.) A. L?ve. Hybrids of T. sartorii (Boiss. &Heidr.) A. L?ve with T. scirpeum and T. junceum, as well as the hybrid between T. curvifolium and Pseudoroegneria geniculata ssp. scythica (Nevski) A. L?ve, were made and chromosome pairing at metaphase I was studied. The karyotype analyses of mitotic cells stained by aceto-orcein were conducted for both hybrids and the four target species. The Giemsa C-banding following acetocarmine staining was carried out for the above species and the triploid hybrid T. curvifolium x T. bessarabicum (Savul &Rayss) A. L?ve. Meiotic data indicate that all target species have two sets of the basic genome J, but they behave like true allopolyploids because of bivalentization. Karyotypes of T. curvifolium and its triploid hybrid with T. bessarabicum indicate that T. curvifolium contains two different versions of the Jb genome, designated as Jb3 and Jb4, rather than two Je genomes as previously believed. Thinopyrum scirpeum and T. elongatum (4x) have similar karyotypes. Both are segmental allotetraploids carrying two forms of the Je genome. Their genome formulae are Je2 Je3 and Je1 Je3, respectively. Thinopyrum distichum has a karyotype similar to T. junceiforme, which has the Jb2 Je2 genome formula. However, the two species differ in C-banding patterns, reflecting their geographical separation. Thinopyrum junceum is a hexaploid with two pairs of Jb2 genomes and one pair of the Je2 genome, and it has a C-banding pattern similar to that of T. junceiforme, which has one pair each of the Jb2 and Je2 genomes.     

Genetic and genomic relationships in Leymus Hochst

Genetic and genomic relationships among three taxonomically related species of Leymus, northern European L. arenarius (octoploid, 2n = 56), northern American/Pacific L. mollis (tetraploid, 2n = 28) and central Eurasian L. racemosus (tetraploid, 2n = 28), were examined using molecular and cytogenetic methods. The amplified fragment length polymorphism (AFLP) analysis clearly differentiated Icelandic populations of L. arenarius from Alaskan populations of L. mollis. The former group is more genetically homogeneous than the latter. Leymus arenarius in Iceland has a common gene pool and a relatively recent origin. The Alaskan L. mollis, on the other hand, is probably a glacial survival that has accumulated high level of genetic variation and has differentiated into subspecies. Analysis of the 18S-26S ribosomal genes, by restriction fragment length polymorphism (RFLP) and fluorescence in situ hybridization (FISH), revealed a very close relationship between the octoploid northern European L. arenarius and the tetraploid Eurasian L. racemosus, such that the former could have originated from the latter, probably via interspecific hybridization. Leymus-specific DNA sequences were isolated and used for analyzing genetic relatedness among five Leymus species and four Psathyrostachys species. The RFLP analysis of retrotransposon sequence pLm44 and ribosomal clone pTa71 clearly revealed a close relationship between these two genera, i.e. higher variation was found within genera than between them. The results support the previous notion that Leymus is autopolyploid having all genomes being designated Ns as in Psathyrostachys, but a major taxonomic revision of this group would require analysis of more species.     

Phylogenetic Analysis of Leymus (Poaceae: Triticeae) Inferred from Nuclear rDNA ITS Sequences

To investigate the phylogenetic relationships of polyploid Leymus (Poaceae: Triticeae), sequences of the nuclear rDNA internal transcribed spacer region (ITS) were analyzed for 34 Leymus accessions representing 25 species, together with three Psathyrostachys species (Ns genome), two Pseudoroegneria (St genome) species, Lophopyrum elongatum (E(e) genome), and Thinopyrum bessarabicum (E(b) genome). The phylogenetic analyses (maximum likelihood and Bayesian inference) supported two major clades, one including 21 Leymus species and three Psathyrostachys species, the other with nine Leymus species and four diploid species. The ITS RNA secondary structure of the Leymus species was compared with that of their putative diploid donor. It is suggested that (1) the species from the same areas or neighboring geographic regions are closely related to each other; (2) L. coreanus, L. duthiei, L. duthiei var. longearistatus, and L. komarovii are closely related to other Leymus species, and it is reasonable to transfer these species from the genus Hystrix to Leymus; (3) the ITS sequences of Leymus are evolutionarily distinct; (4) the different Leymus species and different distribution of a species derived their Ns genome from different Psathyrostachys species; and (5) there is a close relationship among Leymus, Pseudoroegneria, Lophopyrum, and Thinopyrum, but it is difficult to presume that the St, E(e), and E(b) genome may be the Xm genome donor of the Leymus species.     

