A review of chromosome cytology in Hyacinthaceae subfamily Ornithogaloideae (Albuca, Dipcadi, Ornithogalum and Pseudogaltonia) in sub-Saharan Africa

The chromosome cytology of Hyacinthaceae subfamily Ornithogaloideae is reviewed within the framework of a recent molecular-based classification, with particular emphasis on its center of diversity in sub-Saharan Africa. We also provide new chromosome counts for sections that are unknown or poorly known cytologically. Albuca subgen. Namibiogalum (9 spp.) probably has an ancestral base number of x = 10 but subgen. Albuca (± 70 spp), subgen. Monarchos (9 spp.) and subgen. Osmyne (36 spp.) have x = 9. The pattern in subgen. Urophyllon (3 spp.) is remarkable: although x = 6 is likely, the species in the section exhibit a range of 2n = 12, 10, 8, 6 and 4 (exclusive of polyploidy). All karyotypes have three large chromosome pairs and a variable number of small chromosomes. Pseudogaltonia (2 spp.) has x = 9 and Dipcadi (26 spp.) possibly x = 9 in series Uropetalum and x = 6 in series Dipcadi, which exhibits a pattern of descending dysploidy leading to n = 3 in D. marlothii. In Ornithogalum (± 130 spp.) chromosome numbers are known for only 24 of the ± 84 sub-Saharan species, mostly from subgen. Aspasia and subgen. Ornithogalum sect. Linaspasia, both of which have x = 6, and from subgen. Galtonia, which has x = 8. In contrast, x = 7 is basic for the Eurasian sects. Honorius and Melophis, and x = 18 seems likely for sect. Cathissa. Sect. Ornithogalum, the cytology of which we does not examine in detail, may have x = 9. Polyploidy is apparently rare in the sub-Saharan African ornithogaloids, in marked contrast to the high frequency of polyploidy among Eurasian species. In Albuca just 3 or possibly 4 sub-Saharan species (9% or 13% of those counted) are exclusively polyploid and 5 more have diploid and polyploid races; and in sub-Saharan Ornithogalum, only the tropical O. gracillimum is exclusively polyploid, and the western southern African O. hispidum has diploid and polyploid races.     

Production and characterization of arboreous and fertile <Emphasis Type="Italic">Solanum melongena</Emphasis> + <Emphasis Type="Italic">Solanum marginatum </Emphasis>somatic hybrid plants

In crop plants the shift from being annuals to perennials may allow future agricultural systems requiring less energy inputs. The practicability of this was tested for Solanum melongena. Leaf protoplasts of S. melongena (2n = 2x = 24) and one of the related arborescent species Solanum marginatum (2n = 2x = 24) were electrofused and fertile somatic hybrids with arborescent habit regenerated. The magnetic cell sorter (MACS) technique was used for the selection of heterokaryons. The hybrid nature of 18 regenerated plants was assessed on the banding patterns generated by inter-simple sequence repeat PCR. When taken to maturity in the greenhouse, hybrids grew more vigorously compared to the parental species. Their morphological traits were intermediate between those of S. melongena and S. marginatum. Hybrids flowered and produced an average of 85% stainable viable pollen and fertile fruits. The somatic hybrids were maintained in the greenhouse for more than 3 years and continued to produce flowers developing into two types of fruits with plentiful seeds. Fruits were either striated green containing non-germinable seeds or yellow with fully germinable seeds. Their S₁ progenies showed common features with S₀ hybrids, including fertility and arborescent habit. Cytologically, somatic hybrids exhibited the expected chromosome number of 2n = 4x = 48, while chromosome pairing during microsporogenesis was associated with a low frequency of intergenomic pairing. It is concluded that an arborescent perennial species has been obtained by somatic hybridization. The usefulness of this species per se or in eggplant breeding will depend not only on the transmission of the arborescent habit to cultivated eggplant varieties, but also on the variability that should be created from backcrossing the S. melongena + S. marginatum hybrids to S. melongena.     

