江西发现绵枣儿多倍体复合体多细胞型混合居群

对来自江西省新干县三湖镇绵枣儿(Scilla scilloides)多倍体复合体两个居群263个植株进行了细胞学调查,发现4个整倍体细胞型,即BB、BBB、BBBB和ABBB。其中BBBB和ABBB为大陆上首次记录。至此在东亚大陆上已经发现该多倍体复合体12个整倍体细胞型中的9个。这表明东亚大陆也是该复合体细胞型分化的重要地区之一。此地出现ABBB和BBB,而未出现ABB以及非整倍体可能是说明自第四纪以来BB细胞型在大陆和朝鲜半岛的分布区正在收缩;大陆上AABB已经接近消失;而大陆、朝鲜半岛和济州岛(韩国)的BB和AABB混合居群只是残遗。大陆与日本列岛居群细胞型组成的相似性可能暗示着它们的某种历史关系,但是并不能支持日本分布的绵枣儿是从中国引入的“史前归化”植物。     

Geographical distribution of cytotypes and genomic structures in natural populations of the<Emphasis Type="Italic">Scilla scilloides</Emphasis> complex in Korea

Cytotype distribution, B-chromosome frequency, and genomic constitution in the natural populations of theScilla scilloides complex in Korea were analyzed. Plants with various cytotypes were found: AA (2n = 16), BB (18), AAB (25), ABB (26), ABB (34), ABBB (35), BBBB (36), AABBB (43), AAABBB (51) and AAAABBBB (68). Allotetraploid AABB plants predominated with a frequency of 68.3%, and were found to distribute all over the Korean peninsula and Cheju-do. In diploids, the type AA plants distributed throughout the Korean midwest, while the type BB plants were limited to Cheju-do. Two other cytotypes, ABBB and AABBB, were found only in the southern part of the Korean peninsula including Cheju-do. Chromosomal variations, aneuploidy, and centromeric shifts were also found in the natural populations. The cytotypes AAB and AAAABBBB are reported here for the first time. B-chromosomes were found in 149 (85.6%) of 174 populations, the highest frequency being 81.8% in BB populations. The number of B-chromosomes per plant ranged from 1 to 31, and IB plants predominated (21.0%). Subtypes, with respect to the number and composition of B-chromosomes, indicated that sexual reproduction is still prevalent in AABB populations.     

Cytogenetics of theScilla scilloides complex

Four natural populations ofScilla scilloides Druce were analysed cytogenetically. The Ikeda and Taegu populations were pure populations composed of BB and AABB plants, respectively. The other two were mixed populations; the Shozugawa consisted of ABB, AABB and ABBB and the Nakabaru BB, BBB, ABB, AABB and ABBB. These populations have been maintained by sexual reproduction of sexually stable BB and AABB, and asexual propagation of sexually unstable ABB, BBB and ABBB. In addition, many kinds of aneuploids were observed at the high frequency of 18.1% in the Shyozugawa and 6.8% in the Nakabaru. Occurrence of these aneuploids would be due to sexual propagation of the sexually unstable ABB and ABBB plants. From the spatial distributions of each type within the mixed populations, it was apparent that sexual propagation of ABB plants was effective in the presence of the sexually stable BB and/or AABB individuals.     

Supplemental Cytogeographical Studies on the Scilla sinensis （Louriro） Merrill Complex in China

Seven hundred and fourteen individuals of the Scilla sinensis (Louriro) Merrill complex were sampled from 21 populations at 19 localities in 10 provinces and two cities of China and investigated cytologically in order to test its cytotype composition and cytogeographical variation still further. Five cytotypes were found (i.e. AA (2n = 16), BB (2n = 18), AABB (2n = 34), AAA (2n = 24), and AABB 9 (2n =43)). Until now, nine of the 12 cytotypes have been identified on the continent, which indicates that it is an important place for the cytotypic differentiation of the complex. The cytotype AABB 9 is a new record on the continent. AAA was recorded for the second time in this complex. The majority of populations are composed of only one cytotype (AA, BB, or AABB), whereas another four populations consist of more than two cytotypes involving AA, BB, AAA, AABB, and AABB 9. The distribution of BB covers the entire region south to Huai River, whereas AA populations are distributed in Taiwan, Fujian, Hong Kong, and Guangdong and are isolated from its major distribution. The appearance of AABB 9 is possibly evidence for the degeneration of the subglobular bulb type of AABB and the fact that AAA and AAAA exist in the interior border regions of the range of this complex probably suggests that drought is a major factor promoting plant polyploidization. In addition, the mean numbers of B chromosomes per individual, the percentage of individuals with the B chromosome in each population, and the correlation coefficients between the latter and their latitude/longitude were calculated. Generally speaking, the B chromosome number is higher in BB populations compared with AA populations and there is a prominent tendency that the B chromosome content in either AA or BB populations correlates with the latitude and longitude of the populations.     

