Comparative cytogenetic investigation of the two subspecies of the grass Alloteropsis semialata (Poaceae)

A cytotaxonomic investigation was undertaken to assess the taxonomic status of the grasses Alloteropsis semialata subsp. eckloniana and A. semialata subsp. semialata , distinguished because of a C3 photosynthetic pathway in subsp. eckloniana and a C4 pathway in subsp. semialata. Of the 30 analysed specimens of population A, 14 (46.7%) were diploid, 14 (46.7%) hexaploid, one octoploid (3.3%) and one dodecaploid (3.3%). Of the 21 specimens of population B, 14 (66.7%) were diploid, three (14.3%) hexaploid and four (19.0%) octoploid. All the diploids belonged to subsp. eckloniana , while all the polyploids belonged to subsp. semialata. Meiosis of the diploids appeared normal, with nine bivalents and a mean metaphase I chiasma frequency of 12.7 per cell. The hexaploids displayed a large range of chromosome pairing associations, although a high percentage of bivalents was recorded (89.9%). Three of the hexaploids showed 100% bivalent pairing, but the largest multivalent found in other hexaploids was a hexavalent pairing. The three octoploids analysed had 93.5% bivalent pairing. B chromosomes were found in five diploids, two hexaploids and one octoploid. Slow-moving bivalents, two in the diploids and up to four in the polyploids segregated late at anaphase I in most specimens.     

Nuclear variation withinAvenae

The amount of DNA per nucleus and the dry mass of isolated nuclei does not vary significantly between the six diploid species investigated. No differences were found either between tetraploid species or between hexaploids. Between the diploids, tetraploids and hexaploids taken as groups there was respectively a 1, 2 and 3 ratio maintained for the amount of DNA. For chromosome volume significant differences were found between the tetraploids but not in 2x or 6x series. Correlations are shown between chromosome volume, nuclear DNA content and nuclear dry mass. In polyploids the nuclear dry mass is less than expected from the values obtained for the diploids.Contribution No. 20 from the Canada Agriculture Research Station, Ste Foy, P.Q.     

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。     

Polyploidy in Dalea formosa (Fabaceae) on the Chihuahuan Desert

Dalea formosa consists of diploids (n = 7), tetraploids (n = 14), and hexaploids (n = 21), the polyploids restricted to the Chihuahuan Desert region or its immediate borders. There is very little morphological differentiation between the three chromosome races and, therefore, the polyploids are assumed to be primarily autoploid. Tetraploids discovered were few and were very similar to hexaploids; the two ploidy levels were combined as “polyploids” for analyses of geographically and cytologically correlated morphological variation. Pollen length generally was found to be greater in known polyploids than in known diploids. Through the use of pollen length and geographic origin, chromosomally unknown specimens were estimated as to ploidy level. This produced four groups, known diploids and polyploids, and putative diploids and polyploids, which were then subjected to stepwise discriminant analysis (SDA) to search for other morphological characters that might indicate ploidy level, to evaluate the assignments to putative ploidy level in unknown plants, and to assess correlation of these plants of putative ploidy level to geographic regions. SDA also indicated that pollen length, among ten morphological features, is the primary discriminator between ploidy levels, and that putative polyploids are confined primarily to the Chihuahuan Desert. Chromosomally unknown specimens that were originally assigned to one ploidy level, but were classified by SDA as another, are viewed as indicative of areas where further cytological sampling is particularly needed. These areas are southeastern Arizona, where pollen among known diploids is comparatively large, northeastern New Mexico, where polyploids might occur off the Chihuahuan Desert, east edge of the Chihuahuan Desert in Texas, a cytologically poorly sampled contact zone between diploids and polyploids, and central Coahuila, where no cytological sampling has been done. Canonical variate analysis is used to aid in the visualization of the general morphological relationship between diploids and polyploids.     

