Karyotype analysis and chromosome evolution in South American species of Lathyrus (Leguminosae)

The karyotypes of 10 species and one variety of South American Lathyrus were determined and compared with those obtained of five entities from the Northern Hemisphere. Although all the species have a chromosome number of 2n = 14, they could be differentiated by their karyotype formula and quantitative parameters of the karyotypes. Phenetic distance and principal component analysis showed that in spite of the differences observed among entities, they can be grouped in clusters that coincide with the taxonomic sections established by F. K. Kupicha and with the life cycle of the species. South American species form a homogeneous group and can be distinguished by the presence of a subtelocentric pair, which has a macrosatellite in the long arm, and the lack of a short metacentric pair characteristic of most species of the Northern Hemisphere. From an evolutionary point of view, variation in total chromosome length without major changes in the karyotype formula suggests that changes in the amounts of genomic DNA are proportional to the relative length of each chromosome arm and that species of Notolathyrus evolved in a concerted fashion. Variation in genome size, however, is congruent with morphological variation of some reproductive organs as well as with the life cycle and minimum generation time, as predicted by the nucleotype hypothesis.     

Karyotype analysis and chromosome evolution in southernmost South American species of Crotalaria (Leguminosae)

Karyotypes of 18 populations belonging to eight of the southernmost species of South American Crotalaria were examined. All had x  = 8, except C. incana which had x  = 7. Diploid, tetraploid and octoploid species were found. The octoploid is a new record and is the highest ploidy level detected so far in the genus. Species within the same ploidy level could be distinguished by quantitative karyotype parameters and by the size and position of the satellites. From an evolutionary point of view, our results support a decrease of the basic chromosome number by dysploidy. Polyploidy is one of the more important mechanisms of chromosome change, even though some structural rearrangements may explain variations of the karyotype formula. Gigas effects, due to polyploidy, were evident in the length of the stomatal guard cells and in pollen size, but were less clear in seed weight. Nucleotype effects, caused by the DNA quantity in a nucleus, were also evident for pollen and stomatal guard cell size and seed weight, although only in diploids with x  = 8 for the latter character. Chromosome data are further discussed in the light of the infrageneric systematic arrangement.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 150 , 329–341.     

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.     

Karyotypes,heterochromatin distribution and rDNA patterns in South American Grindelia (Asteraceae)

Grindelia is a genus with a complex evolutionary history with reticulate evolution. We studied the karyotype, fluorescent banding, and fluorescence in situ hybridization (FISH) using 18–5.8–26 S and 5 S ribosomal DNA probes to survey karyotypic diversity of South American Grindelia species. Chromosome basic numbers were x = 6 (with several ploidy levels: 2x, 4x, and 6x). All the Grindelia studied conserved the patterns of CG-rich heterochromatin and 18-5.8-26 S rDNA. The third m sat-chromosome pair was homeologous in Grindelia. Chromosome variation, although not always large, accompanied the evolutionary divergence of the taxa studied. The Grindelia studied formed two species groups: (1) G. globularifolia and G. pulchella var. pulchella, (2) G. buphthalmoides, G. cabrerae var. alatocarpa and var. cabrerae, G. chiloensis, G. orientalis, and G. prostrata. These groups do not show any morphological affinities and their phylogenetic relationships are not clearly resolved, suggesting that these groups have recently diverged.     

Karyotype diversification and evolution in diploid and polyploid South American Hypochaeris (Asteraceae) inferred from rDNA localization and genetic fingerprint data

Background and Aims: Changes in chromosome structure and number play an importantrole in plant evolution. A system well-suited to studying differentmodes of chromosome evolution is the genus Hypochaeris (Asteraceae)with its centre of species' diversity in South America. AllSouth American species uniformly have a chromosome base numberof x = 4 combined with variation in rDNA number and distribution,and a high frequency of polyploidy. The aim of this paper isto assess directions and mechanisms of karyotype evolution inSouth American species by interpreting both newly obtained andprevious data concerning rDNA localization in a phylogeneticcontext. Methods: Eleven Hypochaeris species from 18 populations were studiedusing fluorescence in situ hybridization (FISH) with 35S and5S rDNA probes. A phylogenetic framework was established fromneighbour-net analysis of amplified fragment length polymorphism(AFLP) fingerprint data. Key Results: A single 5S rDNA locus is invariably found on the short armof chromosome 2. Using 35S rDNA loci, based on number (one ortwo) and localization (interstitial on the long arm of chromosome2, but sometimes lacking, and terminal or interstitial on theshort arm of chromosome 3, only very rarely lacking), sevenkaryotype groups can be distinguished; five of these includepolyploids. Karyotype groups with more than one species do notform monophyletic groups. Conclusions: Early evolution of Hypochaeris in South America was characterizedby considerable karyotype differentiation resulting from independentderivations from an ancestral karyotype. There was marked diversificationwith respect to the position and evolution of the 35S rDNA locuson chromosome 3, probably involving inversions and/or transpositions,and on chromosome 2 (rarely 3) concerning inactivation and loss.Among these different karyotype assemblages, the apargioidesgroup and its derivatives constitute by far the majority ofspecies.     

