Further evidence for early sex chromosome differentiation of Anuran species

Chromosome banding and meiotic evidence show that XX/XY systems found in two Eupsophus species (Amphibia-Leptodactylidae) represent early stages of sex chromosome differentiation. Pair 14 is heteromorphic in E. migueli males and represents the heterochromosomes. In E. roseus this pair is metacentric and does not show heteromorphism. Paracentromeric constitutive heterochromatin is present in all chromosomes except in the E. migueli and E. roseus metacentric Y chromosomes. Constitutive heterochromatin loss is the structural modification responsible for Y chromosome differentiation. Pericentric inversions may have modified the morphology of the X chromosome of Eupsophus species.Departamento de Ciencias Ecológicas, Facultad de Ciencias, Universidad de ChileDepartamento de Biología Celular y Genética, Facultad de Medicina, Universidad de Chile     

白眉长臂猿(Hylobates hoolock leuconedys)的染色体研究

本文对两只雄性白眉长臂猿的染色体的C带、G带及Ag-NORs分布进行了较详细的分析,证实染色体数2n=38,并对该种的分类地位提出了一些新看法。     

The effect of heterochromatin on synapsis of the sex chromosomes of Peromyscus (Rodentia, Cricetidae)

The pairing behavior of the sex chromosomes in male and female individuals representing seven species of Peromyscus was analyzed by electron microscopy of silver-stained zygotene and pachytene configurations. Six species possess submetacentric or metacentric X chromosomes with heterochromatic short arms. Sex-chromosome pairing in these species is initiated during early pachynema at an interstitial position on the X and Y axes. Homologous synapsis then progresses in a unidirectional fashion towards the telomeres of the X short arm and the corresponding arm of the heterochromatic Y chromosome. The distinctive pattern of synaptic initiation allowed a late-synapsing bivalent in fetal oocytes to be tentatively identified as that of the X chromosomes. In contrast to the other species, Peromyscus megalops possesses an acrocentric X chromosome and a very small Y chromosome. Sex-chromosome pairing in this species is initiated at the proximal telomeric region during late zygonema, and then proceeds interstitially towards the distal end of the Y chromosome. These observations suggest that the presence of X short-arm heterochromatin and corresponding Y heterochromatin interferes with late-zygotene alignment of the pairing initiation sites, thereby delaying XY synaptic initiation until early pachynema. The pairing initiation sites are conserved in the vicinity of the X and Y centromeres in Peromyscus, and consequently the addition of heterochromatin during sex-chromosome evolution essentially displaces these sites to an interstitial position.     

Heterochromatin variation and sex chromosome polymorphism in Nesokia indica: a population study

Nesokia indica, the Indian mole rat, exhibits extensive variability (polymorphism) for the constitutive heterochromatin of the X and Y chromosomes. These polymorphic X and Y types range from a large metacentric chromosome to a small acrocentric one and occur in different frequencies in the population. On the assumption that there is random mating among individuals carrying these various X and Y chromosomes, the population shows Hardy-Weinberg proportions for the genotypes. However, notwithstanding the partial or total loss of constitutive heterochromatin of the X and Y chromosomes in a few individuals, its retention in most of the animals seems obligatory to the population at large. Hence, we suggest that the C-heterochromatin plays a "regulatory" role in the population dynamics of this species.     

Chromosome homology and heterochromatin in goat,sheep and ox studied by banding techniques

Peripheral blood lymphocyte metaphase chromosomes of three Bovoidean species have been studied using Quinacrine fluorescence and Giemsa banding techniques to give Q-, G-, and C-banding patterns. Q- and G-banding characteristics, coupled with chromosome length, enabled all of the chromosomes in each of the chromosome complements to be clearly distinguished, although some difficulties were encountered with the very smallest chromosomes. A comparison of G-banding patterns between the species revealed a remarkable degree of homology of banding patterns. Each of the 23 different acrocentric autosomes of the domestic sheep (2n=54) was represented by an identical chromosome in the goat (2n=60) and the arms of the 3 pairs of sheep metacentric autosomes were identical matches with the remaining 6 goat acrocentrics. A similar interspecies homology was evident for all but two of the autosomes in the ox (2n=60). This homology between sheep metacentric and goat acrocentric elements confirms a previously suggested Robertsonian variation. The close homology in G-banding patterns between these related species indicates that the banding patterns are evolutionarily conservative and may be a useful guide in assessing interspecific relationships. —The centromeric heterochromatin in the autosomes of the three species was found to show little or no Q-or G-staining, in contrast to the sex chromosomes. This lack of centromeric staining with the G-technique (ASG) contrasts markedly with results obtained with other mammalian species. However, with the C-banding technique these regions show a normal intense Giemsa stain and the C-bands in the sex chromosomes are inconspicuous. The amount of centromeric heterochromatin in the sheep metacentric chromosomes is considerable less than in the acrocentric autosomes or in a newly derived metacentric element discovered in a goat. It is suggested that the pale G-staining of the centromeric heterochromatin in these species might be related to the presence of G-Crich satellite DNA.     

