云南山茶花四倍体的首次发现及其科学意义

本文对分布于云南和四川金沙江河谷的云南山茶花C. reticulata及其两个近缘种(怒江山茶C.saluenensis和西南山茶C．pitardii)进行了细胞学研究。34个居群的云南山茶花中,21个居群是四倍体类型(2n=60),11个居群是六倍体类型(2n=90),另2个居群为二倍体(2n=30),云南山茶花的四倍体类型为首次发现,并且进行了核形态研究。四倍体和六倍体的花粉母细胞减数分裂染色体构型在大多数居群都为二价体(四倍体中30个二价体,六倍体中45个二价体),少数居群或是个体除二价体为主外还出现单价体和四价体,六倍体类型没有出现六价体构型。根据减数分裂的构型,我们认为,四倍体和六倍体分别为异源四倍体和异源六倍体,少数四价体的存在表明染色体有部分同源性。所有四倍体和六倍体的体细胞间期核特征和前期染色体形态特征基本相似。中海拔(1800m)以上的四倍体的云南山茶花外部形态特征与六倍体类型比较相似,而低海拔(1100～1800 m)的四倍体类型的外部形态特征则有些不同,但核形态结构是比较相似的。四倍体与六倍体类型地理分布是连续的,并与近缘的二倍体种怒江山茶、西南山茶重叠分布。     

探讨云南山茶起源的一线曙光—野生二倍体类型在金沙江流域的发现

新近发现的云南山茶原始二倍体类型分布于金江河谷流域的云南华坪悬及四川盐边县,海拔１８００－２８００ｍ。其植被模型为亚热带常绿阔叶林与云南松的混交林,生境较为湿润。云南山茶原始二倍体类型的形态特征与广布的六倍体类群非常相似,而与怒江山茶及西南山茶的形态特征不同。二倍体的花粉母细胞在减数分裂中期Ｉ形成１５个二价体,间期的细胞核结构为球形前染色体型,分裂前期染色体为中间型,其核型公式为２ｎ＝３０＝２２ｍ     

The origin and meiotic behaviour of hexaploid northern wheatgrass (Agropyron dasystachyum)

The occurrence of hexaploid (2n = 6x = 42) forms in some otherwise natural tetraploid populations of Agropyron dasystachyum (2n = 4x = 28) was cytologically detected and studied. The hexaploid plants are morphologically similar to the tetraploids except for a small reduction in the anther size. The general survey of chromosome numbers of natural Northern Wheatgrass (A. dasystachyum 2n = 4x = 28) populations derived from eight different regions of Alberta indicated that the occurrence of hexaploid variants was not restricted to a single locality. A comparative study of chromosome pairing in the natural and the synthetic hexaploids revealed that the naturally occurring 42-chromosomed plants of A. dasystachyum originated as a result of fertilization between unreduced (SSHH) and the natural (SH) gametes, both coming from the tetraploid form of A. dasystachyum. Based on chromosome pairing, the genomes of the natural hexaploid A. dasystachyum have been designated as SSSHHH. The natural hexaploids appear to intercross among themselves and also with tetraploids producing euploid and aneuploid hybrids. The possible evolutionary significance of these findings is briefly discussed.     

A CYTOGEOGRAPHIC STUDY OF CHAENACTIS DOUGLASII (COMPOSITAE,HELENIEAE)

Analysis of 512 plants derived from 200 populations shows that the widely distributed western North American Chaenactis douglasii species-complex consists of diploids (n = 6), triploids, tetraploids, and hexaploids. Microsporocytes were the source of most of the chromosome counts. About 9% of all plants examined had one or more full-sized extra chromosomes. Multivalents, usually a ring or chain of four chromosomes, were almost entirely restricted to polyploids, where one or more were identified in 38% of the tetraploids and 33% of the hexaploids. With two exceptions, diploids and polyploids were not found in the same population. Frequencies of diploid, triploid, tetraploid, and hexaploid populations were, respectively, 34, 1.5, 55 and 9.5%. With significant exceptions, diploid populations predominate in the Pacific and Rocky Mountain Systems, whereas polyploid ones are most frequent in the intervening plateaus. Ploidy level is correlated with age of substrate, rather than with climate, elevation, vegetation, or soil type. Range, morphology, ploidy level, and meiotic behavior suggest that var. achilleifolia tetraploids and hexaploids are descendents of hybrids between other variants of the complex. The diploid-tetraploid-hexaploid geographic distribution and the age of the substrates where each tends to occur suggest that the complex evolved in late Cenozoic time in response to major climatic and geologic changes that induced migration and hybridization. The hybrid derivatives, stabilized by polyploidy and tolerant of increasing aridity, came to occupy newly available habitats in areas disturbed by volcanic activity and glacial or glacial-related processes.     

