六个不同类型荞麦花花粉粒形态的电镜观察比较研究

对二倍体甜荞长花柱类型 (ES2 s)、二倍体甜荞短花柱类型 (ES2 S)、四倍体甜荞长花柱类型 (ES4s)、四倍体甜荞短花柱类型 (ES4S)、四倍体有翅细野荞 (GR4HI)、四倍体无翅细野荞 (GR4HO)等 6个不同荞麦类型花的花粉形态学进行了电镜观察。结果表明 :这些荞麦类型的花粉粒都是椭圆形 ,都有网状纹饰、3孔沟等特征。二倍体甜荞染色体加倍后形成的四倍体甜荞 ,相对于二倍体甜荞而言 ,花粉粒显著增大和明显变圆。无论二倍体还是四倍体甜荞 ,其长花柱型花的花粉粒都比其对应短花柱型花的花粉粒要小。尽管四倍体甜荞和四倍体细野荞在染色体数目和倍性上一样 ,但是它们在花粉粒的大小上有显著差异。四倍体细野荞有翅类型和无翅类型的花粉粒大小和形态差异不显著     

A new cytotype of Jacobaea vulgaris (Asteraceae): frequency,morphology and origin

Jacobaea vulgaris subsp. vulgaris (syn. Senecio jacobaea subsp. jacobaea) constitutes an intricate polyploid complex distributed in Europe. Four cytotypes have been reported in this species, three with euploid (diploid, tetraploid and octoploid; 2n=20, 40 and 80) and one with aneuploid (2n=32) chromosome numbers. Here we report that the diploid chromosome number (2n=20) reported from Bulgaria is due to misidentification with Jacobaea aquatica. On the other hand, we have discovered a new, hexaploid (2n=6x=60) cytotype within J. vulgaris subsp. vulgaris using flow cytometry. The new cytotype occurs within four sympatric populations of otherwise tetraploid and octoploid plants in Pannonia (one locality in the eastern Czech Republic and two localities in southwestern Slovakia) and in Podillya (one locality in western Ukraine). The frequency of hexaploid individuals within 76 studied populations is very low (only 10 of 693 analysed plants), and hexaploids probably represent hybrids between tetraploid and octoploid plants. Three mixed populations with hexaploid plants were subjected to detailed morphological and pollen fertility analyses. Multivariate morphometric analysis reveals partial separation of tetraploid and octoploid plants, whereas hexaploid individuals are similar in morphology to octoploids. In comparison with tetraploids, octoploids and hexaploids exhibit slightly longer ray florets, involucral bracts and tubular florets and more hairy outer achenes. Hexaploid plants display larger pollen grains and lower pollen fertility compared to tetraploids and octoploids.     

Genome duplication and the evolution of conspecific pollen precedence

Conspecific pollen precedence can be a strong reproductive barrier between polyploid and diploid species, but the role of genome multiplication in the evolution of this barrier has not been investigated. Here, we examine the direct effect of genome duplication on the evolution of pollen siring success in tetraploid Chamerion angustifolium. To separate the effects of genome duplication from selection after duplication, we compared pollen siring success of synthesized tetraploids (neotetraploids) with that of naturally occurring tetraploids by applying 2x, 4x (neo or established) or 2x + 4x pollen to diploid and tetraploid flowers. Seed set increased in diploids and decreased in both types of tetraploids as the proportion of pollen from diploid plants increased. Based on offspring ploidy from mixed-ploidy pollinations, pollen of the maternal ploidy always sired the majority of offspring but was strongest in established tetraploids and weakest in neotetraploids. Pollen from established tetraploids had significantly higher siring rates than neotetraploids when deposited on diploid (4x(est) = 47.2%, 4x(neo) = 27.1%) and on tetraploid recipients (4x(est) = 91.9%, 4x(neo) = 56.0%). Siring success of established tetraploids exceeded that of neotetraploids despite having similar pollen production per anther and pollen diameter. Our results suggest that, while pollen precedence can arise in association with the duplication event, the strength of polyploid siring success evolves after the duplication event.     