Isolation, characterization, and analysis of Leymus-specific DNA sequences.

Genomic Southern hybridization using labeled total genomic DNA of Leymus mollis as probe showed intense hybridization signals on all restriction enzyme digested DNA from five species of Leymus Hochst., and four species of Psathyrostachys Nevski. Experiments using the same L. mollis probe, but with unlabeled blocking DNA from Psathyrostachys, showed no hybridization at all. These two genera evidently had the same genomic content. Southern hybridization without blocking allowed identification of DNA fragments abundant in Leymus and Psathyrostachys. Fragments potentially specific to Leymus were cloned. Five repetitive DNA clones from L. mollis and L. arenarius were characterized: pLmIs1, pLmIs44, pLmIs51, pLmIs53, and pLaIs56. These clones hybridized to both Leymus and Psathyrostachys on Southern blots - no clone hybridized to only one of these genera. Both Southern blot and fluorescence in situ hybridization (FISH) experiments showed that all the clones contained dispersed repetitive sequences. They painted all and whole chromosomes uniformly except at centromeres, telomeres, and nucleolar organiser regions. Three of these clones, i.e., pLmIs1, pLmIs44, and pLmIs53, were essentially specific to Leymus and Psathyrostachys - little or no hybridization was detected in other genera such as Triticum, Hordeum, Thinopyrum, or Elymus. Sequence analysis further revealed that the clones were part of retroelements. In particular, the clone pLmIs44 produced hybridization profiles suitable for analysis of genetic relatedness among species. The present study shows that Leymus and Psathyrostachys share the same basic genome, Ns, and therefore provides strong evidence for combining these two genera.     

猬草及其近缘属植物单拷贝核Pgk1基因序列的系统发育分析

文章对禾本科小麦族猬草属及其近缘属Thinopyrum(Eb)、Lophopyrum(Ee)、拟鹅观草属(St)、新麦草属(Ns)、大麦属(H)、赖草属(NsXm)和披碱草属(StH)植物共23个类群的单拷贝核Pgk1基因序列进行系统发育分析, 探讨猬草属及其近缘属植物的系统发育关系。序列分析发现Pgk1基因序列在L. arenarius和Psa. juncea中有81 bp的Stowaway家族DNA转座元件插入, 而在Hy. duthiei、Hy. duthiei ssp. longearistata和L. akmolinensis中有29 bp Copia家族的反转录转座元件插入。最大似然和贝叶斯推断进行的系统发育分析表明: (1)猬草属模式种Hy. patula与披碱草属、拟鹅观草属和大麦属具有密切的亲缘关系; (2)猬草属的其他物种Hy. duthiei、Hy. duthiei ssp. longearistata、Hy. coreana和Hy. komarovii与新麦草属和赖草属植物亲缘关系密切。研究结果支持将Hy. patula从猬草属组合到披碱草属中, 而Hy. duthiei、Hy. duthiei ssp. longearistata、Hy. coreana和Hy. komarovii应组合到赖草属中。     

Genome analysis of Elytrigia caespitosa, Lophopyrum nodosum, Pseudoroegneria geniculata ssp. scythica, and Thinopyrum intermedium (Triticeae: Gramineae).

To elucidate the genome constitutions of the tetraploid (2n = 4x = 28) species Elytrigia caespitosa, Lophopyrum nodosum, and Pseudoroegneria geniculata ssp. scythica and the hexaploid (2n = 6x = 42) Thinopyrum intermedium, meiotic pairing was studied in these species as well as 10 hybrids. Karyotype analysis with aceto-orcein stained root-tip cells was performed for the four species and the hybrids of T. bessarabicum with E. caespitosa, P. geniculata ssp. scythica, and T. intermedium. Karyotype analysis by Giemsa C-banding was carried out with the three tetraploid species and the two triploid hybrids involving T. bessarabicum. The species behaved as strict allopolyploids. All hybrids were male sterile with few stainable pollen grains. It is concluded from the results that the three tetraploid species have the genome formula JeJeSS and T. intermedium has the formula JeJeJeJeSS. The chromosomes of the Je and S genomes in these species had C-banding patterns differing from each other and from those of the extant diploid species. Based on these findings, the four species investigated should be placed in the same genus or the same section of a genus. However, new combinations are not proposed at this time pending future taxonomic investigation of the genome constitution of Elytrigia repens (L.) Nevski.     