Chromosome numbers and pollen fertility in the Senecio nemorensis group (Compositae) in the Carpathians

New chromosome numbers for two species from the Senecio nemorensis group: S. dacicus (2n = 40) and S. ucranicus (2n = 40) have been ascertained. The counts for S. germanicus Wallr. subsp. germanicus (2n = 40), S. hercynicus Herborg subsp. hercynicus (2n = 40), S. ovatus (P. Gaertn. et al.) Willd. subsp. ovatus (2n = 40) occurring in the Carpathians are also reported. The study confirmed only the known tetraploid chromosome number for the taxa of this group. The pollen fertility ranged from 82.09 to 92.99% in all examined species and subspecies, including their hybrids.     

Cytology and systematics in JapaneseArisaema (Araceae)

Chromosome numbers are determined from 37 populations attributed to 22 taxa of JapaneseArisaema. Of them, chromosome numbers ofA. limbatum var.conspicuum (2n=26),A. minus (2n=26),A. nambae (2n=28) andA. seppikoense (2n=26) are determined for the first time. New chromosome numbers, 2n=26, are reported forA. aequinoctiale, A. limbatum, A. stenophyllum, A. undulatifolium andA. yoshinagae. Three modes of basic chromosome numbers,x=14,x=13 andx=12, occur in JapaneseArisaema. Precise karyotypic comparisons of 20 taxa reveal that taxa withx=14 andx=13 share 26 major chromosome arms and have an obvious chromosomal relationship. One of two submeta-centric chromosomes inx=13 corresponds to two telo-centric chromosomes inx=14. InA. ternatipartitum with 2n=6x=72, ten out of 12 basic chromosomes are the most similar in size and arm ratio with larger ten chromosomes ofA. ringens among JapaneseArisaema examined. A basic chromosome number ofx=14 is the commonest in the genusArisaema and the remaining basic chromosome numbers,x=13 andx=12, seem to be derived through dysploidal reduction by translocating large segments of major arm of telo-centric chromosome onto other minor arm of telo-centric followed by loss of the remainings including a centromere, and by loss of two telo-centrics fromx=14, respectively. Some systematic problems of JapaneseArisaema are discussed based on new cytological data.Arisaema hatizyoense, A. minus andA. nambae are accepted as independent species.     

First report of chromosome numbers of the Carlemanniaceae (Lamiales)

The Carlemanniaceae comprises two small genera that are restricted to East Asia: the Carlemannia and Silvianthus. These genera were previously placed in the Rubiaceae or Caprifoliaceae, but are now considered a distinct family that is probably related to the Oleaceae in the Lamiales. The family is still poorly understood with respect to its morphological characteristics. Here, we present the first report of the chromosome numbers of the family using species from both genera, i.e., Carlemannia tetragona, Silvianthus bracteatus ssp. bracteatus, and S. bracteatus ssp. clerodendroides. The species were compared with the chromosome numbers of Oleaceae and associated families using a Bayesian tree that was generated from rbcL and ndhF sequence data from Genbank. C. tetragona had 2n = 30 (x = 15), whereas the two subspecies of Silvianthus had 2n = 38 (x = 19). Comparisons of chromosome numbers support the distinctness of the Carlemanniaceae, not only from the Oleaceae (x = 11, 13, 23), but also from the Tetrachondraceae (x = 10, 11), a family that is possibly related to the Carlemanniaceae and/or Oleaceae in the Lamiales. The notable difference in chromosome number between Carlemannia and Silvianthus, as well as the differences in other characteristics (pollen, seed, and fruit morphology), suggests that the family split early in its evolution.     