DNA content in South American endemic species of Lathyrus

The genome size was surveyed in 13 Notolathyrus species endemic to South America by flow cytometry and analyzed in an evolutionary and biogeographic context. A DNA content variation of 1.7-fold was registered, and four groups of species with different DNA content were determined. Although, the 2C values were correlated with the total chromosome length and intrachromosomal asymmetry index (A₁), the karyotype formula remained almost constant. The conservation of the karyotype formula is in agreement with proportional changes of DNA in the chromosome arms. Species with annual life cycle and shorter generation time had the lowest DNA content and the data suggest that changes in DNA content involved reductions of genome size in the perennial to annual transitions. The variation of 2C values was correlated with precipitation of the coldest quarter and, to some extent, with altitude. Additional correlations with other variables were observed when the species were analyzed separately according to the biogeographic regions. In general, the species with higher DNA content were found in more stable environments. The bulk of evidence suggests that changes on genome size would have been one of the most important mechanisms that drove or accompanied the diversification of Notolathyrus species.     

Karyotype analysis in South American species of Myrtaceae

In Myrtaceae (Myrteae), the diploid chromosome number 2 n  = 2 x  = 22 is the most common, although variations of ploidy level occur, with some triploid (2 n  = 3 x  = 33) and tetraploid (2 n  = 4 x  = 44) records. Karyotype details in this group are scarce because the chromosomes are small (< 2 μm). In this work, we carried out a karyotypic analysis of 15 species of Myrtaceae grouped in different subtribes and genera. Measurements of chromosome length (long arm, L ; short arm, S ) were taken and several karyotypic parameters were calculated for each species. The karyotypes in fleshy-fruited taxa (Myrteae) were more varied than in the other previously analysed dry-fruited group ( Eucalyptus , Eucalypteae), in which the chromosomes were exclusively metacentric.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society, 2007, 155 , 571–580.     

Chromosome banding in Amphibia. XXV. Karyotype evolution and heterochromatin characterization in Australian Mixophyes (Anura,Myobatrachidae)

The mitotic chromosomes of the Australian ground frogs Mixophyes fasciolatus and M. schevilli were analyzed by means of banding techniques and restriction endonuclease digestions. Chromosomal differentiation in these two species occurred exclusively by considerable changes in the amount of telomeric and centromeric heterochromatin, whereas the sizes and locations of interstitial heterochromatic regions, the sizes of all euchromatic segments as well as the positions of centromeres remained nearly identical during karyotype evolution. The major heterochromatic regions in the karyotypes of M. fasciolatus and M. schevilli amount to 30.2% and 20.7%, respectively. They consist of AT base pair-rich repetitive DNA sequences that are brightly labeled by AT-specific fluorochromes and display quenched fluorescence after staining with GC-specific fluorochromes. The heterochromatic regions can be differentiated by treatment of metaphase chromosomes and interphase cell nuclei with various restriction enzymes which either disclose the complete set of C-band patterns in the karyotypes of both species, or else reveal several subsets of these C-bands.     

Karyotype analysis of the southernmost South American species of Stemodia (Scrophulariaceae)

Abstract

The aim of the present study was to examine the somatic chromosomes of seven species of Stemodia (Scrophulariaceae). The karyotypes of Stemodia ericifolia (2n = 22 = 20m + 2 sm), S. hassleriana (2n = 22 = 20m + 2sm), S. hyptoides (2n = 22 = 20m + 2sm), S. hyptoides (2n = 44 = 40m + 4sm), S. lobelioides (2n = 44 = 40m + 4sm) and S. stricta (2n = 22 = 20m + 2sm) were analyzed for the first time. All the species studied showed a predominance of metacentric chromosomes and a lower proportion of submetacentric pairs. The chromosomes in all the species were found to be small with a mean chromosome length of 1.42 μm, varying from 0.77 μm in S. hyptoides (2n = 66) to 2.10 μm in S. lanceolata. The differences in the asymmetry of the karyotypes were small, for which it is possible to assume that the great diversification of the genus has been accompanied by very small changes in the structure of the chromosomes.     