Phylogeny of Hystrix and related genera (Poaceae: Triticeae) based on nuclear rDNA ITS sequences

The taxonomic status of Hystrix and phylogenetic relationships among Hystrix and its related genera of Pseudoroegneria (St), Hordeum (H), Psathyrostachys (Ns), Elymus (StH), Leymus (NsXm), Thinopyrum bessarabicum (E(b)) and Lophopyrum elongatum (E(e)) were estimated from sequences of the internal transcribed spacer (ITS) region of nuclear ribosomal DNA. The type species of Hystrix, H. patula, clustered with species of Pseudoroegneria, Hordeum, Elymus, Th. bessarabicum and Lo. elongatum, while H. duthiei ssp. duthiei, H. duthiei ssp. longearistata, H. coreana and H. komarovii were grouped with Psathyrostachys and Leymus species. The results indicate that: (i) H. patula is distantly related to other species of Hystrix, but is closely related to Elymus species; (ii) H. duthiei ssp. duthiei, H. duthiei ssp. longearistata, H. coreana and H. komarovii have a close affinity with Psathyrostachys and Leymus species, and H. komarovii might contain the NsXm genome of Leymus; and (iii) the St, H and Ns genomes in Hystrix originate from Pseudoroegneria, Hordeum and Psathyrostachys, respectively, while the Xm in Hystrix and Leymus has a complex relationship with the E or St genomes. According to the genomic system of classification in Tiritceae, it is reasonable to treat Hystrix patula as Elymus hystrix L, and the other species of Hystrix as species of a section of Leymus, Leymus Sect. Hystrix.     

Genetic diversity and phylogeny in Hystrix (Poaceae, Triticeae) and related genera inferred from Giemsa C-banded karyotypes

The phylogenetic relationships of 15 taxa from Hystrix and the related genera Leymus (NsXm), Elymus (StH), Pseudoroegneria (St), Hordeum (H), Psathyrostachys (Ns), and Thinopyrum (E) were examined by using the Giemsa C-banded karyotype. The Hy. patula C-banding pattern was similar to those of Elymus species, whereas C-banding patterns of the other Hystrix species were similar to those of Leymus species. The results suggest high genetic diversity within Hystrix, and support treating Hy. patula as E. hystrix L., and transferring Hy. coreana, Hy. duthiei ssp. duthiei and Hy. duthiei ssp. longearistata to the genus Leymus. On comparing C-banding patterns of Elymus species with their diploid ancestors (Pseudoroegneria and Hordeum), there are indications that certain chromosomal re-arrangements had previously occurred in the St and H genomes. Furthermore, a comparison of the C-banding patterns of the Hystrix and Leymus species with the potential diploid progenitors (Psathyrostachys and Thinopyrum) suggests that Hy. coreana and some Leymus species are closely related to the Ns genome of Psathyrostachys, whereas Hy. duthiei ssp. duthiei, Hy. duthiei ssp. longearistata and some of the Leymus species have a close relationship with the E genome. The results suggest a multiple origin of the polyploid genera Hystrix and Leymus.     

Genomic and genetic relationships among species of Leymus (Poaceae: Triticeae) inferred from 18S-26S ribosomal genes

The 18S-26S ribosomal genes in three closely related species of Leymus (Poaceae: Triticeae) were examined using fluorescence in situ hybridization (FISH) and restriction fragment length polymorphism (RFLP). Both approaches revealed a close relationship between L. arenarius (8x = 56, northern European) and L. racemosus (4x = 28, central Eurasian), whereas L. mollis (4x = 28, northern American/Pacific) was distinct. Each species had three homologous pairs of major rDNA loci: a1, a2, and a3 for L. arenarius; m1, m2, and m3 for L. mollis; and r1, r2, and r3 for L. racemosus. Leymus arenarius had in addition three minor loci, a4, a5, and a6. The major loci of L. arenarius and L. racemosus were identical, indicating that the former species could have originated from the latter, via interspecific hybridization and/or polyploidy. The rDNA-RFLPs further indicated relationships of these species to other species of Leymus (L. karellini, 8x = 56 and L. angustus, 12x = 84) and Psathyrostachys (P. fragilis, P. huashanica, P. juncea, and P. lanuginosa, which are all diploids). A phenogram constructed from 20 BamHI, EcoRI, and DraI rDNA fragments revealed closer relationship between the two genera, Leymus and Psathyrostachys, than that among species within a genus.     