蟛蜞菊属和孪花菊属(菊科-向日葵族)的细胞学研究

研究了菊科向日葵族鳢肠亚族蟛蜞菊属(Sphagneticola O. Hoffm.)和孪花菊属(Wollastonia DC. ex Decne.)各2种植物的染色体数目和染色体形态。蟛蜞菊[S. calendulacea (L.) Pruski]的染色体数目为2n=50, 核型公式为2n=18m+30sm+2st,南美蟛蜞菊[S. trilobata (L.) Pruski]的染色体数目为2n=56, 核型公式为2n=24m+28sm+4st, 孪花菊[W. biflora (L.) DC.]的染色体数目为2n=30,核型公式为2n=24m+4sm+2st,山孪花菊[W. montana (Blume) DC.]的染色体数目为2n=74, 核型公式为2n=37m+31sm+6st。根据上述结果并结合以前的有关资料,推测蟛蜞菊属的染色体基数可能为x=14和x=25,而不应是x=15。该属的3个新世界热带种[S. brachycarpa (Baker) Pruski、S. gracilis (Richard) Pruski和南美蟛蜞菊]可能都基于x=14, 其中S. gracilis为二倍体(2n=2x=28), S. brachycarpa和南美蟛蜞菊为四倍体(2n=4x=56); 唯一的亚洲种(蟛蜞菊)可能是基于x=25的二倍体(2n=2x=50)。染色体资料不支持将山孪花菊(x=37)这一植物置于孪花菊属(x=15)中。     

Karyomorphology ofCrossostylis (Rhizophoraceae)

We present the first report on somatic chromosome numbers and morphology in eight of 13 recorded species ofCrossostylis, one of inland genera of Rhizophoraceae. The chromosome number ofCrossostylis is 2n=28 in all species examined; therefore, the genus hasx=14, a number which is the smallest and unknown elsewhere in the family. Based onCrossostylis raiateensis, we further present that 24 of 28 chromosomes at metaphase have centromeres at median position, and the remaining four at submedian or subterminal position. The chromosome morphology seems to imply thatCrossostylis might be a tetraploid with the original base numberx=7, but an extensive study in the other inland genera is needed to find such a small chromosome number.     

扇脉杓兰和无距虾脊兰的核型分析

为了解扇脉杓兰(Cypripedium japonicum Thunb.)和无距虾脊兰(Calanthe tsoongiana T. Tang et F. T. Wang)的核型,采用根尖压片法对扇脉杓兰和无距虾脊兰的染色体数目和核型进行了研究。结果表明,扇脉杓兰体细胞的染色体数为22,核型公式为2n=2x=22=16m+2sm+2st+2t,染色体相对长度组成为2n=22=2L+6M2+12M1+2S,核不对称系数为60.01%,核型分类为2B型;而无距虾脊兰体细胞的染色体数为40,核型公式为2n=2x=40=28m+10sm+2st,染色体相对长度组成为2n=40=8L+10M2+16M1+6S,核不对称系数为59.84%,核型分类为2B型;两者核型都较为对称。其中,无距虾脊兰的核型为首次报道。这为扇脉杓兰和无距虾脊兰的进化地位和种质保护提供了细胞学证据。     

CHROMOSOME NUMBERS IN THE COMPOSITAE: COLOMBIAN SPECIES

Chromosome numbers are presented for 76 species belonging to 35 genera of Compositae from Colombia. Thirty-nine species and three genera, Espeletia (x = 19), Steiractinia (x = 14), and Vasquezia (x = 19), are reported for the first time. New base numbers or chromosome series are recorded in Baccharis (B. nitida, n = 25), Calea (C. caracasana, n = 24), and Liabum (L. mega-cephalum, n = 10).     

Cytogenetic and molecular characterization of morphologically variable Sisyrinchium micranthum (Iridaceae) in southern Brazil

Sisyrinchium micranthum is native to southern Brazil and shows a wide range of morphological variation in this region. This study was performed to compare individuals of this species in terms of traits such as morphology, chromosome number, meiotic behaviour, pollen viability and genetic diversity using inter simple sequence repeat markers. First, general morphological aspects were evaluated and plants were classified into three morphological categories (CI, CII and CIII) to analyse the cytogenetic and genetic data. Three chromosome numbers corresponding to three ploidies were found: 2n = 2x = 16, 2n = 4x = 32 and 2n = 6x = 48. All morphological types showed regular meiotic behaviour and high meiotic index values and pollen viability. CII is the most frequent in southern Brazil. The most common haploid chromosome number, n = 8, has not been reported previously in this species. The percentage of polymorphic loci ranged from 6.45 to 61.29% for each accession. Molecular and cytogenetic analyses indicate that some S. micranthum accessions may have more than one ploidy. This study represents the first effort to characterize this taxonomically complex species based on cytogenetic and genetic aspects. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 169 , 350–364.     