New Molecular and Morphometric Evidence for the Validation of Eupsophus calcaratus and E. roseus (Anura: Leptodactylidae) in Chile

Molecular, morphometric, and distributional data are used to establish the taxonomic identity and differentiation of the South-American leptodactylid frogs Eupsophus calcaratus and E. roseus. Their geographical distribution in Chile suggests that the differentiation of these species occurred allopatrically. Molecular and morphometric data indicate a clear taxonomic identity of both taxa.     

Chromosome evolution in the erythrinid fish, Erythrinus erythrinus (Teleostei: Characiformes)

The genus Erythrinus belongs to the family Erythrinidae, a neotropical fish group. This genus contains only two described species, Erythrinus erythrinus being the most widely distributed in South America. Six samples of this species from five distinct Brazilian localities and one from Argentina were studied cytogenetically. Four groups were identified on the basis of their chromosomal features. Group A comprises three samples, all with 2n = 54 chromosomes, a very similar karyotypic structure, and the absence of chromosome differentiation between males and females. One sample bears up to four supernumerary microchromosomes, which look like 'double minute chromosomes' in appearance. Groups B-D comprise the three remaining samples, all sharing an X(1)X(1)X(2)X(2)/X(1)X(2)Y sex chromosome system. Group B shows 2n = 54/53 chromosomes in females and males, respectively, and also shows up to three supernumerary microchromosomes. Groups C and D show 2n=52/51 chromosomes in females and males, respectively, but differ in the number of metacentric, subtelocentric, and acrocentric chromosomes. In these three groups (B-D), the Y is a metacentric chromosome clearly identified as the largest in the complement. The present results offer clear evidence that local samples of E. erythrinus retain exclusive and fixed chromosomal features, indicating that this species may represent a species complex.     

白颊长臂猿染色体的研究

本文对我国云南南部的白须长臂猿（Ｈ．ｌｅｕｃｏｇｅｎｙｓ）染色体的Ｇ带、Ｃ带、晚复制带及Ａｇ－ＮＯＲｓ进行了较为详细的研究。它的２ｎ＝５２,核型公式为４４（Ｍ或ＳＭ）＋６（Ａ）,ＸＹ（Ｍ,Ａ）。Ｃ带表明一些染色体着丝点Ｃ带弱化;有的染色体出现插入的和端位的Ｃ带;Ｘ染色体两臂有端位Ｃ带,Ｙ染色体是Ｃ带阳性和晚复制的。Ａｇ－ＮＯＲｓ的数目,雌体有４个,雄体有５个,Ｙ染色体上具ＮＯＲ。本文对白颊长臂猿与其它长臂猿间的亲缘关系、核型进化的可能途径进行了讨论。     

Karyotypes and systematics of Asian high-mountain voles, genus Alticola (Rodentia, Arvicolidae). Results of Mongolian-German biological expeditions since 1962, No. 211.

Six species/subspecies of Asian high-mountain voles, genus Alticola, were studied cytogenetically via conventional staining and C- and G-banding. The karyotypes are very similar. The standard karyotype, as in A. strelzovi strelzovi, consists of 56 chromosomes. These are split into 25 acrocentric pairs, one large subtelocentric pair, one small metacentric pair, a large acrocentric X chromosome, and a small Y chromosome, which varies in shape. Constitutive heterochromatin is almost entirely restricted to small centromeric regions. A small submetacentric pair of autosomes in both subspecies of A. semicanus and a medium-sized Y chromosome in A. argentatus severtzovi are of importance in systematics. The data suggest that A. barakshin, A. semicanus, and A. argentatus are separate species.     