The Discovery of Tetraploid Camellia reticulata and Its Implication in Studies on the Origin of This Species

Camellia reticulata and its two allied species, C. saluenensis and C. pitardii,all from the Jinshajiang Valley of Yunnan and Sichuan, were cytologically studied in this work. Among 34 populations of C.reticulata studied here, 21 were found to be tetraploids (2n=60), 11 hexaploids (2n = 90), and the remaining two diploids (2n = 30). Tetraploid was reported in C. reticulata for the first time and its karyomophology was studied. In meiosis of pollen mother cells of the tetraploids and the hexaploids, only bivalents were observed in most of the populations, although in several populations or individuals a few univalents or tetravalents appeared. In the hexaploids, hexavalents were not observed. Thus, the meiotic chromosome cofigurations showed that the tetraploids and hexaploids in C.reticulata allopolyploids, and the appearance of a few tetravalents in some populations might indicatehomology of chromosomes in different genomes to some extent. The characteristics of the resting nuclei and the prophase chromosomes of all the tetraploids and hexaploids were quite similar to each other. The tetraploids of C.reticulata from the localities with the altitude of over 1800 m were gross-morphologically similar to the hexaploids, but those growing between 1100 ～1800 m were different to some extent from the hexaploids. In geographical distribution, the tetraploids and the hexaploids were continuous, and overlapped with C. saluenensis and C. pitardii.     

不同小麦进化材料生育后期光合特性和产量

以二倍体野生一粒小麦(Triticum boeoticum)、栽培一粒小麦(T. monococcum)、节节麦(Aegilops tauschii)和黑麦(Secale cereale)、四倍体野生二粒小麦(T. dicoccoides)、栽培二粒小麦(T. dicoccum)、硬粒小麦(T. durum)、六倍体普通小麦(T. aestivum)‘扬麦9号’和‘扬麦158’及八倍体小黑麦(Triticale)为材料,采用盆栽试验研究了不同小麦进化材料生育后期旗叶光合特性的演变及产量的差异。结果表明,与六倍体普通小麦和八倍体小黑麦相比,二倍体和四倍体材料在开花前具有较高的光合速率(P_n)、气孔导度(G_s)、最大光能转换效率(F_v/F_m)和实际光化学效率(Φ_PSⅡ)。开花以后,二倍体和四倍体材料受非气孔因素的影响,光合能力下降较快;除黑麦外,旗叶光合速率在开花10 d后都低于普通小麦和小黑麦,胞间CO₂浓度(C_i)迅速增加,F_v/F_m、Φ_PSⅡ和叶绿素含量快速下降。二倍体和四倍体材料开花前单株总叶面积和旗叶叶面积较大,花后下降迅速,功能期短;单株穗数也较多,但穗粒数、千粒重、产量和收获指数却显著低于普通小麦。因此,小麦长期进化过程中,普通小麦花后较高的光合能力及较长的光合持续期是提高千粒重,进而提高产量的重要生理基础。     

Polyploidy associated with altered and broader ecological niches in the Claytonia perfoliata (Portulacaceae) species complex

? Premise of the study: Polyploids are often hypothesized to have distinct and broader niches than their diploid progenitors. Differences in geographic distributions between diploid and polyploids are frequently used to infer niche differentiation and increased breadth, but they are seldom used to test these hypotheses explicitly. ? Methods: Niche overlap and breadth were compared for diploids, tetraploids, and hexaploids of three taxa in the Claytonia perfoliata complex (C. parviflora, C. perfoliata, and C. rubra) with the use of species distribution models. Resampling and randomization approaches were used to test hypotheses of niche differentiation, breadth, and conservatism. ? Key results: Niche differentiation was detected between polyploid and diploid cytotypes assigned to the same taxon (e.g., C. parviflora 2× vs. 4×) but not between hexaploids and tetraploids within a taxon (e.g., C. parviflora 4× vs. 6×). Individual hexaploid cytotypes had broader ecological niches than individual diploid cytotypes. However, as a group the three hexaploid taxa did not exceed the combined niche breadth of the three diploids, suggesting that polyploidy does not result in transgressive niche breadth for this group. Niche overlap was lowest among diploids and was highest among the three hexaploid cytotypes, consistent with introgression associated with polyploidy resulting in greater ecological similarity. Although cytotypes possessed nonidentical niches, after accounting for environmental differences among ranges, cytotypes were more similar than expected, suggesting niche conservatism and similar responses to environmental characteristics. ? Conclusions: These results suggest that polyploids occupy distinct and broader niches relative to diploids but that cytotypes also share fundamentally similar responses to environmental variation across ploidy levels.     