Karyotype studies in South American species of Solanum subgen. Leptostemonum (Solanaceae)

Mitotic chromosome numbers and karyotypes of 13 South American species (12 native and one naturalized) from four sections of SOLANUM subgen. LEPTOSTEMONUM were studied. Chromosome numbers of S. ACERIFOLIUM, S. AENICTUM, S. CONDITUM, S. CONSIMILE, S. INCARCERATUM, and S. PLATENSE are reported for the first time. The number 2n = 24 was found in most species, while 2n = 22 was found in S. MAMMOSUM and S. PLATENSE. The latter is the second SOLANUM with this unusual number. Satellites are always present and were visible in more than 50 % of the cells studied. Karyotypes are symmetrical: M and SM chromosomes are common, whereas ST chromosomes are rare. The karyotypes of S. AENICTUM, S. MAMMOSUM, and S. PANICULATUM are comparatively asymmetrical. Species can be distinguished by a combination of chromosome number, karyotype formulae, karyotype length, the position of satellites in a particular chromosome pair, and asymmetry indices. The phenogram obtained does not reflect the sectional arrangements or the systematic affinities of the species studied. In sect. ACANTHOPHORA, increased asymmetry is associated with derived characters (strong andromonoecy, winged seeds, mammiform fruits). Diversification in the subgenus is suggested to be related to visible chromosome rearrangements and cumulative, cryptic structural changes may have also played a relevant evolutionary role.     

Karyotype relationships among four South American species of Urvillea (Sapindaceae: Paullinieae)

A cytogenetic study was conducted on four species of the genus Urvillea (Sapindaceae, Paullinieae): U. chacoensis Hunz., U. filipes Radlk. and U. ulmacea Kunth of the Urvillea section and U. laevis Radlk. of the Stenelytron section. The chromosome numbers in U. chacoensis (2n = 22) and U. laevis (2n = 24) were confirmed, and new chromosome numbers are reported for U. filipes with 2n = 22 and U. ulmacea with 2n = 88. Additionally, data on interphase nuclear structure, chromosome banding patterns (C-Giemsa and C-CMA₃/DAPI) and FISH with rDNA probes are also presented. The distribution of AT- and GC-rich regions and the physical mapping of ribosomal genes (45S and 5S rDNA sites) were established for the first time in these Urvillea species. Sections of Urvillea are cytogenetically differentiated according to basic chromosome number, where x = 11 in the section Urvillea and x = 12 in the section Stenelytron. This first section displayed an important karyotypic feature, the occurrence of large AT- and GC-rich bands at terminal chromosomal regions. The Urvillea section showed polyploidy and its species were differentiated by their banding patterns. Urvillea chacoensis showed several terminal AT-rich bands, while terminal AT- and GC-rich bands were both found in U. ulmacea. However, the section Stenelytron did not exhibit this banding pattern. The 45S rDNA sites appeared always associated with GC-rich regions and they were numerically variable among species, being located or not the same chromosome 5S rDNA sites. Variation in the repetitive DNA distribution and their role in karyotype differentiation among these Urvillea species are discussed.     

Karyotype and mitochondrial 16S gene characterizations in seven South American Cichlasomatini species (Perciformes, Cichlidae)

The family Cichlidae constitutes most of the freshwater fish fauna of South America; its taxonomy is at present mainly based on morphological characters. Here, relationships among seven Cichlasomatini species have been investigated by studying their karyotype structure and by sequencing a 520 bp fragment of the mitochondrial 16S gene. Molecular data sets point to a high affinity of Cichlasoma amazonarum with Aequidens sensu stricto group, in particular with Aequidens tetramerus . Aequidens never form a single monophyletic clade: molecular trees group together ' Aequidens pulcher ' and ' Aequidens rivulatus ' and suggest their close relationship with Bujurquina and Laetacara , rather than with the A. sensu stricto group. Both molecular and karyotypic data confirm that Cleithracara maronii belongs to a distinct clade, thus supporting its generic differentiation based on morphological characters. Chromosome number, karyotype structure and molecular data suggest that Laetacara dorsigera is related to Bujurquina vittata and confirm their generic level of differentiation. From a cytotaxonomic point of view, a karyotype of 2 n  = 48 with most acrocentric or subacrocentric chromosomes could be the ancestral one from which the others might have derived.     