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.     

Identification of the parental chromosomes of the wheat-alien amphiploid Agrotana by genomic in situ hybridization.

Labelled total genomic DNA from four alien species, Thinopyrum ponticum (Host) Beauv. (2n = 70, genomes J1J1J1J2J2), Th. bessarabicum (Savul. &Rayss) Love (2n = 14, genome J), Th. elongatum (Host) Beauv. (2n = 14, genome E), and Haynaldia villosa (L.) Schur. (2n = 14, genome V), were used as probes in combination with blocking wheat DNA for in situ hybridization of the chromosomes of Agrotana, a wheat-alien hybrid (2n = 56) of unknown origin. The results showed that genomic DNA probes from Th. ponticum and Th. bessarabicum both clearly revealed 16 alien and 40 wheat chromosomes in Agrotana, indicating that the J genome present in these two species has a high degree of homology with the alien chromosomes in Agrotana. Biotinylated genomic DNA probe from Th. elongatum identified 10 chromosomes from Agrotana, while some regions of six other chromosomes yielded a weak or no signal. The probe from H. villosa produced no differential labelling of the chromosomes of Agrotana. The genomic formula of Agrotana was designated as AABBDDJJ. We suggest that the alien parent donor species of Agrotana is Th. ponticum rather than Th. bessarabicum. Genomic relationships of the three Thinopyrum species are discussed in relation to the distribution of GISH signals in the chromosomes of Agrotana.     

Coexistence and hybridization of Leymus mollis and L. arenarius in Greenland, and demarcation of the species by endospermal prolamins, leymins

The occurrence of Leymus arenarius as a constituent of the Greenlandic flora is confirmed. The different patterns of hairiness of the rachis segments characteristic of L. mollis and L. arenarius , respectively, were found to correlate with endospermal protein patterns fractionated with SDS-PAGE (sodium dodecylsulphate, polyacrylamide gel electrophoresis). The characteristics of its protein patterns were substantiated by a transect study of L. arenarius through Fennoscandia. The northern populations of L. mollis and L. arenarius displayed no or relatively little variation in endospermal protein patterns within plants or sites. This may reflect a prevailing self-pollination, founder effect, clonal spreading of stands and selection or any combination thereof. The key proteins of 120 to 140 kg mole^-1 demarcating L. mollis and L. arenarius were shown to be soluble in 50% isopropanol at pH 8.0, and hence to be prolamins, which were also found to carry glycosylation. The spontaneous, sterile hybrids of these species in Greenland have an intermediary morphology. A hybrid clone may form relatively wide stands in South and Southwest Greenland.     

Characterization and distribution of an interspersed repeated nucleotide sequence from Lophopyrum elongatum and mapping of a segregation-distortion factor with it

Repeated nucleotide sequence pLeUCD2 cloned from Lophopyrum elongatum is highly abundant in the genomes of diploid and polyploid wheatgrass species of genera Lophopyrum, Thinopyrum, Pseudoroegneria, Agropyron, Elymus, Elytrigia, and Pascopyrum but undetectable by Southern blot hybridization in Triticum and representative species of Dasypyrum, Hordeum, Psathyrostachys, Secale, Taeniatherum, Heteranthelium, and Leymus in the tribe Triticeae. The DNA fragment inserted in pLeUCD2 is 277 bp long and AT rich (65%), and contains numerous inverted and palindromic repeats. In situ DNA hybridization substantiated a previous hypothesis that the sequence is interspersed in the wheatgrass genomes. Heterogeneity and clustering of like repeats of the pLeUCD2 family along wheatgrass chromosomes was used to map a segregation-distortion factor, designated Sd-1, proximal to the Lr19 locus in recombinant chromosomes of L. ponticum chromosome 7Ag and wheat chromosome 7D.     