Pollen competition as a unilateral reproductive barrier between sympatric diploid and tetraploid Chamerion angustifolium

Speciation requires the evolution of barriers to gene exchange between descendant and progenitor populations. Cryptic reproductive barriers in plants arise after pollination but before fertilization as a result of pollen competition and interactions between male gametophytes and female reproductive tissues. We tested for such gametic isolation between the polyploid Chamerion angustifolium and its diploid progenitor by conducting single (diploid or tetraploid) and mixed ploidy (1 : 1 diploid and tetraploid) pollinations on both cytotypes and inferring siring success from paternity analysis and pollen-tube counts. In mixed pollinations, polyploids sired most (79%) of their own seeds as well as those of diploids (61%) (correcting for triploid block, siring success was 70% and 83%, respectively). In single donor pollinations, pollen tubes from tetraploids were more numerous than those from diploids at four different positions in each style and for both diploid and tetraploid pollen recipients. The lack of a pollen donor x recipient interaction indicates that the tetraploid siring advantage is a result of pollen competition rather than pollen-pistil interactions. Such unilateral pollen precedence results in an asymmetrical pattern of isolation, with tetraploids experiencing less gene flow than diploids. It also enhances tetraploid establishment in sympatric populations, by maximizing tetraploid success and simultaneously diminishing that of diploids through the production of inviable triploid offspring.     

Morphology, distribution, and chromosome counts of two varieties of Hedychium villosum (Zingiberaceae)

Polyploidy is a major mechanism of adaptation and speciation in plants. Two varieties of Hedychium villosum, var. villosum and var. tenuiflorum, primarily differ in plant and flower size. Chromosome number suggests that var. tenuiflorum is diploid (2n = 34) and var. villosum is tetraploid (2n = 68). Although the flowers of the two varieties do not have any difference in floral shape, each can be easily distinguished morphologically because the shoots, leaves, and flowers of the tetraploid var. villosum are consistently larger than the diploid var. tenuiflorum. The two varieties each possess distinct geographic ranges and habitats, and no sympatric distribution has been found.The tetraploid var. villosum has a broader geographic distribution range and more diverse ecological habitats than the diploid var. tenuiflorum. The two varieties are also completely reproductively isolated due to the non-overlap of their flowering times. The two varieties should be recognized as two distinct species as they fulfill the requirements of various species concepts. Thus, we suggested that the tetraploid var. villosum should be kept as Hedychium villosum and the diploid var. tenuiflorum should be renewed to Hedychium tenuiflorum.     

Morphology,distribution,and chromosome counts of two varieties of Hedychium villosum(Zingiberaceae)

Polyploidy is a major mechanism of adaptation and speciation in plants.Two varieties of Hedychium villosum,var.villosum and var.tenuiflorum,primarily differ in plant and flower size.Chromosome number suggests that var.tenuiflorum is diploid(2n=34)and var.villosum is tetraploid(2n=68).Although the flowers of the two varieties do not have any difference in floral shape,each can be easily distinguished morphologically because the shoots,leaves,and flowers of the tetraploid var.villosum are consistently larger than the diploid var.tenuiflorum.The two varieties each possess distinct geographic ranges and habitats,and no sympatric distribution has been found.The tetraploid var.villosum has a broader geographic distribution range and more diverse ecological habitats than the diploid var.tenuiflorum.The two varieties are also completely reproductively isolated due to the non-overlap of their flowering times.The two varieties should be recognized as two distinct species as they fulfill the requirements of various species concepts.Thus,we suggested that the tetraploid var.villosum should be kept as Hedychium villosum and the diploid var.tenuiflorum should be renewed to Hedychium tenuiflorum.     