First chromosome number determinations in south-eastern South American species of Lupinus L. (Leguminosae)

Chromosome numbers are presented for the first time for 30 accessions of nine south-eastern South American Lupinus species. Chromosome numbers of 2 n = 32 and 34 were found for L. bracteolaris (three out of five accessions with 2 n = 32) and L. linearis (two out of three accessions with 2 n = 32), and of 2 n = 36 for L. gibertianus , L. lanatus, L. magnistipulatus , L. multiflorus , L. rubriflorus , L. reitzii and L. uleanus . All the South American species examined have relatively low chromosome numbers when compared with most of the Old World and North American species. Our results, where 2 n = 36 is the rule, are in sharp contrast to the data for North American Lupinus species and reveal the following: (1) low chromosome numbers are the rule, at least in the southern part of eastern South America; (2) cytologically, the eastern South American species form a group distinct from the North American taxa; (3) high levels of polyploidy have not played as important a role in evolution and speciation in eastern South America as in North America; (4) the predominance of low chromosome numbers in eastern South American species and the existence of similar numbers in two of the six rough-seeded Old World species support the hypothesis that in the evolution of the genus the eastern South American species branched off first, followed by the rough-seeded group.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Socety , 2002, 139 , 395–400.     

Karyotype analysis and physical mapping of 45S rDNA in eight species of Sophora,Robinia, and Amorpha

The karyotype analysis and physical locations of 45S rDNA were carried out by means of fluorescence in situ hybridization in three species,and two forms of Sophora,two species of Robina,and one species of Amorpha.S.japonica L.,S.japonica L.f.oligophylla Franch.,S.japonica L.f.pendula Loud.,and S.xanthantha C.Y.Ma.are all tetraploids with 2n=28.There were four 45S rDNA sites in pericentromeric regions of two Pairs of chromosomes in each of them.S.rubriflora Tsoong.is a triploid with 2n=21,and three sites were located in each satellite of group 5 chromosomes.In R.pseudoacacia L.(2n=2x=22),we examined four intensive signals in telomeric regions of two pairs of satellite chromosomes.In R.hispida L.(2n=2x=30),there were four other signals in centromeric regions besides those like in R.pseudoacacia.Amorpha fruticosa L.has most chromosomes(2n=40)among the eight materials,however,there were only six 45S rDNA loci and they laid in centromeric regions,and satellites of three pairs of chromosomes.45S rDNA is a valuable chromosomal landmark in karyotype analysis.The distribution and genomic organization Of rDNA in the three genera were also discussed.     

Pollen morphology of Lathyrus (Leguminosae) taxa belonging to Lathyrus, Orobastrum and Cicercula sections from Turkey

The pollen morphology of 20 wild taxa belonging to Lathyrus (Syn: Eulathyrus), Orobastrum (Taub.) Boiss. and Cicercula (Medic.) Gren. & Godr. sections of Lathyrus L. grown in Turkey (L. rotundifolius Wild. subsp. miniatus (Bieb. ex Steven) P.H. Davis, L. grandiflorus Sibth. & Sm., L. saxatilis (Vent.) Vis., L. vinealis Boiss. & No?, L. inconspicuus L. var. inconspicuus, L. inconspicuus var. stenophyllus (Boiss.) Rech. f., L. tauricola P.H. Davis, L. woronowii Bornm., L. hierosolymitanus Boiss., L. cassius Boiss., L. gorgoni Parl. var. gorgoni, L. pseudo-cicera Pamp., L. sativus L., L. blepharicarpus Boiss., L. stenophyllus Boiss. & Heldr., L. belinensis Maxted & Goyder, L. phaselitanus Hub.-Mor. & P.H.Davis, L. chrysanthus Boiss., L. chloranthus Boiss., and L. trachycarpus (Boiss.) Boiss were examined by light microscopy (LM) and scanning electron microscopy (SEM) in this study. The pollen grains were 3-zonocolporate, spheroidal, subprolate, and prolate (P/E?=?0.99–1.48) types, and were medium in size (equatorial view: rectangular or elliptical-obtuse-convex; polar view: circular, triangular and quinquangular-obtuse-convex). The smallest pollen grains belonged to L. tauricola (P?=?30.94/E?=?31.20) and the largest to L. grandiflorus (P?=?50.60/E?=?36.40). The ornamentation was reticulate and reticulate-granulate in the mesocolpium, and usually psilate in the apocolpium. Some photographs included in this work were taken using both LM and SEM.     