中国刺猬的染色体研究

本文采用全血培养和骨髓染色体制片法,对分布于我国南京市郊和济南市郊的刺猬染色体进行了组型、C-带、G-带和银染色的观察分析,并与东欧、西欧两种刺猬比较,它们之间的核型及带型差异显著;又将南京、济南及金清波(1985)报道的河南新乡三地分布的刺猬进行比较,它们的核型及带型也显示出一定差异,这种多态性在分类和进化上有一定的意义。     

Sister chromatid exchange points in the heterochromatin and euchromatin regions of Chinese hedgehog chromosomes

Summary The Chinese hedgehog has a diploid chromosome number of 48 in which there are eleven pairs of telo- or subtelocentric autosomes, twelve pairs of meta- or submetacentric autosomes, a metacentric X chromosome and a telocentric Y chromosome. The heterochromatin is almost completely distributed in five large distal segments of chromosomes nos. 9 to 12 and no. 18. There is no positive C-band in the centromeres of the chromosomes except for the X chromosome which has a small, weakly stained C-band in the centromere. In Chinese hedgehog cells 52.1% of SCEs are found at the junction between the euchromatin and the heterochromatin, 39.5% in the heterochromatin and 8.4% in the auchromatin. The SCE number per unit C-band is double the SCE number per unit euchromatin. The SCE rate in the heterochromatin or euchromatin regions is not proportional to their chromosome length and can be quite different between different pairs of the chromosomes. Our results indicate that there is a non-uniform distribution of the SCEs in the Chinese hedgehog cells.     

Cytogenetic analysis of Epicauta atomaria (Meloidae) and Palembus dermestoides (Tenebrionidae) with Xyp sex determination system using standard staining, C-bands, NOR and synaptonemal complex microspreading techniques

The mitotic and meiotic chromosomes of the beetles Epicauta atomaria (Meloidae) and Palembus dermestoides (Tenebrionidae) were analysed using standard staining, C-banding and silver impregnation techniques. We determine the diploid and haploid chromosome numbers, the sex determination system and describe the chromosomal morphology, the C-banding pattern and the chromosome(s) bearing NORs (nucleolar organizer regions). Both species shown 2n = 20 chromosomes, the chromosomal meioformula 9 + Xyp, and regular chromosome segregation during anaphases I and II. The chromosomes of E. atomaria are basically metacentric or submetacentric and P. dermestoides chromosomes are submetacentric or subtelocentric. In both beetles the constitutive heterochromatin is located in the pericentromeric region in all autosomes and in the Xp chromosome; additional C-bands were observed in telomeric region of the short arm in some autosomes in P. dermestoides. The yp chromosome did not show typical C-bands in these species. As for the synaptonemal complex, the nucleolar material is associated to the 7th bivalent in E. atomaria and 3rd and 7th bivalents in P. dermestoides. Strong silver impregnated material was observed in association with Xyp in light and electron microscopy preparations in these species and this material was interpreted to be related to nucleolar material.     

Chromosome plasticity in Ctenomys (Rodentia Octodontidae): chromosome 1 evolution and heterochromatin variation

A chromosome 1 (Cr1) pericentric inversion is described in six of seven species in the genus Ctenomys (tuco-tucos) from Uruguay. The inversion was inferred from G-band analyses of subtelocentric Cr1 hypothesised to be derived from the ancestral metacentric condition. Cr1 varies across species in heterochromatin amount and localisation including a metacentric chromosome without positive C-bands in C. torquatus, a subtelocentric chromosome with heterochromatic short arms in C. rionegrensis, and a subtelocentric chromosome negative after C-banding in five of the species analysed here. Pachytene chromosomes from C. rionegrensis, a species with the highest heterochromatin content, and C. torquatus, one of the species with the lowest heterochromatin content, were analysed in order to assess possible mechanisms of heterochromatin evolution. This analysis revealed the presence of three heterochromatic chromocenters in C. rionegrensis where bivalents converge, while in C. torquatus only one chromocenter was observed. In both species, highly repetitive DNA was observed, localised in chromocenters after “in situ” hybridisation. Heterochromatin associated protein M31 was localised in chromocenters of both species after immuno-detection. The spread of heterochromatin in Ctenomys chromosomes could be produced by chromatin exchanges at the chromocenter level. We propose the exchange of this DNA associated proteins between non-homologous chromosomes in pachytene to be the responsible for the spread of heterochromatin through the karyotypes of species like C. rionegrensis     