Supplemental Cytogeographical Studies on the Scilla sinensis (Louriro) Merrill Complex in China

Seven hundred and fourteen individuals of the Scilla sinensis (Louriro) Merrill complex were sampled from 21 populations at 19 localities in 10 provinces and two cities of China and investigated cytologically in order to test its cytotype composition and cytogeographical variation still further. Five cytotypes were found (I.e. AA (2n = 16), BB (2n = 18), AABB (2n = 34), AAA (2n = 24), and AABB 9 (2n = 43)). Until now, nine of the 12 cytotypes have been identified on the continent, which indicates that it is an important place for the cytotypic differentiation of the complex. The cytotype AABB 9 is a new record on the continent. AAA was recorded for the second time in this complex. The majority of populations are composed of only one cytotype (AA, BB, or AABB), whereas another four populations consist of more than two cytotypes involving AA, BB, AAA, AABB, and AABB 9. The distribution of BB covers the entire region south to Huai River, whereas AA populations are distributed in Taiwan, Fujian, Hong Kong, and Gungdong and are isolated from its major distribution. The appearance of AABB 9 is possibly evidence for the degeneration of the subglobular bulb type of AABB and the fact that AAA and AAAA exist in the interior border regions of the range of this complex probably suggests that drought is a major factor promoting plant polyploidization. In addition, the mean numbers of B chromosomes per individual, the percentage of individuals with the B chromosome in each population, and the correlation coefficients between the latter and their latitude/longitude were calculated. Generally speaking, the B chromosome number is higher in BB populations compared with AA populations and there is a prominent tendency that the B chromosome content in either AA or BB populations correlates with the latitude and longitude of the populations.     

Cytogenetics of the Scilla scilloides complex

There are two cytogenetically well differentiated genomes, A (x=8) and B (x=9), in the Scilla scilloides complex. The principal cytogenetic types form an aneuploidal series of chromosome numbers, i.e., AA (2n=16), BB (2n=18), ABB (2n=26), BBB (2n=27), AABB (2n=34), ABBB (2n=35), BBBB (2n=36), and AABBB (2n=43). These types are widespread in the Japanese islands, excepting AA which is confined to Korea. On the contrary diploid BB is not known from Korea. However, polyploids AABB and AABBB are known from both Japan and Korea.Plants of the complex do not grow in wild lands or montane regions, but in close relation to man's activities, e.g., in agricultural lands, on river banks, along roadways and railway lines, and in graveyards. Natural populations are, as a rule, a mixture of many different cytogenetic types.The present study was supported financially by a Grant-in-Aid for Fundamental Scientific Research from the Japanese Ministry of Education; No. (A) 0810.     

Giemsa-C带揭示的绵枣儿多倍体复合体染色体变异

绵枣儿是分布在东亚的多倍体复合体。以Giemsa-C带法对分布在中国的绵枣儿24个居群进行了染色体变异分析。AA和BB细胞型大部分居群未发现变异,分别具a2和b1染色体的近着丝点着丝粒带。AABB虽无居群内变异,但居群间有所不同。其中柳林、商南和徐州居群带型相同,带纹丰富。元宝山居群只有b1的近着丝粒带,而灵岩山居群具a2和b1的近着丝粒带。根据已有的研究资料推测,绵枣儿b1和a2上的近着丝粒带显示的应当是高度重复的rRNA基因;大庆和文县居群一个体该位点的杂合可能是由于部分rRNA拷贝的同源染色体间转移;AABB起源后随时间推移会发生rRNA重复位点的丢失;柳林等居群丰富的带纹可能是其刚刚起源时重复基因关闭的表征,随时间推移同样可能会丢失。另外大量的实验证据表明,BB比AA细胞型可能更易于发生染色体结构变异。     

The distribution of diploids and polyploids of the Scilla scilloides complex in Korea