Molecular variation in Leymus species and populations

Icelandic populations of European lymegrass [Leymus arenarius (L.) Hochst.] were examined using amplified fragment length polymorphism (AFLP) and restriction fragment length polymorphism (RFLP) of the major ribosomal genes (18S-5.8S-26S rDNA), in comparison with Alaskan populations of its closely related species L. mollis (Trin.) Pilger. The AFLP profiles emerged as two distinct entities, clearly separating the two species, and based on species-specific bands it was simple to distinguish these two morphologically similar species. The rDNA-RFLPs also differentiated the species. Within species, the Icelandic L. arenarius was more homogeneous than the Alaskan L. mollis, and its variation was dispersed over geographically different populations, suggesting a common gene pool. The variation among the Alaskan L. mollis was more extensive and its interrupted pattern may be the result of gene introgression at subspecies level. Within a 40-year-old population of L. mollis established in Iceland from Alaskan material, the molecular profiles separated old and new genotypes. Both AFLP and rDNA revealed the new genotypes to be extremely similar. This rapid change in allele frequency is thought to be the result of adaptation to a new environment.     

Salt tolerance in the Triticeae: K/Na discrimination in some perennial wheatgrasses and their amphiploids with wheat

Perennial wheatgrasses of the genera Thinopynim, Leymus andPsathyrostachys were subjected to high and low salinities todetermine whether the enhanced K/Na discrimination trait wasexpressed in these members of the Triticeae. At low salinities,where the enhanced K/Na discrimination trait is most evident,the leaf cation concentrations were consistent with the operationof the trait. At higher salinities sodium exclusion and potassiumretention were greater than in tetraploid wheat. Experimentswith hexaploid or octoploid wheat x Thinopynim amphidiploidsshowed that the presence of the D genome of wheat greatly affectedcation discrimination, and pointed to the absence of enhancedK/Na discrimination in the perennial wheatgrasses. Results obtainedwith Thinopynim elongatum addition lines, and the uptake of²²Na into 'low-salt' amphiploid seedlings, are consistent withthe absence of the trait in Thinopynim. Key words: Wheat, Thinopyrum, Leymus, amphiploids, salt     

RAPD method for the identification of intergeneric asymmetric somatic hybrid plants of wheat]

Randomly amplified polymorphic DNA (RAPD) method was used to identify the hybrid nature of three kinds of intergeneric asymmetric somatic hybrid plants of wheat: wheat (Triticum aestivum) + Haynaldia villosa, Wheat + Leymus chinensis and wheat + Agropyron elongatum. It was shown from the electrophoresis profiles that the genome of somatic hybrid plants contained specific section genome of both parents after DNA amplification with arbitrary primers. A specific RAPD product (DNA fragment of 0.77 kbp) of A. elongatum generated with primer OPJ-12 was isolated, purified, labeled and used as a probe. Southern blot from OPJ-12 primer-generated specific section genome of the hybrid (T. aestivum + A. elongatum) hybridized to this probe (0.77 kbp) proved that they are homologous in nature. This paper also discussed the advantage of RAPD method in identification of hybrid plants, especially asymmetric somatic hybrids.     

Phylogenetic relationships in Leymus (Triticeae; Poaceae): Evidence from chloroplast trnH-psbA and mitochondrial coxII intron sequences

Leymus Hochst. is a polyploid genus with a diverse array of morphology, cytology, ecology, and distribution in Triticeae. To investigate the phylogenetic relationships and maternal genome donor of polyploid Leymus, the chloroplast trnH-psbA region and mitochondrial coxII intron sequences of 33 Leymus taxa were analyzed with those of 36 diploid perennial species representing 19 basic genomes in Triticeae. The results showed that reticulate evolution occurred in Leymus species, with the cytoplasmic lineage of Leymus contributed by different progenitors. Interspecific relationships of Leymus were also elucidated on the basis of orthologous comparison. Our data suggested that: (i) due to incomplete lineage sorting and/or difference in the pattern of chloroplast and mitochondrial inheritance, the genealogical conflict between the two genealogical patterns suggest the contribution of Psathyrostachys Nevski, Agropyron J. Gaertn, Eremopyrum (Ledeb.) Jaub. & Spach, Pseudoroegneria (Nevski) Á. Löve, Thinopyrum Á. Löve, and Lophopyrum (Host) Á. Löve to the cytoplasmic lineage of Leymus; (ii) there is a close relationship among Leymus species from the same area or neighboring geographic regions; (iii) L. coreanus (Honda) K. B. Jensen & R. R.-C. Wang, L. duthiei (Stapf) Y. H. Zhou & H. Q. Zhang ex C. Yen, J. L. Yang & B. R. Baum, L. duthiei var. longearistatus (Hack.) Y. H. Zhou & H. Q. Zhang ex C. Yen, J. L. Yang & B. R. Baum, and L. komarovii(Roshev.) C. Yen, J. L. Yang & B. R. Baum are closely related to other Leymus species, and it is reasonable to transfer these species from the genus Hystrix Moench to Leymus; (iv) Leymus species from North America are closely related to L. coreanus from the Russian Far East and L. komarovii from northeast China but are evolutionarily distinct from Leymus species from Central Asia and the Qinghai–Tibet Plateau. The occurrence of multiple origin and introgression could account for the rich diversity and ecological adaptation of Leymus species.     