Molecular and cytogenetic characterization of an Oryza officinalis–O. sativa chromosome 4 addition line and its progenies

The wild species Oryza officinalis Wall. ex Watt (2n = 24, CC) is a valuable genetic resource for rice (O. sativa L., 2n = 24, AA) breeding and genomics research. Genomic in situ hybridization (GISH) and molecular approaches were used to determine the nature and composition of the additional chromosome in a monosomic alien addition line (MAAL) of O. officinalis and its backcross progenies. The extra wild species chromosome in the MAAL (2n = 2x = 25) was a mosaic one, comprising of the long arm of chromosome 4 from O. officinalis and the short arm from O. sativa. Comparative analysis showed that O. sativa and O. officinalis shared high synteny of restriction fragment length polymorphism (RFLP) markers and low synteny of simple sequence repeat (SSR) markers. A DNA methylation alteration was revealed at C619 in the MAAL and progenies. Analysis of progenies of the MAAL indicated that introgression segments were small in size and introgression was not evenly distributed along the long arm. One recombination hot spot between C513 and RG177 was identified, which is in a gene-rich region.     

The phylogenetic position of Apterosperma (Theaceae) based on morphological and karyotype characters

Classifications of Theaceae have usually placed the endangered monotypic genus Apterosperma in tribe Schimeae (x=18), whereas recent molecular phylogenetic evidence supports its transfer to tribe Theeae (x=15). Molecular data have not resolved the phylogenetic position of Apterosperma within Theeae. We investigated the chromosome number and karyotype of Apterosperma in the context of molecular and morphological phylogenetic evidence to provide further insight into the placement of Apterosperma within Theaceae. The chromosome number and karyotype was found to be 2n = 30 = 26m + 4sm, consistent with the transfer of Apterosperma to tribe Theeae. When the chromosome data were incorporated into a data set of 46 other nonmolecular characters, Apterosperma was placed as the first-diverging lineage within the clade comprising tribe Theeae. This supports its placement based on molecular data. The low intrachromosomal asymmetry (type 1A) of Apterosperma, presumably ancestral for the family, is also consistent with this placement. Character optimization strongly supports a base chromosome number of x=15 for tribe Theeae. Because of variable and sometimes conflicting chromosome count reports of species in tribes Schimeae and Stewartieae, the base chromosome number of Theaceae could be either x=15 or 17.     

Comparative karyotype analysis in Haplopappus Cass. and Grindelia Willd. (Asteraceae) by double FISH with rRNA specific genes

The genera Grindelia Willd. and Haplopappus Cass. belong to the family Asteraceae - Astereae and are distributed in America and South America, respectively. Previous cytotaxonomic studies showed for South American species of Grindelia 2n=12 and for Haplopappus 2n=10 and 2n=12. Both Grindelia species (G. anethifolia, G. prunelloides), newly analyzed with molecular-cytological methods, exhibited symmetric karyotypes (AsI %=55.46 and 55.95) with metacentric chromosome sets (5m + 1m-sat) and 2n=12 chromosomes. The NOR was detected after fluorescence in situ hybridization (FISH) with 18/25S rDNA in the satellite chromosome 2. In contrast H. Happlopappus glutinosus, H. grindeloides and H. stolpii showed exclusively a higher asymmetric index (66.83%, 67.01% and 68.87%, respectively) with submetacentric chromosome sets (4sm + 1sm–sat). The sat-chromosomes 3 of H. glutinosus and H. grindelioides were both significantly different in their length from chromosomes 2 and 4. Furthermore in Grindelia the FISH with 5S rDNA could estimate signals in the short arms of chromosomes 3 or 4, that were not significantly differentiated in their length. Contrary to these findings in Grindelia, the position of 5S rDNA in Haplopappus was detected in the long arms of chromosome 1 (H. grindelioides and H. stolpii) and chromosome 2 (with two different loci) and chromosome 4 of H. glutinosus. The lengths of all measured chromosome arms with 5S rDNA were significantly different to those of the neighbours in the karyotypes. The two-color FISH of 5S and 18/25S rDNA had provided clear karyotypic markers for three (Haplopappus glutinosus) and two (H. grindelioides and H. stolpii) chromosomes. The number and position of rDNA signals were relatively highly conserved in the investigated five species without the double marked chromosome 2 of H. glutinosus, which shows an evolutionary dynamic of this 5S rRNA specific gene cluster. This investigation supports the assumption that the evolution of New World members of Grindelia and Haplopappus has not been accompanied by large karyotypic changes, but small chromosomal rearrangements have undoubtedly occurred (e.g. 5S rDNA localizations).     