THE HORDEUM VIOLACEUM COMPLEX OF IRAN

Fifty-seven Iranian collections of Hordeum violaceum Boiss. & Huet, a perennial forage grass, contained diploid (2n = 14), tetraploid (2n = 28), and hexaploid (2n = 42) chromosome races. All collections came from moderate to high elevations in the Alborz and Zagros mountains and adjacent plateau areas of Iran. Each chromosome race had a discrete distribution, and the hexaploids were the most widespread. The diploids were cytologically regular, except for a chromosome interchange that occurred in about half of the plants. The tetraploids and hexaploids behaved cytologically as autopolyploids. The hexaploids were taller, coarser and later-flowering than the diploids and tetraploids, and they had fewer but thicker culms and larger seeds. The tetraploids were the leafiest and most productive, making them the most desirable from an agronomic standpoint. All races were more or less self-sterile, a characteristic that sets H. violaceum apart from most other Hordeum species. The taxonomic status of H. violaceum and its closest relatives, H. turkestanicum Nevski and H. brevisubulatum Link, is uncertain because of close morphological similarities and the occurrence of chromosome races in each taxon.     

Dry Matter Yields and Photosynthetic Rates of Diploid and Hexaploid Triticum Species

The above ground dry matter yields of two wild diploid Triticumspecies averaged 76 per cent of that of two hexaploid breadwheat varieties in field trials carried out over five years.A cultivated diploid species, T. monococcum, gave similar drymatter yields to the bread wheat varieties but had a longergrowth cycle. The flag leaves of wild diploid species had higher rates ofphotosynthesis than those of the bread wheat varieties bothwhen expressed per unit area of leaf or per unit weight of chlorophyll.Photosynthetic rates of other organs, expressed per unit weightof chlorophyll were also greater for the wild diploids thanfor hexaploids. For snoots at the stage when their flag leaveswere fully expanded, the investment in photosynthetic machinery,as measured by chlorophyll concentration, was less in the twowild diploids than in the hexaploids. This compensated for thehigh photosynthetic rate of the former, such that the specificgrowth rates, assessed by carbon-14 fixation per unit shootdry matter, were similar. Triticum spp., wheat, dry matter yield, photosynthesis, carbon-14, ploidy     

Morphological and cytotype variation of wild kiwifruit (Actinidia chinensis complex) along an altitudinal and longitudinal gradient in central‐west China

The Actinidia chinensis complex, a group of commercially important fruits (kiwifruit), is a complex of functionally dioecious lianas of variable ploidy. To understand the cytogeography better and to facilitate breeding in this complex, we examined the ploidy and morphological variations in 16 natural populations of A. chinensis var. chinensis and A. chinensis var. deliciosa across an ecogeographical gradient. Four ploidy levels were found, var. chinensis consisting of diploids and tetraploids and var. deliciosa consisting of tetraploids, pentaploids and hexaploids. Hexaploids were centred in the western Yun‐Gui plateau, tetraploids coexisted with hexaploids or diploids in the middle Yun‐Gui plateau and the Wuling‐Xuefeng mountains, and diploids occurred in the eastern Wuling‐Xuefeng mountains and the Hunan foothills. These findings indicate a gradual, clinal transition from hexaploid to diploid across the elevational and longitudinal gradient. The clear geographical segregation of diploids and hexaploids may have arisen from their differential ecological adaptation in response to altitude and climate, whereas the coexistence of cytotypes (2x–4x, 4x–6x and 4x–5x–6x) might be a result of reproductive barriers, with a particular contribution from the postzygotic reproductive isolation between ploidy races. The geographical pattern and morphological variation of cytotypes suggest a hybrid zone between the varieties in the Wuling‐Xuefeng mountains. The differences in cytotypes which have arisen as a result of ecological adaptation, distribution and morphological characteristics will provide important baseline data for the selection of germplasm and the breeding of kiwifruit. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 72–83.     

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.     

