Unusual occurrence of a ZZ/ZW sex-chromosome system and supernumerary chromosomes in Characidium cf. fasciatum (Pisces, Characiformes, Characidiinae)

Individuals of two populations of the fish Characidium cf. fasciatum were cytogenetically studied and showed a basic diploid number of 50 chromosomes. Some fishes were found to have 51 to 54 chromosomes due to the presence of one to four small subtelocentric/acrocentric supernumerary chromosomes. When analyzed by conventional Giemsa staining, male and female specimens of C. cf. fasciatum from the Quinta stream and Pardo River presented the same basic karyotypic macro- and microstructure, consisting of 32 metacentric and 18 submetacentric chromosomes. Ag-NORs were terminally located on the long arms of two submetacentric chromosome pairs. Constitutive heterochromatin was identified by C-banding as small pericentromeric blocks in the majority of the chromosomes, and B-chromosomes were found to be heterochromatic. The occurrence of one totally heterochromatic submetacentric chromosome restricted to females and considered as an unusual feature in fish karyotypes led to the identification of a ZZ/ZW sex-chromosome system. The implications of chromosomic differentiation observed in the genus Characidium are discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

A chromosomal study on Greek populations of the genusApodemus (Rodentia, Murinae) reveals new data on B chromosome distribution

During this study, 94 specimens (51 males, 43 females) of the wood mouseApodemus sylvaticus (Linnaeus, 1758), the yellow-necked mouseA. flavicollis (Melchior, 1834) and the western broad-toothed mouseA. epimelas (Nehring, 1902) from 20 localities of Greece were karyologically examined. The first two species were found to be widely distributed and their otherwise very similar karyotype (2n=48, FN=48) could be clearly distinguished, based on C-banding pattern. The third species appeared to have a more limited geographical distribution and its karyotype was distinguished from that of the previous two species, since it contained two pairs of small metacentric autosomes (2n=48, FN=52). The chromosomal study further revealed that nine individuals ofA. flavicollis possessed supernumerary B chromosomes (2n=49–50, FN=49–50). Meiotic chromosome preparations revealed that in contrast to sex chromosomes and autosomes, B chromosomes do not participate in bivalent formation. On the other hand, no supernumerary chromosomes were found in the studiedA. sylvaticus andA. epimelas material.     

Studies on metatherian sex chromosomes. IX. Sex chromosomes of the greater glider (Marsupialia: Petauridae)

The greater glider, currently but incorrectly known as Schoinobates volans, is widely distributed in forested regions in eastern Australia. All animals studied from six different localities had 20 autosomes but there were four chromosomally distinct populations. At Royal National Park, N.S.W., all female greater gliders studied had 22 chromosomes including two large submetacentric X chromosomes with subterminal secondary constrictions in their longer arms. This form of X chromosome occurred also at Bondo State Forest, Myall Lakes and Coff's Harbour, N.S.W., and at Eidsvold, Qld. At Coomooboolaroo, Qld, the X chromosome was also a large submetacentric but a secondary constriction occurred in the shorter arm. Two chromosomally distinct types apparently occur in Royal National Park, one with XY males as in all other populations, and one with XY1Y2 males. Y or Y1, but not Y2, chromosomes were eliminated from the bone marrow in all populations but were present in spermatogonia, primary spermatocytes and cultured fibroblasts. Animals from Bondo State Forest had three or more acrocentric or metacentric supernumerary chromosomes.     

A comparative study of mitotic and meiotic chromosomes of the Montandon's newt,Triturus montandoni (Urodela: Salamandridae) from Poland and Rumania

A karyological analysis was carried out on two populations ofTriturus montandoni, one from Poland and another from Rumania. For both samples, morphometric characteristics and C-banding pattern of mitotic chromosomes are provided. Only slight differences between specimens from two geographic localities were found. Data on chiasma frequency and distribution are presented for male meiosis. No sex-related heteromorphism was found and for none of the chromosomal arms was a consistent absence of chiasmata recorded. There was a relatively high proportion of spermatocyte metaphases I with chiasmata on both arms of all chromosomes in all specimens studied. It is concluded that there are no well-defined sex chromosomes in the chromosomal complement of the maleT. montandoni. The findings are compared with previous studies on chromosome morphology, C-banding pattern, and meiosis in closely related species,T. vulgaris andT. helveticus.     