Karyotype analysis in some species of Consolida (Ranunculaceae) from Iran

Consolida (dc .) S. F. Gray belongs to Ranunculaceae. The genus includes about 52 species worldwide. Here we report the diploid chromosome number and chromosome size and morphology for six Consolida species. For C. anthoroidea, C. leptocarpa, C. paradoxa and C. rugulosa the diploid chromosome number is reported for the first time. All investigated species have a diploid chromosome number of 2n = 2x = 16, except for C. persica having 2n = 2x = 14. The karyotypes of all six taxa are asymmetric, consisting of all four major chromosome types: metacentric, submetacentric, subtelocentric and telocentric chromosome type. However, considering the karyotype formula, all six species could be distinguished. In all taxa, metacentric chromosome pair 1 possesses a satellite. The only exception is C. rugulosa having an additional satellite positioned on metacentric chromosome pair 2. Karyotype data allow the separation of Aconitella from Consolida. Karyotype data plus morphological evidence support the reduction of C. paradoxa to formae level of C. rugulosa. (© 2013 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

我国入侵植物薇甘菊(菊科)的细胞学研究

为了解入侵植物薇甘菊(Mikania micrantha)的细胞学特性,研究了薇甘菊在我国广东东南部4居群和台湾2居群的染色体数目和染色体形态。结果表明,所有居群的薇甘菊染色体数目均为2n=36,第一对染色体为近中部着丝粒染色体,其长臂中部具有次缢痕,显著大于其余染色体。广东深圳大学和深圳水库居群的核型公式为2n=20m+14sm+2st;广东汕头、阳江和台湾屏东居群为2n=22m+12sm+2st;台湾宜兰居群为2n=18m+16sm+2st。此前有报道薇甘菊深圳大学居群的染色体数目为2n=38,推断为制片过程中第一对染色体从次缢痕处断裂而导致的计数错误。因此,薇甘菊虽存在非整倍性和多倍性变化,但在广东东南部和台湾的入侵居群中目前仅发现基于x=18的二倍体(2n=36),该种在这些地区的成功入侵与多倍性无关。     

Karyotype evolution in South American species of Hypochaeris (Asteraceae, Lactuceae)

The genus Hypochaeris (Asteraceae, Lactuceae) contains ten species in Europe, three in Asia, and approximately 50 in South America. Previous cytotaxonomic studies have shown two groups of taxa: (1) European species with different basic chromosome numbers and differentiated karyotypes, and (2) South American species with x=4 and uniform asymmetric and bimodal karyotypes. Karyotypic data are synthesized for South American species of Hypochaeris with new information for six Chilean species: H. acaulis, H. apargioides, H. palustris, H. spathulata, H. tenuifolia and H. thrincioides. Four main groups can be distinguished based on presence and localization of secondary constrictions – SCs (bearing Nucleolar Organizer Regions – NORs) on chromosomes 2 and 3, and 18S–25S and 5S rDNA loci number, localization, and activity. We propose karyotypic evolution of South American Hypochaeris (x=4) from H. maculata-like (x=5) European ancestors. The original South American karyotype would have possessed two SCs, one on the long arm of chromosome 2, and the other on the short arm of chromosome 3 (in terminal position). Further evolution would have involved inversion within the short arm of chromosome 3 and inactivation/loss of the SC on chromosome 2.     