Tempo and mode of recurrent polyploidization in the Carassius auratus species complex (Cypriniformes,Cyprinidae)

Polyploidization is an evolutionarily rare but important mechanism in both plants and animals because it increases genetic diversity. Goldfish of the Carassius auratus species complex can be tetraploids, hexaploids and octaploids. Polyploidization events have occurred repeatedly in goldfish, yet the extent of this phenomenon and its phyletic history are poorly understood. We explore the origin, tempo and frequency of polyploidization in Chinese and Japanese goldfish using both mitochondrial (mtDNA) and nuclear DNA sequences from up to 1202 individuals including the outgroup taxon, Cyprinus carpio. Analyses of de novo nuclear gene data resolve two clusters of alleles and the pattern supports the prior hypothesis of an ancient allotetraploidization for Carassius. Alleles shared by tetraploid and hexaploid individuals indicate recent autoploidizations within the C. auratus complex. Sympatric tetraploids and hexaploids share mtDNA haplotypes and these frequently occur independently within six well-supported lineages and sublineages on a small spatial scale. Gene flow estimates (F_st values) indicate that hexaploids differ only slightly from sympatric tetraploids, if at all. In contrast, allopatric populations of tetraploids and hexaploids differ from one another to a far greater extent. Gene flow between sampled localities appears to be limited. Coalescence-based time estimations for hexaploids reveal that the oldest lineage within any sampled locality is around one million years old, which is very young. Sympatric, recurrent autoploidization occurs in all sampled populations of the C. auratus complex. Goldfish experience polyploidization events more frequently than any other vertebrate.     

金沙江中游地区红山茶组植物的Giemsa C－带研究

研究了金沙江中游地区红山茶组植物的ＧｉｅｍｓａＣ－带。该地区的红山茶植物以四倍体为主,个别居群为二倍体或六倍体。居群间的Ｃ－带差异明显,Ｃ－带多出现在染色体端部。在四倍体和六倍体的Ｃ－带带型中,只能找到２条显相同Ｃ－带的同源染色体,通过与其它地区的红山茶植物进行比较,发现红山茶组植物的倍性从华东,华南经贵州,四川向云南逐渐增高,显Ｃ－带的染色体与染色体总数之比随信性增加而减少。文中指出华东或华南可能是红山茶组植物的起源地,而金沙江中游地区是其现代分化中心,这一地区红山茶的多倍体类群可能是异源起源的。     

Calamagrostis hakonensis (Poaceae): Distribution and differentiation of cytotypes

Chromosome counts were made for a total of 540 collections ofCalamagrostis hakonensis from 66 localities ranging from Kyushu to Hokkaido. Distribution and habitat preferences of the cytotypes involved are described. Sympatric occurrence of tetraploids (=semidiploids, 2n=28) and higher polyploids, as well as that of septaploids and plants at hexaploid and/or octoploid levels, was confirmed in several localities, and mixtures of hexaploids and octoploids within a population were frequently observed. Plants at hexaploid and octoploid levels were the most abundant and widespread. An examination of pollen of the voucher specimens showed that tetraploids (amphimictic) had good pollen, while higher polyploids (apomictic) were generally devoid of pollen and very rarely produced moderately good pollen, the grains of higher polyploids being larger than those of tetraploids. Distributions of tetraploids and higher polyploids were more precisely delineated through an examination of pollen of many herbarium specimens. In relation to the processes by which the complicated internal structure ofC. hakonensis has been established, the following subjects are discussed: infraspecific hybridization between ecotypically differentiated populations, enrichment of variability through occasional sexual reproduction expected in plants with more than 2n=42, persistence of the variants by apomictic reproduction, and probable roles of some extinct taxa.     