四倍体南瓜的诱变和特性研究

以中国南瓜种‘上海小闸黄狼’南瓜为试材,用秋水仙素为诱变剂,对‘黄狼’种子进行处理,得到了同源四倍体南瓜。在4种浓度处理4h后,发现获得同源四倍体的最佳浓度为0．2％和0．3％。变异材料染色体数加倍（2n=4x=80）。研究发现,与二倍体相比,四倍体单位面积叶面气孔数变少,叶色变深,叶裂变浅,叶尖变钝,叶变大,茎变粗,大量出现畸形雄花,果实变小,果形变直,果柄变短,单瓜种子平均数由340粒减为68粒,抗早衰性更强,脯氨酸含量和过氧化物歧化酶活性更高,丙二醛含量更低。四倍体南瓜伤流量明显比二倍体高,根系活力强于二倍体。     

Correlation between distyly and ploidy level in Damnacanthus (Rubiaceae)

Somatic chromosomes were observed in 661 individuals of 14 taxa, nine species and five varieties, of Damnacanthus (Rubiaceae). Chromosome numbers are reported for the first time for 13 taxa. Diploid (2n = 22) and tetraploid (2n = 44) counts were obtained. Distyly is reported for the first time for four species, D. angustifolius, D. henryi, D. labordei, and D. officinarum. A strong correlation exists between chromosome number and occurrence of distyly. Regardless of taxa in Damnacanthus, distylous populations are diploid, and monomorphic populations are tetraploid. Flowers of the monomorphic populations observed have a long style and short stamens with few exceptions. Polyploidization may have caused the breakdown of distylous to monomorphic flowers. In D. indicus, leaves from the tetraploid populations tend to be larger than those from the diploid populations. Populations of tetraploid D. indicus were distributed in more northern areas than those of the diploid. Three types of sympatric distribution were found for the varieties of D. indicus in Japan: diploid and tetraploid, two diploids, and two tetraploids. Based on the present chromosome number study, the taxonomy of the varieties of D. indicus should be revised.     

Gender polymorphism in flowers of Silene vulgaris (Moench) Garcke (Caryophyllaceae)

Silene vulgaris occurs in Israel as a stolonifcrous tetraploid which forms mostly sexually polymorphic populations. Fully hermaphrodite and fully male-sterile plants are linked by a range of intermediate forms which are gynomonoecious and bear hermaphrodite, male-sterile, and partially male-sterile flowers. Male-sterile flowers differ from hermaphrodite ones by having a narrower calyx, a shorter corolla, a thicker style, and a considerably larger stigmatic area with longer papillae which are more markedly bulbous basally. In individual anthers of hermaphrodite flowers, diameters of stained pollen grains showed a continuous variation from 36 to 80 μm with a modal value of 56. About 30% of pollen was found to be non-stainable. Reduction in male fertility occurs at different levels, from small or inviable pollen grains through abortion of single anthers, abortion of entire androecia, and through different degrees of gynomonoecy to totally male-sterile individuals. The proportion of stainable pollen to ovules in hermaphrodite flowers alone was found to be less than 100. From the data presented, less than a third of all ovules produced in the different kinds of flower develop into seeds. This may reflect an inadequacy of pollen, which the local population of S. vulgaris can tolerate because of its stoloniferous habit.     

中国桔梗多倍体诱导与鉴定

在离体培养条件下,比较了不同浓度、不同处理时间的秋水仙素对中国桔梗（Platycodon grandiflorus A．CD）进行染色体加倍的诱导效果。结果表明：用含0．1％秋水仙素处理40h后诱导频率可达50％,诱导效果最佳。经秋水仙素诱导后形成的多倍体植株与原二倍体植株比较,在形态上,多倍植株叶片变宽变大,叶色变深,茎变粗且节距长,气孔增大而单位叶面积气孔数目减少。对变异植株进行细胞学研究发现,体细胞中期染色体数目为2n=4x=36,而原二倍体的染色体数目为2n=2x=18,基数x=9,因此,变异植株（2n=4x=36）为四倍体。前者的核型公式为2n=4x=14m＋20sm＋2st,核型属于2B;后者的核型公式为2n=2x=7m＋10sm＋1st,核型也属于2B。检测发现有少数个体有非整倍体变异。     