Chromosomal Diversity and Karyotype Evolution in South American Macaws (Psittaciformes,Psittacidae)

Most species of macaws, which represent the largest species of Neotropical Psittacidae, characterized by their long tails and exuberant colours, are endangered, mainly because of hunting, illegal trade and habitat destruction. Long tailed species seem to represent a monophyletic group within Psittacidae, supported by cytogenetic data. Hence, these species show karyotypes with predominance of biarmed macrochromosomes, in contrast to short tailed species, with a predominance of acro/telocentric macrochromosomes. Because of their similar karyotypes, it has been proposed that inversions and translocations may be the main types of rearrangements occurring during the evolution of this group. However, only one species of macaw, Ara macao, that has had its genome sequenced was analyzed by means of molecular cytogenetics. Hence, in order to verify the rearrangements, we analyzed the karyotype of two species of macaws, Ara chloropterus and Anodorhynchus hyacinthinus, using cross-species chromosome painting with two different sets of probes from chicken and white hawk. Both intra- and interchromosomal rearrangements were observed. Chicken probes revealed the occurrence of fusions, fissions and inversions in both species, while the probes from white hawk determined the correct breakpoints or chromosome segments involved in the rearrangements. Some of these rearrangements were common for both species of macaws (fission of GGA1 and fusions of GGA1p/GGA4q, GGA6/GGA7 and GGA8/GGA9), while the fissions of GGA 2 and 4p were found only in A. chloropterus. These results confirm that despite apparent chromosomal similarity, macaws have very diverse karyotypes, which differ from each other not only by inversions and translocations as postulated before, but also by fissions and fusions.     

Karyotype characterization of five Brazilian species of Echinodorus (Alismataceae) with chromosomal banding and 45S rDNA FISH

The karyotypes of five Brazilian species of Echinodorus; E. bolivianus, E. grandiflorus, E. longipetalus, E. macrophyllus and E. tenellus (Alismataceae); were studied using C-banding, CMA₃/DAPI banding and fluorescence in situ hybridization with a 45S rDNA probe. There were few differences in the G-C rich regions of the five species, but marked differences were seen in the number and position of C-bands, A-T rich regions and 45S rDNA sites. Overall, these characteristics were species-specific, with the exception of E. bolivianus and E. tenellus, which were highly similar in all of the karyotypic characteristics studied.     

Intra- and interspecific variation in genome size in Lathyrus (Leguminosae)

2C nuclear DNA amounts for 24 species of Lathyrus (section Lathyrus ) were determined using flow cytometry. A greater than two-fold variation was observed, ranging from 10.2 pg in L. basalticus to 24.2 pg in L. latifolius. In general, the perennial species had greater DNA amounts than the annuals. Significant intraspecific variation was observed in five species of Lathyrus (from 10.1% in L. annuus to 28% in L. tingitanus). A positive correlation was observed between DNA values obtained by flow cytometry and those previously determined by microdensitometry. Finally, the distribution of DNA amounts in species within section Lathyrus appears to be continuous.     

Karyotype structure, supernumerary chromosomes and heterochromatin distribution suggest a pathway of karyotype evolution in Dactylorhiza (Orchidaceae)

The relationships between Dactylorhiza romana and D. saccifera from southern Italy were analysed. These two species, both with 2 n = 2 x = 40 chromosomes and belonging to different sections of the genus, were distinguishable on the basis of karyotype structure and heterochromatin amounts and distribution. Their C-banded karyotypes differed considerably. D. saccifera showed most chromosomes with banded regions in the short arms, whereas in D. romana the bands were located mostly at telomeric regions of longer arms. Several individuals of D. romana had one or two very large heterochromatic supernumerary chromosomes. Based on evidence resulting from karyotype structure and heterochromatin distribution in the two species and on the genetic distances derived from the comparison of ITS sequences, it is suggested that D. romana represents a primitive form with respect to D. saccifera and is a possible intermediate step in the evolution of the genus Dactylorhiza from the 42-chromosome Orchis group. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 138 , 85–91.     

Evidence for the hybrid origin of Lathyrus plitmannii (Leguminosae)

ABSTRACT

The hypothesis that Lathyrus plitmannii may be of hybrid origin was tested on a morphological data set, derived from measurements of 50 herbarium specimens, by employing the computer program HYWIN. Results showed that all the specimens of L. plitmannii were putative hybrids and that the highest ranked pair of specimens that best matches the morphologies of its parents was L. gorgonei and L. pseudocicera. Examination of the distribution patterns of the three species shows that L. plitmannii is confined to regions common to both putative parents and that hybrids could be formed, assuming that genetic barriers were overcome. Additional supporting evidence includes overlaps in the edaphic requirements and the habitats of the three species. Lathyrus plitmannii is recognised as a distinct species that should be regarded as derivative of hybridisation between L. gorgonei and L. pseudocicera.