A total of 3 384 individuals of Scilla scilloides Druce were sampled from 40 natural populations at 35 different localities over the Korean Peninsula and analysed cytogenetically. The following cytogenetic types were found: AA (19.6%), AABB (72.4%), ABBB (4.3%), AABBB (3.0%) and aneuploids (0.8%). Of these, AA was found in 9 populations along the west side of the Korean Peninsula facing the Yellow Sea and on Yeongdo and Jeju Islands. Type AABB, the most common type in the complex, was found in 33 natural populations on the Korean Peninsula and Jeju Island. The distributions of ABBB and AABBB are restricted to the southern part of the Korean Peninsula and Jeju Island; they were found in 8 natural populations. In southern Korea the above four types showed overlapping distribution and two or three were often found growing together.Genome A, occurring in the three allopolyploids in the Korean Peninsula, was the same as that in allopolyploids in Japan. However, genome A occurring as AA diploids was karyotypically different from that in the allopolyploids. On the basis of distribution patterns and cytological features, the origin of the allopolyploids is discussed.     

Physical mapping by FISH and GISH of rDNA loci and discrimination of genomes A and B inScilla scilloides complex distributed in Korea

The chromosomal locations of the 18S-26S (45S) and 5S rDNA loci in cytotypes AA, BB, and AABB ofScilla scilloides Complex from Korea were physically mapped using multicolor fluorescencein situ hybridization (McFISH). Genomicin situ hybridization (GISH) was also performed to distinguish between the AA and BB genomes in allotetraploid AABB plants. One 18S-26S rDNA locus was detected in both AA (a2) and BB (b1 ); one locus also was found in the allopolyploid AABB (b1 ). This demon-strated the loss of that locus in genome A. GISH with biotin-labeled DNA from the BB genome and digoxigenin-labeled 18S-26S rDNA probes revealed that the 18S-26S rDNA in AABB plants was localized in the nucleolus organizer region (NOR) of genome B. One and two 5S rDNA loci were found in diploids AA and BB, respectively. As expected, all three 5S rDNA loci were detected in the AABB plants. The sequence identities of the 5S rDNA genes among cytotypes AA and BB, AA and AABB, and BB and AABB were 99%, 95%, and 95%, respectively. These authors contributed equally to this paper     

Origins, distribution, and local co-occurrence of polyploid cytotypes in Solidago altissima (Asteraceae)

There is growing realization that intraspecific polyploidy in plants has important implications for the evolution of plant diversity and for plant and plant-herbivore community dynamics. Nonetheless, the rate at which polyploid cytotypes arise and their fate in local mixed-cytotype populations are not well understood. Although within- and especially among-population cytotype variation has been documented in many plants, particularly at the largest (continental) spatial scales, local and regional distributions of cytotypes have been well characterized only for a handful of species. Furthermore, few if any studies have addressed both local and regional patterns in the same plant species with sufficient power to detect cytotype variation on both scales. We assessed the co-occurrence and frequency distributions of diploid, tetraploid, and hexaploid cytotypes of Solidago altissima (Asteraceae) at local and regional spatial scales, sampling across a zone of presumed ploidy-form overlap in the midwestern United States. Determining ploidy by flow cytometry, we found many local populations containing two or three cytotypes, with the relative frequencies of cytotypes varying dramatically even among neighboring populations. AFLP marker data suggest that polyploid cytotypes of S. altissima likely have multiple origins from different diploid lineages. Our results are consistent with recent perspectives on polyploidization as an evolutionarily dynamic phenomenon with frequent multiple origins of higher-ploidy cytotypes.     

The distribution of diploids and polyploids of theScilla scilloides complex in the northeastern district of China

A cytological analysis was made of 1504 individuals ofScilla scilloides Druce from the northeastern district of China. In terms of genome combination four cytotypes were found: AA (90.95%), AAAA (0.07%), AABB (8.92%) and AABB+2 (0.07%). In the cytotypes A and B denote genome A (x=8) and genome B (x=9), respectively. Most of 18 natural populations examined consist of one cytotype: indeed, all individuals examined of 15 of the populations are AA diploids, those of one population AABB allotetraploids, and individuals examined of two other populations have two cytotypes each, AA and AAAA, and AABB and AABB+2. In karyotype the genome A of the AA and AAAA was different from that of the AABB, and a direction of morphological change of chromosomes in the genome A is discussed in light of results of hybridization experiments reported elsewhere. Taken together with information available for the distribution of different cytotypes in China, Korea and Japan, the results of the present study support the view of Maekawa thatScilla scilloides was native to mainland China and lately introduced to Japan.     