A simple and effective method for the micropropagation and in vitro induction of polyploidy and the effect on floral characteristics of the South African iris,Crocosmia aurea

South Africa is home to approximately 10% of the world's flora, many of which are endemic to the country. A large number of South African genera have been improved for horticultural use and many of these are economically important as cut flowers or ornamentals on international markets. The genus Crocosmia, an attractive member of the family Iridaceae, has potential both as an ornamental plant and for cut flower production, although market potential of the species may be increased by improving the size of the flowers and inflorescence. Polyploidy has been used as a tool in the improvement of ornamental plants and has led to the development of several improved ornamental species. This study established a micropropagation protocol for Crocosmia aurea, using seed as the source material. Tetraploidy was induced by treating seeds with colchicine. These seeds were subsequently germinated and multiplied in vitro using the established protocol. The resulting tetraploid plantlets were successfully hardened-off and used to study the effect of the induced tetraploidy on the plant characteristics. The tetraploid (4n) plants were found to have longer, wider leaves as well as longer inflorescence stems and fewer, but larger, flowers than their diploid (2n) counterparts. These polyploid selections have potential in the ornamental/floriculture trade.     

Changes in the endopolyploidy pattern of different tissues in diploid and tetraploid Phalaenopsis aphrodite subsp. formosana (Orchidaceae)

Endopolyploidy is frequently observed during development in plant species. Patterns of endopolyploidy are diverse in the various organs of different plant species. However, little is known about the role of endopolyploidization and its significance in orchids. This study was undertaken to determine the extent of endopolyploidy in different tissues of the diploid and tetraploid genotypes of Phalaenopsis aphrodite subsp. formosana and to examine the factors that contribute to increased ploidy levels. Endopolyploidy occurs in various tissues of diploid and tetraploid orchids, at different developmental stages and under different culture conditions, as determined by flow cytometry. In this study, different patterns of endopolyploidy were observed in parts of the protocorms, leaves, roots and flowers. Endopolyploidy was found in all tissues studied except the pollinia and the tetraploid ovaries. A higher degree of endopolyploidy was observed in mature tissues compared to young tissues, greenhouse-grown plants compared to in vitro plants and diploid plants compared to tetraploid plants. We discuss the relationships between endopolyploidization and several factors related to plant growth, as well as some practical considerations of these findings.     

Breeding system evolution in Tarasa (Malvaceae) and selection for reduced pollen grain size in the polyploid species

Polyploidy, primarily allopolyploidy, has played a major role throughout flowering plant evolution with an estimated 30-80% of all extant angiosperms carrying traces of ancient or recent polyploidy. One immediate and seemingly invariant phenotypic consequence of genome doubling is larger cell size in polyploids relative to their diploid progenitors. In plants, increases in pollen grain and guard cell sizes exemplify this rule and are often used as surrogate evidence for polyploidy. Tarasa (Malvaceae), a genus of 27 species primarily distributed in the high (>3000 m) Andes, has numerous independently generated tetraploid species, most of which have pollen grains smaller than their putative diploid parents. The tetraploids are also unusual because they are annual, rather than perennial, in habit. Data correlate these apparent anomalies to a change in the breeding system within the genus from xenogamy (outcrossing) in the diploid species to autogamy (inbreeding) in the tetraploids, leading to a convergence in reduced floral morphology. The harsh environment of the high-elevation Andean habitats in which all the tetraploid annuals are found is implicated as a critical factor in shaping the evolution of these unusual polyploids.     