Rapid displacement of a monoecious plant lineage is due to pollen swamping by a dioecious relative

Interspecific hybridization is recognized as a potentially destructive process that represents a major threat to biodiversity. The rate of population displacement by hybridization can be rapid, but underlying mechanisms are often obscure. One hypothesis is that a species may be driven to extinction by interspecific gene flow, or pollen swamping, when hybrids are inviable or sterile. Here, we document the rapid movement of two zones of contact between monoecious hexaploid and dioecious diploid populations of the wind-pollinated plant Mercurialis annua (Euphorbiaceae) in northeastern and northwestern Spain, where diploids have displaced hexaploids by about 80 and 200 km, respectively, over a period of four decades. By using experimental mating arrays, we show that hybridization is highly asymmetrical in favor of the diploids, mainly because they disperse substantially more pollen, as expected in a comparison between an obligate outcrosser and a facultative selfer. Self-fertilization, which is expected to reduce the proportion of sterile hybrids produced in mixed ploidy populations, allowed the hexaploids to avoid the effects of pollen swamping only slightly, and in a density-dependent manner. Our results thus provide a mechanistic explanation for the rapid movement of both contact zones of M. annua in Spain.     

Chromosome studies of progenies of tetraploid female rainbow trout

Summary Nine induced tetraploid females were artificially inseminated by UV-irradiated sperm collected from diploid males, in order to induce the gynogenetic development of their ova. Most of the resulting embryos were diploid (or minor aneuploids). Several gynogenetic tetraploids, likely to issue from unreduced ova, were also detected in these progenies. The same females fertilized by normal sperm of diploid males gave a majority of triploids and several pentaploids, while the fertilization by normal sperm of tetraploid males gave rise to a majority of tetraploids and one hexaploid. The same crosses, after the eggs had been heat-shocked to double the maternal genetic contribution, yielded about three-quarters pentaploids and one quarter haploids (normal sperm of diploids), or three-quarters hexaploids and one quarter diploids (normal sperm of tetraploids). These haploids and diploids are likely to result from androgenesis.     

Polyploids and their structural and PHYSIOLOGICAL CHARACTERISTICS RELATIVE TO WATER DEFICIT IN BETULA PAPYRIFERA (Betulaceae)

The morphology and leaf anatomy, water relations, gas exchange, and growth characteristics of Betula papyrifera seedlings grown in a greenhouse representing three ploidy levels (diploids, pentaploids, and hexaploids) have been compared in an effort to understand the role of polyploidy in habitat differentiation and adaptation to water deficit. The polyploids are more tolerant of water deficit than their diploid relatives. The polyploids have fewer stomata per unit area and smaller stomatal indices than the diploids. They also have thicker upper and lower epidermis and more pubescence, which tend to reduce water loss. The polyploids have lower osmotic potential at saturation (_p.sat) than corresponding diploids, suggesting they are more able to maintain turgor in the case of low tissue water potential than the diploids. The photosynthesis of the diploids is more sensitive to water deficit. A 2-hr artificial water stress treatment caused complete cessation of photosynthesis in the diploids. They are more drought-avoiding than the polyploids as evidenced by closing their stomata to limit water loss at a relatively higher tissue water potential (-1.425 MPa). The pentaploids and hexaploids maintained, although to a small extent, photosynthesis when their tissue water potentials dropped to -1.86 MPa and -2.07 MPa, respectively.     

Sympatric diploid and hexaploid cytotypes of Senecio carniolicus (Asteraceae) in the Eastern Alps are separated along an altitudinal gradient

We explored the fine-scale distribution of cytotypes of the mountain plant Senecio carniolicus along an altitudinal transect in the Eastern Alps. Cytotypes showed a statistically significant altitudinal segregation with diploids exclusively found in the upper part of the transect, whereas diploids and hexaploids co-occurred in the lower range. Analysis of accompanying plant assemblages revealed significant differences between cytotypes along the entire transect but not within the lower part only, where both cytotypes co-occur. This suggests the presence of ecological differentiation between cytotypes with the diploid possessing the broader ecological niche. No tetraploids were detected, indicating the presence of strong crossing barriers. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Elucidating polyploidization of bermudagrasses as assessed by organelle and nuclear DNA markers