Studies on the Asian Eupatorias

Analysis of 265 plants derived from 13 sites in the Rokko Mountains shows that the widely distributed eastern AsianEupatorium chinense var.simplicifolium consists of seven cytotypes. These polymorphic karyotypes comprise four levels of ploidy based on X=10 and partial deficiency, occurring either alone or in combination. The polyploid cytotypes exhibit anomalous meiosis and highly variable pollen stainability in contrast to the nearly-normal behaviour of the diploid. The polyploid cytotypes were revealed as being agamospermous. They are readily distinguishable from the diploid cytotypes by several morphological characters and by growth habits closely related to their respective niches. Two or more cytotypes occurred in all sites examined, the most common being the coexistence of triploid and tetraploid cytotypes. The frequencies of occurrence of the pentaploid plants and the cytotypes involving a deficient chromosome were relatively low. While the diploid cytotype is restricted to fragile gravitational slopes and rocky areas which are poor in species and lack tall competitors, the polyploids occur widely in the grasslands, the roadsides or the forest-margins, closely associated with tall grasses and forbs such asMiscanthus sinensis. Based on these data, theE. chinense var.simplicifolium complex is divided into two distinct groups: the diploid cytotype and the polyploid cytotypes.     

Cytotype segregation on regional and microgeographic scales in snow buttercups (Ranunculus adoneus: Ranunculaceae)

Polyploid speciation is an important source of angiosperm diversity. Insights into the origin and establishment of new polyploid species may be gained by studying the distributions of ancestral and derivative cytotypes at multiple spatial scales. Diploid (2n = 16) and tetraploid (2n = 32) snow buttercups (Ranunculus adoneus: Ranunculaceae) occur in the alpine of the central and southern Rocky Mountains. Root-tip squashes and flow cytometry were used to determine the ploidy of 1618 individuals from 35 populations. Samples from 31 of the 35 sites were entirely of one cytotype, either diploid or tetraploid. Diploid and tetraploid snow buttercups have nonoverlapping regional distributions. Where both cytotypes occur on the same site, the two are spatially segregated despite no apparent change in habitat. Triploid snow buttercups were only found at a diploid/tetraploid contact zone, while two hexaploid plants were found in tetraploid populations. Tetraploid establishment once or twice in the history of the species complex could account for the regional distribution of the two cytotypes. Habitat differentiation between cytotypes or reproductive exclusion of minority cytotypes may explain the observed segregation at both microgeographic and regional scales.     

Chloroplast DNA Intraspecific Phylogeography of Prairie Cordgrass (Spartina pectinata Bosc ex Link)

Chloroplast DNA (cpDNA) sequences are appropriate for studying intraspecific phylogeography. Comparing cpDNA phylogeny with different cytotypes provides insights into the origin and establishment of polyploid species. We have previously defined the geographic distribution of cytotypes in prairie cordgrass throughout the different regions of the United States. In this study, chloroplast haplotype variation is explored in 54 populations of prairie cordgrass, using nine noncoding chloroplast regions. These chloroplast analyses were combined with the cytotype surveys of prairie cordgrass to infer the phylogeography and to elucidate the origin of the different cytotypes. In this study, we identified three haplotypes, Prairie Cordgrass Group 1 (PCG1), Prairie Cordgrass Group 2 (PCG2), and Prairie Cordgrass Group 3 (PCG3). In general, related haplotypes were strongly associated with geographic distribution. Useful insertions–deletions (indels) were also found among prairie cordgrass populations. The PCG1 haplotypes collected inthe East North Central to the New England regions of the US, were polyploid (tetraploid, hexaploid, and octoploids), while the PCG2 haplotypes, found in southern South Dakota, Iowa, and Missouri, were primarily octoploids, but also included a small number of tetraploids. The PCG3 haplotypes were octoploids and were collected in North Dakota, South Dakota, and Minnesota.     