In vitro induction of tetraploid plants from diploid Zizyphus jujuba Mill. cv. Zhanhua

Tetraploid plants of Zizyphus jujuba Mill. cv. Zhanhua were obtained with in vitro colchicine treatment. Shoot tips from in vitro-grown plants were treated with five different concentrations of colchicine (0.01, 0.03, 0.05, 0.1, 0.3%) in liquid MS medium (Murashige and Skoog 1962), and shaken (100 rpm) at 25 °C in darkness for 24, 48, 72 or 96 h, respectively. Tetraploids were obtained at a frequency of over 3% by using 0.05% colchicine (48 h, 72 h) and 0.1% colchicine (24 h, 48 h) treatment as determined by flow cytometry. Cytological and morphological evidence confirmed the results of flow cytometric analysis. The chromosome number of diploid plants was 24 and that of tetraploid plants was 48. The stomata sizes of tetraploid plants were significantly larger than those of diploid plants, while the frequency of stomata were reduced significantly. Similarly, the chloroplast number of guard cells of tetraploid plants increased significantly. The selected tetraploid plants were grafted onto mature trees of Z. jujuba Mill. cv. Zhanhua in the field, resulted in thicker stems, rounder and succulent leaves, larger flowers and a delay in florescence time (3–4 days later) than diploid plants.     

Pollen anomaly - a clue to natural hybridity in Argemone (Papaveraceae)

Studies in the pollen morphology of Argemone mexicana (a diploid species, 2n=28), A. ochroleuca (a tetraploid, 2n=56), the wild populations of which growing side by side, and of a triploid (2n=42) observed in the population of A. ochroleuca, have been carried out using light and scanning electron microscope. Pollen grains in both the species are 3 (-4)-zonocolpate with reticulate exine surface, and are distinguishable only on the basis of pollen size, the tetraploid species having larger pollen grains. In the triploid plant, a large number of grains deviates from the normal 3-zonocolpate condition showing a number of apertural types, such as 1-aperturate-operculate, 2-colpate, 3-syncolpate, 4-zonocolpate and spiraperturate. Besides, planaperturate 3-colpate-trilobed grains also occur. The triploid is also characterised by 'minigrains' (size 13-19 μm), a feature not observed in A. mexicana and A. ochroleuca. Since these species are reported to be genetically close, the occurrence of the triploids (forming nearly 5% of the population of A. ochroleuca) indicates the possibility of a natural hybridization between A. ochroleuca and A. mexicana, which is well pronounced on its anomalous pollen features serving as marker for such a hybrid occurring in nature.     

Cytotype distribution at a diploid-tetraploid contact zone in Chamerion (Epilobium) angustifolium (Onagraceae)

In North America, the geographic distributions of diploid and tetraploid Chamerion (formerly Epilobium) angustifolium overlap in a narrow zone along the southern border of the boreal forest and along the Rocky Mountains. We examined the frequency and distribution of diploid and tetraploid cytotypes in a narrow (5 km) zone of sympatry across an elevational gradient and in putatively uniform diploid and tetraploid reference populations on the Beartooth Pass, in the Rocky Mountains of southern Montana-northern Wyoming. All five reference populations sampled were dominated by a single cytotype, but only one was completely uniform. In the zone of sympatry, 27 transects were sampled every 2 m for a total of 238 plants. Reproductive status (vegetative, flower buds, open flowers) was recorded, and the ploidy of each plant was determined by flow cytometry. Diploid and tetraploid plants predominated (36 and 55%, respectively) but were heterogeneously distributed among the transects. Six of the 27 transects were fixed for a single cytotype (four transects, diploid; two transects, tetraploid), and in seven others either diploids or tetraploids predominated (frequency >75%). Triploids represented 9% of the total sample and occurred most frequently in transects containing both diploids and tetraploids (G = 3.4, df = 2, P = 0.07). Diploids were more often reproductive (in bud, flower, or fruit) than either triploids or tetraploids (G = 12.0, df = 2, P < 0.001) and were the only cytotype to have produced open flowers. These results suggest that the zone of sympatry is best characterized as a mosaic rather than a cline, with diploid and tetraploids in close proximity and that the distribution of polyploidy is regulated by ecological sorting in a heterogeneous physical environment.     