Chromosome evolution in Australian rodents

The chromosome complements of 188 specimens of 29 species of Australian murid rodents belonging to the subfamilies Pseudomyinae and Hydromyinae and the Uromys/Melomys group have been compared. At least one specimen of 18 different species was successfully C-banded. — The autosomal complements of many (9) diverse Pseudomyinae, one species of Melomys and one Hydromyinae proved to be identical, comprising 48 elements in the diploid set, the two smallest autosomal pairs of which are metacentric. No other karyotype is common to more than one species. From this we conclude that these three groups have been derived from a common ancestor which also possessed such a karyotype. The genus Zyzomys is exceptional since it possesses only 44 elements and lacks the two smallest metacentrics. — Karyotypic evolution within this apparently single phyletic line has been remarkably conservative, only three rearrangements being required to derive the most divergent karyotype. Moreover most of the observed rearrangements involve pericentric inversions and only one example of a fusion was found. Considerable differences in heterochromatin content, as determined by C-banding, occur between species however. Two species proved exceptional in this respect, namely Notomys cervinus and Uromys caudimaculatus. N. cervinus possesses numerous heterochromatic short arms. In U. caudimaculatus, there is a striking difference between northern and southern populations; in the former heterochromatin is present principally in the telomeric areas of the conventional A-chromosomes whereas in the latter it is found as separate supernumerary chromosomes. — In contrast to the autosomes, the X and Y chromosomes show high inter- and intra-specific variability in both size and morphology. All of this variability can be explained in terms of variation in heterochromatin content. Moreover the amount of heterochromatin in the X and Y chromosomes is highly correlated both within and between species. The Y chromosome of Uromys caudimaculatus is, however, distinctive in that it lacks C-banding.     

Habit differentiation and chromosome evolution inPleione (Orchidaceae)

Karyotypic and heterochromatin studies suggest a basic division of the orchid genusPleione into two groups, one represented by the clearly epiphytic species and the other including both species with terrestrial trends as well as those that are truly terrestrial. The epiphytic group possesses only (sub) metacentric chromosomes and is characterised by a considerable amount of terminal heterochromatin while the terrestrial group has some subtelocentric chromosomes and only small amounts of centromeric heterochromatin. It is concluded that a major phyletic split in the mode of chromosome change occurred during the transition from the epiphytic to the terrestrial habitat.     

Sex chromosome evolution in frogs—helpful insights from chromosome painting in the genus Engystomops

The differentiation of sex chromosomes is thought to be interrupted by relatively frequent sex chromosome turnover and/or occasional recombination between sex chromosomes (fountain-of-youth model) in some vertebrate groups as fishes, amphibians, and lizards. As a result, we observe the prevalence of homomorphic sex chromosomes in these groups. Here, we provide evidence for the loss of sex chromosome heteromorphism in the Amazonian frogs of the genus Engystomops, which harbors an intriguing history of sex chromosome evolution. In this species complex composed of two named species, two confirmed unnamed species, and up to three unconfirmed species, highly divergent karyotypes are present, and heteromorphic X and Y chromosomes were previously found in two species. We describe the karyotype of a lineage estimated to be the sister of all remaining Amazonian Engystomops (named Engystomops sp.) and perform chromosome painting techniques using one probe for the Y chromosome and one probe for the non-centromeric heterochromatic bands of the X chromosome of E. freibergi to compare three Engystomops karyotypes. The Y probe detected the Y chromosomes of E. freibergi and E. petersi and one homolog of chromosome pair 11 of Engystomops sp., suggesting their common evolutionary origin. The X probe showed no interspecific hybridization, revealing that X chromosome heterochromatin is strongly divergent among the studied species. In the light of the phylogenetic relationships, our data suggest that sex chromosome heteromorphism may have occurred early in the evolution of the Amazonian Engystomops and have been lost in two unnamed but confirmed candidate species.Subject terms: Cytogenetics, Evolutionary genetics     

Multiple sex chromosome system of X1X1X2X2/X1X2)Y type in lutjanid fish, Lutjanus quinquelineatus (Perciformes)