Comportement des chromosomes surnuméraires et relation avec le taux de mortalité dans une population africaine de Locusta migratoria migratorioides

Transmission of supernumerary chromosomes is studied in adults and embryos of an African population of Locusta migratoria migratorioides; 34% of the animals show one or two B chromosomes (24% one, 10% two). This percentage is the same in both sexes. During mitosis, B chromosomes are very stable. At meiosis, in some cases they show pairing, in other cases they enter as univalents. The eventuality of synapsis or association is discussed. These two kinds of behaviour could not be explained by the presence of two different B chromosomes as is shown from the study of parthenogenetic progeny. Repartition of B chromosomes in the progeny is different depending on whether the male or the female parent supplies them; so the behaviour of these supernumerary should be conditioned, not by their own structure, but by a connection with the cell. — One B chromosome apparently is neutral. When there are two B chromosomes, in case of synapsis, the number of animals with two B in the progeny is slightly lower than previous; in case of asynapsis, there is a higher lethality of individuals and oocytes with two B chromosomes.     

Karyotypes of Rana tagoi Okada with diploid number 28 in the Chausu Mountains of the Minamishinshu district of Nagano Prefecture, Japan (Anura: Ranidae)

Karyotypes of Tago's brown frog Rana tagoi from the Chausu mountains in Minamishinshu of Nagano Prefecture were examined by conventional Giemsa staining, C-banding and late replication (LR)-banding. Chromosome number was 2n = 28 in all cases. The 28 chromosomes consisted of four pairs (1-4) of large biarmed chromosomes, two pairs (5-6) of telocentric chromosomes and eight pairs (7-14) of small biarmed chromosomes. Chromosome pair 11 had a secondary constriction on the long arm. In females, the C-band on the long arm of chromosome pair 6 was detected in both homologs, but was absent from the arms of the homologs of chromosome pairs 5 and 9. In males, C-bands were found in the long arms of both homologs of chromosome pairs 5 and 6, were present only in one homolog of chromosome pair 5 for certain male specimens and found in only one homolog of chromosome pair 9. Specimens of R. tagoi (2n = 28) should thus have two pairs of telocentric chromosomes to provide the same number of chromosome arms, these originating quite likely from chromosome pair 1 in the 26-chromosome specimens by centric fission. Heteromorphic sex chromosomes of the XX-XY type in R. tagoi (2n = 28) in the Chausu mountains were identified. Karyotypes of tail-tip cells from a hybrid tadpole between female R. tagoi (2n = 26) from the Hinohara village in Tokyo and male R. tagoi (2n = 28) from the Chausu mountain population were examined by squash preparation. Chromosome number was 2n = 27 in all tadpoles. The 27 chromosomes consisted of one chromosome set of R. tagoi (2n = 28) and one of R. tagoi (2n = 26).     

Distortional meiotic segregation of a supernumerary chromosome producing differential frequencies in the sexes in the short-horned grasshopper Phaulacridium vittatum

Samples of males reveal the presence of a large, mitotically stable supernumerary chromosome in five populations examined. A stable frequency (11.22±0.3%) of male carriers has been observed over nine years in one population. The supernumerary follows closely the condensation cycle and behaviour of the sex chromosome in spermatogonial mitosis and meiosis. A structure simulating a terminalized chiasma frequently joins the precocious sex and supernumerary chromosomes during meiotic prophase; these two chromosomes move preferentially (70%) to opposite poles during the subsequent stages producing a differential transmission of the supernumerary to the two sexes. It is possible that the stable frequency in the population is maintained by a conserved balance between the two sexes without the need of an elimination system. The evidence supports the interpretation that the supernumerary chromosome is partially homologous with the sex chromosome. The possibility that the supernumerary might evolve into a neo-Y chromosome is suggested.     

Developmental Capacity of Aneuploid Xenopus Species Hybrids

Diploid as well as triploid Xenopus interspecific hybrids generate aneuploid eggs because of the presence, at meiosis, of univalent chromosomes which are presumably distributed at random. Zygotes obtained from such eggs, fertilized by either normal or UV-irradiated sperm, were analysed for their developmental capacities. All monosomics die in the course of embryogenesis, whereby optimum capacities correspond closely with those observed in monosomic mammalian embryos, especially in mice. In contrast, hyperdiploid Xenopus are relatively viable: although many die exhibiting the'haploid syndrome'or various other abnormalities, 8% of them reach metamorphosis, and 1–2% become adults. Of the latter, the karyotype was established in 13 individuals. Among them, 8–16 supernumerary chromosomes were found to be present.     

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.     