Molecular evidence for multiple polyploidization and lineage recombination in the Chrysanthemum indicum polyploid complex (Asteraceae)

The Chrysanthemum indicum polyploid complex comprises morphologically differentiated diploids, tetraploids and hybrids between C. indicum and C. lavandulifolium. The relationships between species and cytotypes within this complex remain poorly understood. Random amplified polymorphic DNAs (RAPDs), intersimple sequence repeats (ISSRs) and chloroplast SSR markers were used to elucidate the genetic diversity and relationships of the C. indicum polyploid complex. Molecular analysis of three diploid and nine tetraploid populations provided strong evidence for recurrent origins and lineage recombination in the C. indicum polyploid complex. The high similarity in molecular marker profiles and cpDNA haplotypes between the diploids and tetraploids distributed in the Shen-Nong-Jia Mountain area of China suggested an autopolyploid origin of the tetraploids, while the tetraploids from other populations may have originated via allopolyploidization. Lineage recombination was revealed by the extensive sharing of chloroplast haplotypes and genetic markers among the tetraploid populations with different origins. Multiple differentiation and hybridization/polyploidization cycles have led to an evolutionary reticulation in the C. indicum polyploid complex, and resulted in the difficulties in systematic classification.     

Intraspecific molecular phylogeny, genetic variation and phylogeography of Reticulitermes speratus (Isoptera: Rhinotermitidae)

Population structure was investigated in Reticulitermes speratus populations in the Korean Peninsula and the Japanese Archipelago. All trees derived from analyses of the combined sequence dataset of two mitochondrial genes, COII and COIII, showed that R. speratus populations cluster into two major clades comprising the Korean/southern Japanese populations and the north-ern Japanese populations. Analysis of population ge-netic structure showed strong genetic partitioning between populations of the two clades. To understand historical migration routes and current distributions, the phylogeographic history of R. speratus was inferred from intra-/interspecific phylogeny and diver-gence times estimated between the clades of the phylogenetic tree. The estimated migration route and divergence time of ancestral R. speratus are congruent with recent paleogeographic hypotheses involving land-bridge connections between the Asian continent and the Japanese Archipelago. We suggest that ancestral R. speratus separated into northern and southern Japanese populations after its migration into the Japanese main islands from East China during the early Pleistocene via the East China Sea basin, which may have been exposed during that period. The Korean populations seem to have diverged recently from southern Japanese populations; this may explain the current distribution of R. speratus in the Japanese Arachipelago, and account for why it is restricted to northern areas of the Tokara Strait.     

Analysis of East Asia Genetic Substructure Using Genome-Wide SNP Arrays

Accounting for population genetic substructure is important in reducing type 1 errors in genetic studies of complex disease. As efforts to understand complex genetic disease are expanded to different continental populations the understanding of genetic substructure within these continents will be useful in design and execution of association tests. In this study, population differentiation (Fst) and Principal Components Analyses (PCA) are examined using >200 K genotypes from multiple populations of East Asian ancestry. The population groups included those from the Human Genome Diversity Panel [Cambodian, Yi, Daur, Mongolian, Lahu, Dai, Hezhen, Miaozu, Naxi, Oroqen, She, Tu, Tujia, Naxi, Xibo, and Yakut], HapMap [ Han Chinese (CHB) and Japanese (JPT)], and East Asian or East Asian American subjects of Vietnamese, Korean, Filipino and Chinese ancestry. Paired Fst (Wei and Cockerham) showed close relationships between CHB and several large East Asian population groups (CHB/Korean, 0.0019; CHB/JPT, 00651; CHB/Vietnamese, 0.0065) with larger separation with Filipino (CHB/Filipino, 0.014). Low levels of differentiation were also observed between Dai and Vietnamese (0.0045) and between Vietnamese and Cambodian (0.0062). Similarly, small Fst''s were observed among different presumed Han Chinese populations originating in different regions of mainland of China and Taiwan (Fst''s <0.0025 with CHB). For PCA, the first two PC''s showed a pattern of relationships that closely followed the geographic distribution of the different East Asian populations. PCA showed substructure both between different East Asian groups and within the Han Chinese population. These studies have also identified a subset of East Asian substructure ancestry informative markers (EASTASAIMS) that may be useful for future complex genetic disease association studies in reducing type 1 errors and in identifying homogeneous groups that may increase the power of such studies.     