Estimation of preferential pairing in tetraploid x diploid hybridizations

Summary Crosses made between tetraploid and diploid, 2n pollen-producing species directly transfer from one-half to the entire diploid genome from the diploid to the tetraploid level, depending on the mechanism of 2n pollen formation and the amount of crossing-over that occurs. Tetraploid plants that result from tetraploid x diploid hybridizations can be further utilized in a breeding program. It is postulated that preferential pairing between homologous chromosomes derived from the original tetraploid or diploid parent occurs in the tetraploid x diploid hybrid. Depending on the genetic divergence of the species involved, preferential pairing of homologous chromosomes may range from zero to one. Theoretical estimates of the amount of preferential pairing and the standard errors of these estimates are derived for cases where the diploid parent produces 2n gametes by either a first division or a second division restitution mechanism.     

An Examination of the Function of Male Flowers in an Andromonoecious Shrub Capparis spinosa

The pollen donor and pollinator attractor hypotheses are explanations for the functions of the male flowers of andromonoecious plants. We tested these two hypotheses in the andromonoecious shrub Capparis spinosa L. (Capparaceae) and confirmed that pollen production and cumulative volume and sugar concentration of nectar do not differ between male and perfect flowers. However, male flowers produced larger anthers, larger pollen grains and smaller ovaries than perfect flowers. Observations on pollinators indicated that two major pollinators (Xylocopa valga Gerst and Proxylocopa sinensis Wu) did not discriminate between flower morphs and that they transferred pollen grains a similar distance. However, there were more seeds per fruit following hand pollination with pollen from male flowers than from perfect flowers. Individuals of C. spinosa with a larger floral display (i.e. bearing more flowers) received more pollen grains on the stigma of perfect flowers. Female reproductive success probably is not limited by pollen. These results indicate that male flowers of C. spinosa save resources for female function and that they primarily serve to attract pollinators as pollen donors.     

秋水仙碱诱变甜菊多倍体的研究

用０．１％秋水仙碱溶液处理甜菊实生苗生长点,可诱变产生甜菊多倍体（四倍体）植株,用８次点滴处理,诱变率可达３１．２５％。染色体数鉴定表明：四倍体甜菊染色体数是２ｎ＝４４,而二倍体甜菊染色体数是２ｎ＝２２．形态学和解剖学的观察表明,四倍体比二倍体甜菊植株的茎矮壮,叶片增大,长度增长２．１倍,宽度扩大２．３倍,叶片加厚１．７倍,叶色更浓绿,叶片气孔数减少,气孔变大。叶片糖苷含量测定表明：四倍体的叶片含量为１５．７％,而二倍体的叶片含量为１０．８％,前者比二倍体叶片含量提高４．９％。     

The Hyparrhenia involucrata—H. subplumosa (Gramineae) complex in Nigeria: morphological and cytological characterization

Hyparrhenia involucrata Stapf is an annual with diploid, aneuploíd, tetraploid and hexaploid chromosome races on a base of x = 10, whilst M. subplumosa Stapf is known to be tetraploid and perennial. The two species overlap considerably in their vegetative and floral attributes. Whilst the floral characteristics (such as number of awns and sex of imperfect flowers) are fairly constant within H. involucrata , they are quite variable in H. subplumosa : the evolutionary significance of some of these variations is discussed. Some of the observations on size, variability and fertility of pollen grains are interpreted using data from chromosome numbers and meiotic behaviour. Preliminary observations on the hybrids between a diploid race of H. involucrata and H. subplumosa indicate that the genomes of the two species are highly homologous and that perennial habit is a dominant trait controlled by a single gene.