Reciprocal hybridization at different times between Senecio flavus and Senecio glaucus gave rise to two polyploid species in north Africa and south-west Asia

The analysis of hybrid plant taxa using molecular methods has considerably extended understanding of possible pathways of hybrid evolution. Here, we investigated the origin of the tetraploid Senecio mohavensis ssp. breviflorus and the hexaploid Senecio hoggariensis by sequencing of nuclear and chloroplast DNA, and by analysis of the distribution of taxon-specific amplified fragment length polymorphism (AFLP) fragments. Both taxa originated from hybridization between the diploid Senecio flavus and Senecio glaucus. Whereas S. glaucus was the female parent in the origin of S. mohavensis ssp. breviflorus, S. flavus was the female parent in the origin of S. hoggariensis. The distribution of AFLP fragments suggests that S. hoggariensis is an allohexaploid species with two diploid genomes of S. glaucus and one diploid genome of S. flavus. The high frequency of S. flavus-specific fragments in S. mohavensis ssp. breviflorus is explained either as the result of introgression between a primary hybrid and S. flavus or as the result of intergenomic recombination in a primary hybrid. These two alternative processes cannot easily be distinguished.     

Triploidy in Equisetum subgenus Hippochaete (Equisetaceae, Pteridophyta)

BACKGROUND AND AIMS: The genus Equisetum is cytologically uniform, having a base chromosome number of x = 108. All previously known species and hybrids that have been counted represent diploids with a sporophytic chromosome number of 2n = 216. Biosystematic studies on Equisetum subgenus Hippochaete revealed evidence that triploids occur in nature. The objective of this study was to confirm that triploid plants exist in the natural environment. METHODS: Flow cytometry was used to establish nuclear DNA values and cytological investigations of meiosis were carried out to obtain information on chromosome number and pairing behaviour. KEY RESULTS: Triploidy exists in three morphologically different hybrid taxa. Two of these are morphologically intermediate between a primary diploid hybrid and a parent, while the third apparently combines genomes from all three Central European Hippochaete species. Nuclear 1C DNA values for the four European Hippochaete species range from 21.4-31.6 pg. For the hybrids, the 1C DNA values not only occupy the same range as the species, but their total DNA amounts agree closely with values predicted by adding the 1C DNA values of each parental genome. Chromosome counts confirm diploidy in the species E. hyemale and E. variegatum and in the hybrid E. xtrachyodon (= E. hyemale x E. variegatum). For the triploids (2n approximately 324), cytological information is presented for the first time. CONCLUSIONS: Triploid taxa may have originated by backcrossing or by crossing of a diploid hybrid with an unrelated diploid species. As tetraploid plants are unknown, these crossings probably involve diploid gametophytes that developed from unreduced diplospores. By repeated crossing events or backcrossing, reticulate evolution patterns arise that are similar to those known for a number of ferns and fern allies.     

长穗偃麦草中E和E1基因组高分子量麦谷蛋白基因启动子部分序列的进化分析

通过PCR克隆的方法,获得了分别来自二倍体长穗偃麦草的E基因组和四倍体长穗偃麦草的E_1基因组的4个高分子量麦谷蛋白亚基(HMW-GS)基因启动子的部分序列。序列分析表明,它们之间的同源性较高,两个x型亚基启动子序列之间只有1个碱基的差异,而两个y型亚基启动子序列完全相同,x和y型亚基启动子序列之间的长度和部分碱基位点都有差异。推测四倍体长穗偃麦草中的E_1基因组可能起源于二倍体的E基因组。与来自小麦族的A、B、D和G基因组部分亚基基因的启动子序列比较表明,小麦族的这一区域在进化上是相当保守的,不同基因组来源的序列同源性都在90％以上。经过对这些序列的聚类分析,表明长穗偃麦草的y型HMW-GS基因与其他亚基基因的进化关系较远,而x型亚基基因与一个来自小麦1B染色体的亚基基因关系最近。