A cytotaxonomic study of theCalamagrostis purpurea —langsdorffii—canadensis complex in the lowlands of hokkaido

A chromosome survey of 278 individuals from 52 localities confirmed, that the principal components of theCalamagrostis purpurea—langsdorffi—canadensis complex in the lowlands of Hokkaido are tetraploid (2n=28) and octoploid (2n=56). A few hexploids (2n=42), were also found, but they may be either a triploid of the tetraploid plants or a hybrid between the tetraploid and octoploid plants. These hexaploids clearly differ in origin from the North American plants with 2n=42 which are generally placed underC. canadensis. The tetraploids and the octoploids are significantly different in pollen size but resemble each other in other morphological features and ecological requirements. These two races are also not disparate in distribution within Hokkaido. The tetraploids in Hokkaido are closely related to the tetraploids known in Alaska, and these tetraploid plants may be continuously distributed from Hokkaido to Alaska. The octoploids in Hokkaido have shown good pollen. They seem to have had a somewhat different course of evolution fromC. purpurea s. str. which principally reproduces in an apomictic manner.     

Phytogeographical studies ofCalamagrostis sachalinensis (Gramineae)

Chromosome numbers were determined on 223 collections ofCalamagrostis sachalinensis from 18 localities in Japan. The plants were found to be tetraploid (2n=28), hexaploid (2n=42) or octoploid (2n=56). A few collections were found to include one or two B-chromosomes. The tetraploid collections were made from central Honshu and Mt. Apoi in Hokkaido, while the hexaploids and the octoploids were detected in many localities. Pollen examination of these collections showed that the tetraploids with but one exception have good pollen and the hexaploids and the octoploids have no pollen or have bad pollen with stainability less than 10%. With the help of pollen examination of a number of herbarium specimens, the distribution of the tetraploids and that of the assemblage of the hexaploids and octoploids were delineated. Morphological studies indicated that the tetraploid, hexaploid and octoploid plants can not be separated in gross and spikelet morphology and that the tetraploids in central Honshu and those in Mt. Apoi are significantly different in leaf features. It was concluded thatC. sachalinensis represents an apo-amphimictic complex, which includes the following four races: 1) tetraploid, amphimictic, having thin leaf blades 5–10 mm broad and growing on the subalpine conifer forest belt and the conifer forest-alpine ecotone in the mountains of central Honshu; 2) tetraploid, amphimictic, having hard leaf blades 2–6 mm broad and growing on the stony, arid and exposed alpine belt on Mt. Apoi in Hokkaido; 3) hexaploid, mainly apomictic, the most variable ecologically, widely distributed; 4) octoploid, mainly apomictic, frequent in the upper montane to alpine belts, probably widely distributed.     

Cytogeography of the South American Turnera sidoides L. complex (Turneraceae, Leiocarpae)

New chromosome number determinations are presented for 48 populations of five subspecies of Turnera sidoides. Chromosome counts in sspp. carnea, holosericea, integrifolia and sidoides confirm previous data while, in ssp. pinnatifida , the numbers 2n=2x = 14 and 2n = 6x = 42 are reported for the first time. The results show that polyploidy is a very frequent phenomenon. Populations are primarily tetraploids (2n = 4x = 28), while diploids (2n = 2x = 14), hexaploids (2n = 6x =42) and octoploids (2rc = 8x = 56) are less frequent. Diploid, tetraploid and hexaploid cytotypes occur in sspp. carnea and pinnatifida , whereas tetraploids and hexaploids are known in ssp. holosericea. Turnera sidoides ssp. integrifolia shows a polyploid series with ploidy levels from 2x to 8x. The ssp. sidoides appears to be uniformly tetraploid, with the exception of an isolated pentaploid (2rc = 5x = 35). On the basis of these counts and previous reports the geographical distribution of the cytotypes has been determined and related to climatic and ecological factors.     