Clarification of relationships among ploidy series of Cynodon accessions could be beneficial to bermudagrass breeding programs, and would enhance our understanding of the evolutionary biology of this warm season grass species. This study was initiated to elucidate polyploidization among Cynodon accessions with different ploidy series collected from Turkey based on chloroplast and nuclear DNA. Forty Cynodon accessions including 7 diploids, 3 triploids, 10 tetraploids, 11 pentaploids, and 9 hexaploids were analyzed using chloroplast DNA restriction fragment-length polymorphism (cpDNA RFLP), chloroplast DNA simple sequence repeat (cpDNA SSR), and nuclear DNA markers based on neighbor-joining (NJ) and principle component analyses (PCA). All three-marker systems with two statistical algorithms clustered the diploids apart from the other ploidy levels. Assuming autopolyploidy, spontaneous polyploidization followed by rapid diversification among the higher ploidy levels than the diploids is likely in Cynodon's evolution. Few tetraploid and hexaploid accessions were clustered with or closely to the group of diploids, supporting the hypothesis above. Eleven haplotypes as estimated by cpDNA RFLP and SSR markers were detected. This study indicated that the diploids had different organelle genome from the rest of the ploidy series and provided valuable insight into relationships among ploidy series of Cynodon accessions based on cp and nuclear DNAs.     

Cytogeography of the Phleum pratense group (Poaceae) in the Carpathians and Pannonia

Cytogeographical variability within the Phleum pratense group in the Carpathians and adjacent part of Pannonian lowland, based on 132 populations analysed by flow cytometry, is described. Only diploid and hexaploid plants were detected among 635 samples from the studied area. Diploids were found to be less frequent (127 plants, 20%) than hexaploids (508, 80%). With the exception of the single pure diploid population, diploids always co-occured with hexaploids (30 localities, 22.7%). The majority of populations (101, 76.5%) consisted of hexaploid plants. Most mixed populations occur in the Western Carpathians (26). In the Eastern Carpathians, mixed populations are much rarer, with three populations in Ukraine and one in Romania. In the Southern Carpathians, only hexaploids occur. The conventional taxonomic concept of the two species, diploid P. bertolonii and hexaploid P. pratense , was followed in spite of their sympatric occurence. Distribution maps based on chromosome number data from previous studies and on ploidy level estimates are given for both species in the studied area. The pattern of different distribution of the two taxa within the Carpathians is discussed.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 475–485.     

Agrostis mertensii (Poaceae) in hokkaido

Populations of theAgrostis flaccida-mertensii complex in Hokkaido were examined as part of the taxonomic study of this complex, which has a wide range and comprises a polyploid series of 2x–8x (x=7). The principal aim was to clarify the taxonomic status of hexaploids growing in Hokkaido. Field explorations were made in the mountains and on the coasts in Hokkaido, and the materials were gathered from 19 localities. The occurrence of four cytotypes (2x, 4x, 6x, 8x), as well as some triploids and septaploids, of this complex in Hokkaido was confirmed, and the ranges of respective cytotypes were delineated. Morphological examinations showed that diploids, triploids and tetraploids are referable toAgrostis flaccida and octoploids toA. mertensii. The hexaploids of Hokkaido were morphologically different fromA. flaccida and also from the related hexaploid species,A. tateyamensis, distributed in Honshu. The hexaploids were quite similar to octoploidA. mertensii in all the morphological features examined and also in habitat preference. Septaploids were sterile hybrids between hexaploids and octoploids. A conclusion was drawn that the hexaploid plant of Hokkaido should be grouped with the octoploid in one and the same species,Agrostis mertensii Trin.Agrostis mertensii in this amended sense is discussed from a cytogeographical viewpoint.     

Cytogenetic characterization of Brazilian Paspalum accessions

Chromosome number, pairing relationship and meiotic behavior were evaluated in 24 Brazilian accessions of different Paspalum species as an initial screening to determine which of them might be useful in an interspecific hybridization program. The analysis showed that six were diploids, 16 tetraploids and two hexaploids. The pairing relationship was typical for the ploidy level and agreed with reported data. However, the meiotic behavior after diakinesis was much more abnormal than expected considering the pairing relationship. There was a high frequency of abnormal tetrads in the majority of accessions.