The karyotype and other chromosomal markers as revealed by C-banding and Ag-staining were studied in Lutjanus quinquelineatus and L. kasmira (Lutjanidae, Perciformes). While in latter species, the karyotype was invariably composed of 48 acrocentric chromosomes in both sexes, in L. quinquelineatus the female karyotype had exclusively 48 acrocentric chromosomes (2n = 48) but that of the male consisted of one large metacentric and 46 acrocentric chromosomes (2n = 47). The chromosomes in the first meiotic division in males showed 22 bivalents and one trivalent, which was formed by an end-to-end association and a chiasmatic association. Multiple sex chromosome system of X₁X₁X₂X₂/X₁X₂Y type resulting from single Robertsonian fusion between the original Y chromosome and an autosome was hypothesized to produce neo-Y sex chromosome. The multiple sex chromosome system of L. quinquelineatus appears to be at the early stage of the differentiation. The positive C-banded heterochromatin was situated exclusively in centromeric regions of all chromosomes in both species. Similarly, nucleolus organizer region sites were identified in the pericentromeric region of one middle-sized pair of chromosomes in both species. The cellular DNA contents were the same (3.3 pg) between the sexes and among this species and related species.     

The founder effect theory: quantitative variation and mdg-1 mobile element polymorphism in experimental populations of Drosophila melanogaster

The chromosomes of 14 specimens of the genus Reithrodon from three different localities of Argentina and two localities of Uruguay were studied using G-and C-banding techniques. Specimens of Uruguay showed a karyotype of 2n=28 chromosomes having a large metacentric X, and a telocentric Y chromosome. This karyotype is very similar to that recently described in a sample from southern Brazil, differing only in the nature of the Y chromosome, which is metacentric in the Brazilian form. All specimens from Argentina showed a 2n=34 karyotype, differing from the Brazilian karyotype by two centric fusions, an acquisition of chromosome material, and at least one pericentric inversion, and by the telocentric nature of both the X and the Y chromosomes. G-and C-banding suggest that the metacentric gonosomes in the Brazilian form resulted from a double autosomal-X-Y Robertsonian translocation. The Uruguayan cytotype is interpreted as derived from a hypothetical neo-X/Y₁Y₂ ancestral form by the secondary loss of the Y₁ chromosome. The karyotypic differences between the Brazilian-Uruguayan and the Argentinian forms afford evidence of species differentiation. It is proposed to assign the former to Reithrodon typicus, and the later to R. auritus.     

Chromosomal polymorphisms due to heterochromatin growth and pericentric inversions in white-bellied rat,Niviventer confucianus,from China

Different cytogenetic techniques were used to analyze the chromosomes of white-bellied rat, Niviventer confucianus from Mt. Tai and Jinan, Shandong Province and Ningshan, Shaanxi Province of China. Shandong populations have 2n = 46 chromosomes with 4 metacentric, 2 subtelocentric, 16 telocentric pairs of autosomes and the submetacentric X and telocentric Y. The chromosomal arm number (NF) of the two populations was 56. Shaanxi population has 2n = 46 chromosomes with 4 metacentric, 1 submetacentric, 1 subtelocentric and 16 telocentric pairs of autosomes and the submetacentric X and telocentric Y. The karyotype of Ningshan population showed NF = 58. As the result of the comparison of C- and G-banding patterns, and compare with other species in the genus Niviventer, we suppose that the chromosomal evolution of Niviventer involved in pericentric inversion and heterochromatin growth. The submetacentric chromosomes of Shaanxi population would be originated from the growth of heterochromatin of the subtelocentric chromosome of Shandong population.     

Evidence for a highly differentiated sex chromosome heteromorphism in the salamander Necturus maculosus (Rafinesque)

The mitotic chromosomes of the neotenic (sensu Gould, 1977, and Alberch et al., 1979) salamander Necturus maculosus (Rafinesque) have been examined using a C-band technique to demonstrate the distribution of heterochromatin. The C-banded mitotic chromosomes provide evidence of a highly differentiated XY male/XX female sex chromosome heteromorphism, in which the X and Y chromosomes differ greatly in size and morphology, and in the amount and distribution of C-band heterochromatin. The X chromosome represents one of the largest biarmed chromosomes in the karyotype and is indistinguishable from similar sized autosomes on the basis of C-band heterochromatin. The Y chromosome, on the other hand, is diminutive, morphologically distinct from all other chromosomes of the karyotype, and is composed almost entirely of C-band heterochromatin. The discovery of an X/Y chromosome heteromorphism in this species is consistent with the observation by King (1912) of a heteromorphic spermatogenic bivalent. Karyological and phylogenetic implications are discussed.