Genetic Diversity in Crucian Carp (Carassius auratus)

A survey of restriction fragment polymorphism in mitochondrial DNA of three subspecies of Carassius auratus throughout four provinces in China was undertaken using 17 restriction enzymes. Two carp, Cyprinus carpio rubbrofuscus and Cyprinus carpio carpio, were included as the outgroup. A total of 16 haplotypes was observed: 5 in tetraploids of C. auratus auratus; 8 in hexaploids of C. auratus auratus; and 2 in C. auratus gibelio and C. auratus cuvieri, respectively. The tetraploids and hexaploids share three common haplotypes as I, V, and VI. C. a. Cuvieri may have diverged first among the three subspecies. Interestingly, C. a. auratus and C. a. cuvieri did not form monophyletic clades, which indicated that the classification of Carassius auratus required further studies. The current hypothesis, that hexaploids originated from tetraploids by a polyploidy event, is less favorable, based on the distribution of haplotypes and the lower diversity in tetraploids than in hexaploids. Our data also indicate that divergence of hexaploids and tetraploids might be recent and mtDNA polymorphism existed before the divergence. Meanwhile, genetic isolation exists between the hexaploids and the tetraploids.     

Cytological and morphological variation in Bellevalia

Material from 45 populations representing 15 species of Bellevalia were analyzed as to chromosome numbers and morphology. For 3 species reports are new, viz. for B. clusiana (2x), B, tauri (2x, 4x), and B. nivalis (4x). In B. saviczii diploids, tetraploids and hexaploids were found. The basic Bellevalia karyotype prevails, but chromosome heteromorphism is rather common at the populational level. B–chromosomes were found in some populations of B. glauca s.l. and B. saviczii. Discussions on karyotype variation, polyploidy, morphologic variation and generic delimitation are presented.     

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.     

DNA ploidy-level variation in native and invasive populations of Lythrum salicaria at a large geographical scale

Aim  This study aimed to document precisely the patterns of DNA ploidy variation in the native and secondary ranges of Lythrum salicaria distribution. The hypothesis that species invasiveness had been induced by a switch in ploidy level was addressed.
Location  Europe, Middle East, North America.
Methods  DNA ploidy levels of 1884 progenies of 578+ plants collected at 124 localities were determined by DAPI flow cytometry.
Results  Large cytotype variation (2 x , 3 x , 4 x and 6 x ) was found across the native area of distribution (64 populations covering 12 European and two Middle Eastern countries). DNA hexaploids were detected for the first time, and rare DNA triploids were reliably confirmed. DNA tetraploids largely prevailed across the native range studied, while DNA diploids and DNA hexaploids were recorded only in Israel and Turkey, respectively. DNA triploid progenies occurred in one population from Hungary (together with DNA tetraploids). Sympatric growth of DNA tetraploids and DNA hexaploids was repeatedly encountered in Turkey. In contrast, cytotype uniformity was a typical feature of the invasive North American plants. Sixty populations, covering 13 states of the USA and provinces of Canada, were characterized by the presence of only DNA tetraploids.
Main conclusions  Several L. salicaria cytotypes (2 x , 3 x , 4 x , 6 x ) occur in the native range of distribution, with much variation concentrated in the Middle Eastern countries, whereas only DNA tetraploids appeared to occur in North America. Our data show that the invasive spread of North American populations was not triggered by differences in ploidy level. Alternative explanations should be sought.     

Numerous mitochondrial DNA haplotypes reveal multiple independent polyploidy origins of hexaploids in Carassius species complex

Evolutionary trajectory and occurrence history of polyploidy have been extensively studied in plants, but they remain quite elusive in vertebrates. Here, we sampled and gathered 4,159 specimens of polyploid Carassius species complex including 1,336 tetraploids and 2,823 hexaploids from a large geographic scale (49 localities) across East Asia, and identified a huge number of 427 diverse haplotypes of mitochondrial control region, in which 74 haplotypes with total occurrence frequency up to 75.498% were shared by hexaploids and tetraploids. Significantly, these diverse haplotypes were clustered into four major lineages, and many haplotypes of hexaploids and tetraploids were intermixed in every lineage. Moreover, the evolutionary trajectory and occurrence history of four different lineages were revealed by a simplified time‐calibrated phylogenetic tree, and their geographic distribution frequencies and haplotype diversity were also analyzed. Furthermore, lineage C and D were revealed to undergo population expansion throughout mainland China. Therefore, our current data indicate that hexaploids should undergo multiple independent polyploidy origins from sympatric tetraploids in the polyploid Carassius species complex across East Asia.     