Phylogeography of a northeast Asian spruce, Picea jezoensis, inferred from genetic variation observed in organelle DNA markers

Range-wide genetic variation of the widespread cold-temperate spruce Picea jezoensis was studied throughout northeast Asia using maternally inherited mitochondrial DNA and paternally inherited chloroplast DNA markers. This study assessed 33 natural populations including three varieties of the species in Japan, Russia, China, and South Korea. We depicted sharp suture zones in straits around Japan in the geographical distribution pattern of mitochondrial haplotypes (GST=0.901; NST=0.934). In contrast, we detected possible extensive pollen flow without seed flow across the straits around Japan during the past population history in the distribution pattern of chloroplast haplotypes (GST=0.233; NST=0.333). The analysis of isolation by distance of the species implied that by acting as a barrier for the movement of seeds and pollen, the sharp suture zones contributed considerably to the level of genetic differentiation between populations. Constructed networks of mitochondrial haplotypes allowed inference of the phylogeographical history of the species. We deduced that the disjunction with Kamchatka populations reflects range expansion and contraction to the north of the current distribution. Within Japan, we detected phylogeographically different types of P. jezoensis between Hokkaido and Honshu islands; P. jezoensis in Honshu Island may have colonized this region from the Asian continent via the Korean peninsula and the species in Hokkaido Island is likely to have spread from the Asian continent via Sakhalin through land bridges. Japanese endemism of mitochondrial haplotypes in Hokkaido and Honshu islands might have been promoted by separation of these islands from each other and from the Asian continent by the straits during the late Quaternary.     

Polyploidy: a missing link in the conversation about seed transfer of a commonly seeded native grass in western North America

The use of local, native plant materials is now common in restoration but testing for polyploidy in seed sources is not. Diversity in cytotypes across a landscape can pose special seed transfer challenges, because the methods used to determine genetically appropriate materials for seed transfer do not account for cytotypic variation. This lack of consideration may result in mixing cytotypes through revegetation, which could reduce long‐term population viability. We surveyed nine populations of a native bunchgrass, Pseudoroegneria spicata, in three EPA Level III Ecoregions in the western United States to determine the frequency of polyploidy, whether there are differences in traits (phenotype, fecundity, and mortality) among plants of different cytotypes, and whether cytotype frequency varies among ecoregions. We assessed trait variation over 2 years in a common garden and determined ploidy using flow cytometry. Polyploidy and mixed cytotype populations were common, and polyploids occurred in all ecoregions. Four of the nine populations were diploid. The other five had tetraploids present: three had only tetraploid individuals whereas two had mixed diploid/tetraploid cytotypes. There was significant variation in traits among cytotypes: plants from tetraploid populations were larger than diploid or mixed populations. The frequency and distribution of cytotypes make it likely that seed transfer in the study area will inadvertently mix diploid and polyploid cytotypes in this species. The increasing availability of flow cytometry may allow ploidy to be incorporated into native plant materials sourcing and seed transfer.     

Reproductive differentiation into sexual and apomictic polyploid cytotypes in Potentilla puberula (Potentilleae, Rosaceae)

Background and Aims

Intraspecific reproductive differentiation into sexual and apomictic cytotypes of differing ploidy is a common phenomenon. However, mechanisms enabling the maintenance of both reproductive modes and integrity of cytotypes in sympatry are as yet poorly understood. This study examined the association of sexual and apomictic seed formation with ploidy as well as gene flow towards sexuals within populations of purely polyploid Potentilla puberula.

Methods

The study is based on 22 populations representing various combinations of five polyploid cytotypes (tetraploid–octoploid) from East Tyrol, Austria. Embryo ploidy and the endosperm/embryo ploidy ratio obtained by a flow cytometric seed screen were used to infer reproductive modes of seed formation and to calculate the male and female genomic contributions to the embryo and endosperm. Self-incompatibility (SI) patterns were assessed and a new indirect approach was used to test for the occurrence of intercytotype matings based on the variation in the male genomic contribution to sexually derived embryos on the level of developed seed.

Key Results

Tetraploids formed seeds almost exclusively via sexual reproduction, whereas penta- to octoploids were preferentially apomictic. Non-random distribution of reproductive modes within maternal plants further revealed a tendency to separate the sexual from the apomictic mode among individuals. Self-incompatibility of sexuals indicated functionality of the gametophytic SI system despite tetraploidy of the nuclear genome. We found no indication for significant cross-fertilization of tetraploids by the high polyploids.

Conclusions

The study revealed a rare example of intraspecific differentiation into sexual and apomictic cytotypes at the polyploid level. The integrity of the sexual tetraploids was maintained due to reproductive isolation from the apomictic higher polyploids. Functionality of the gametophytic SI system suggested that the tetraploids are functional diploids.