A cytological study on Kelloggia (Rubiaceae), an intercontinental disjunct genus between eastern Asia and western North America

A cytological study was carried out for the first time on four populations of the only two species of Kelloggia (Rubiaceae), which occur disjunctly in eastern Asia (K. chinensis Franch.) and western North America (K. galioides Torr.). The consistent mitotic prophase chromosome condensation pattern and interphase nuclei type were determined for both species. The chromosome base number of the genus is suggested to be x=11. The karyotype of 2n=22=2x=16m+6sm was examined for both species. The karyotypical asymmetry of 1A and 2A was found in K. chinensis and K. galioides, respectively. According to the predominant evolutionary direction of karyotype asymmetry in angiosperms, K. galioides (2A) seems slightly more evolved than K. chinensis (1A). Our finding is consistent with the hypothesis of the Old World origin of Kelloggia based on molecular study.     

Genome size and chromosome number in the New Zealand species of Schoenus (Cyperaceae)

Schoenus (Cyperaceae) has holocentric chromosomes. Chromosome numbers were counted and nuclear DNA amounts were measured for all the New Zealand species of the genus. Chromosome numbers ranged from 2n = 8 to c. 2n = 90. Two chromosome races, with 2n = 28 and 2n = 56, were found in S. pauciflorus. Flow cytometry using propidium iodide‐stained nuclei was used to measure genome size. A 14.8‐fold variation in 2C DNA content was found, with values ranging from 1.33 to 19.71 pg/2C nucleus. Phylogenetic trees based on sequence variation in the internal transcribed spacer (ITS) region of the 45S ribosomal DNA locus were constructed using several phylogenetic models to reveal possible evolutionary relationships among the New Zealand Schoenus spp. and a sample of Australian Schoenus spp. Analysis revealed heterogeneity of chromosome number, size and DNA C value within clades. Meiosis in four species showed only bivalent formation at metaphase I. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 169 , 555–564.     

Karyoevelution on the genus Onosma L. (Boraginaceae): quantification of chromosome number heterogeneity

In this survey, chromosome counts of different species belonging to the genus Onosma are summarized and then karyological patterns available including frequency of cytotype occurrence, percentage of taxa with particular basic chromosome number and rate of polyploidy in the genus are evaluated. Quantitative parameters have been used to characterize chromosome number (CN) variation. In order to verify if variation patterns differ between three groups of Onosma, Index of CN Heterogeneity (ICNH) was quantified. In addition, meiotic chromosome numbers of 14 populations belonging to 11 species growing in Iran, namely Onosma araratica (2n = 2x = 16), O. asperrima (2n = 2x = 16), O. bulbotricha (2n = 2x = 18), O. kotschyi (2n = 2x = 16), O. microcarpa (2n = 2x = 16), O. nigricaulis (2n = 2x = 16), O. nervosa (2n = 2x = 16), O. obtusifolia (2n = 2x = 16), O. pachypoda (2n = 2x = 16), O. stenosiphon (2n = 2x = 20) and O. subsericea (2n = 2x = 16), were determined. With the exception of O. microcarpa and O. bulbotricha, all chromosome counts are reported for the first time. (© 2013 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Constitutive heterochromatin, 5S and 18S rDNA genes in Apareiodon sp. (Characiformes, Parodontidae) with a ZZ/ZW sex chromosome system

Karyotype, sex chromosome system and cytogenetics characteristics of an unidentified species of the genus Apareiodon originating from Piquiri River (Paraná State, Brazil) were investigated using differential staining techniques (C-banding and Ag-staining) and fluorescent in situ hybridization (FISH) with 5S and 18S rDNA probes. The diploid chromosome number was 2n = 54 with 25 pairs of meta- (m) to submetacentric (sm) and 2 pairs of subtelocentric (st) chromosomes. The major ribosomal rDNA sites as revealed by Ag-staining and FISH with 18S rDNA probe were found in distal region of longer arm of st chromosome pair 26, while minor 5S sites were observed in the interstitial sites on chromosome pairs 2 (smaller cluster) and 7 (larger one). The C-positive heterochromatin had pericentromeric and telomeric distribution. The heteromorphic sex chromosome system consisted of male ZZ (pair 21) and female middle-sized m/st Z/W chromosomes. The pericentric inversion of heterochromatinized short arm of ancestral Z followed by multiplication of heterochromatin segments is hypothesized for origin of W chromosome. The observed karyotype and chromosomal markers corresponded to those found in other species of the genus.