Meiotic behavior and pollen fertility in triploid and tetraploid natural populations of Campuloclinium macrocephalum (Eupatorieae,Asteraceae)

Campuloclinium macrocephalum DC. is a perennial herb widely distributed in the New World and introduced in South Africa, where it is commonly called “pompom weed”. This species is considered one of the most important weeds of Brazil and one of the problematic invasive plants of South Africa. The meiotic system can be studied to assess the ability of a weed to spread, but only few studies on C. macrocephalum have been realized. In this study, we examined the meiotic behavior and pollen fertility of 14 natural populations of C. macrocephalum from Argentina and Uruguay. Meiotic analysis revealed 2 triploid (2n = 3x = 30), 11 tetraploid (2n = 4x = 40) and 1 mixed population (2n = 2x = 20, 2n = 4x = 40). Both, triploid and tetraploid specimens showed a widely variable meiotic behavior with irregular chromosome pairing showing univalents, bivalents, trivalents (in triploids) and tetravalents (in tetraploids) at diacinesis of first meiotic division. Different abnormalities were observed, such as: laggard chromosomes, chromatin bridges, and out of plate chromosomes at metaphase I. During meiosis I (prophase), some cells showed the phenomenon of cytomixis or chromatin transfer between pollen mother cells. The meiotic indexes suggest that only four populations were normally fertile (over 90 % of fertile pollen), indicating meiotically stable plants. The remaining populations share variable pollen fertility, with triploids ranging from 46.64 to 54.83 % and tetraploids varying from 3.54 to 45.30 %. We suggest that polyploidy seems to be recurrent in C. macrocephalum, promoting partial sterility of pollen grains, generating large numbers of individuals by apomixis promoting invasion of crop fields. This study presents the meiotic behavior of this weed, these could be useful for future studies of biological control in areas with no natural enemies.     

Taxonomic studies of the genusPoa of Japan

With the aim of making clear the boundaries between species in thePoa acroleucahisauchii-nipponica aggregate, chromosomes and morphological features of 746 collections gathered from 125 localities in Japan were examined. For morphological observations, the voucher specimens of 95 collections whose chromosome numbers were reported previously (Tateoka, 1985) were also used. Tetraploids (2n=28) and hexaploids (2n=42), as well as a few pentaploids which were the hybrids between 4× and 6×, were found. By examining morphological features of these collections, two groups were recognized in tetraploids and one in hexaploids. The two tetraploid groups corresponded toPoa acroleuca Steud. andP. hisauchii Honda, and the hexaploid group toP. nipponica Koidz. It was confirmed that the hairiness on the internerve surface of lemma, ligule hairiness and the length ratio of anther/ lemma are the most important features for discriminating between these species; panicle shape, leaf shape and anther length are also helpful for the identification. The ambiguity of the boundaries between species which was hitherto present in the taxonomy of this species aggregate was unrelated to the creation of nature itself but was attributable to the insufficiency of our research work.     

Cytological Study on Acorus L. in Southwestern China,with Some Cytogeographical Notes on A. calamus

Chromosome counts of three Acorus species ( A. calamus L., A. tatarinowii Schott and A.　gramineus Soland.) were performed in the Southwestern China. Both A. tatarinowii and A. gramineus shared the same chromosome numbers 2n=24. Populations of A.calamus in South and Southwestern Yunnan are tetraploids (2n=44) while those in the central and northwestern Yunnan being hexaploids (2n=66). The hexaploids are reported here for the first time. Attention was drawn to the ploidy pattern in A.calamus . It seems that the hexaploids were evolved from their ancestors during the early Tertiary as an impact of orogenesis of the Himalayas.