Duplication of a conditionally dispensable chromosome carrying pea pathogenicity (PEP) gene clusters in Nectria haematococca

A supernumerary chromosome called a conditionally dispensable chromosome (CDC) is essential for pathogenicity of Nectria haematococca on pea. Among several CDCs discovered in N. haematococca, the PDA1 CDC that harbors the pisatin demethylation gene PDA1 is one of the best-studied CDCs and serves as a model for plant-pathogenic fungi. Although the presence of multiple copies is usual for supernumerary chromosomes in other eukaryotes, this possibility has not been examined well for any CDCs in N. haematococca. In this study, we produced strains with multiple copies of the PDA1 CDC by protoplast fusion and analyzed dosage effects of this chromosome. Using multiple methods, including cytological chromosome counting and fluorescence in situ hybridization, the fusion products between two transformants derived from the same strain that bears a single PDA1 CDC were shown to contain two PDA1 CDCs from both transformants and estimated to be haploid resulting from the deletion of an extra set or sets of A chromosomes in the fused nuclei. In phenotype assays, dosage effects of PDA1 CDC in the fusion products were evident as increased virulence and homoserine-utilizing ability compared with the parents. In a separate fusion experiment, PDA1 CDC accumulated up to four copies in a haploid genome.     

Chromosome studies in Andean taxa of Alstroemeria (Alstroemeriaceae)

Meiotic or mitotic chromosomes of seven Alstroemeria taxa, native in Argentina and Chile and with Andean distribution were studied: A. andina ssp. venustula, A. hookeri ssp. cummingiana , A. hookeri ssp. recumbens , A. pallida , A. patagonica , A. pseudospathulata and A. pygmaea . All were diploid with 2 n = 16. Karyotypes of A. andina ssp. venustula and A. pygmaea were analysed, revealing similarity to previously analysed species. Thus, to all existing arguments for not retaining Schickendantzia as a separate genus, we can add another one which merges A. pygmaea with other Alstroemeria species, and does not support its taxonomic uniqueness. In general, the meiotic behaviour was normal, with regular formation of eight bivalents except in A. hookeri ssp. cummingiana , in one plant of which meiotic irregularities at various stages were observed. At the tetrad stage a large percentage of the cells presented micronuclei. The presence of 0–2 supernumerary chromosomes in A. hookeri ssp. recumbens is recorded. The karyotype asymmetry presented by most Alstroemeria species is discussed. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 138 , 451–459.     

Karyotypic variability in Iheringichthys labrosus (Teleostei, Pimelodidae) from the Tibagi River basin (Paraná State, Brazil)

Cytogenetic analyses were carried out in a populational sample of Iheringichthys labrosus from the Guaraúna River (Upper Tibagi River; Paraná State, Brazil) in order to provide a karyotypic comparison with another previously studied population from the Lower Tibagi River, characterized by the presence of 32m + 8sm + 6st + 10a (2n = 56, FN = 102) and occurrence of supernumerary chromosomes (80% of individuals). The 17 specimens of I. labrosus (6 females, 10 males and 1 of unknown sex) from the Upper Tibagi River showed 2n = 56 chromosomes, a karyotype formula of 14m + 32sm + 4st + 6a (FN = 106), without evidence of sex chromosome heteromorphism or supernumerary chromosomes. The heterochromatin was detected at telomeric and centromeric positions in several chromosomal pairs. The Ag-nucleolar organizer regions were heteromorphic and located at terminal position on short arms of the 16th chromosomal pair, suggesting a positive association with heterochromatic regions. The inter-populational karyotypic differentiation reported indicates distinct evolutionary pathways within I. labrosus in the Tibagi River basin.     

B chromosomes in Amazonian cichlid species

B chromosomes are reported in three different Amazonian cichlid species. One to three supernumerary microchromosomes were detected in the peacock bass Cichla monoculus (4 out of 28 specimens) and Cichla sp. (4 out of 13 specimens), and pike cichlids Crenicichla reticulata (2 out of 5 specimens), with no similar standard chromosomal morphology. C-banding revealed that B chromosomes are totally heterochromatic. We suggest two scenarios for the origin of these B chromosomes either by chromosomal breakdowns due to mutagenic action of methyl mercury present in the aquatic environment or by interspecific origin due to hybridization events.     

THE GENUS COLLINSIA. XVI. SUPERNUMERARY CHROMOSOMES

Dhillon , T. S., and E. D. Garber . (U. Chicago, Chicago, Ill.) The genus Collinsia. XVI. Supernumerary chromosomes. Amer. Jour. Bot. 49 (2) : 168–170. 1962.—The pairing and transmission of supernumerary chromosomes in Collinsia solitaria and in hybrids between C. sparsiflora subsp. arvensis and C. bruceae were studied. Two supernumerary chromosomes usually formed a bivalent with 1 chiasma at metaphase I; 4 supernumerary chromosomes occasionally yielded a trivalent and univalent at this stage. Plants with 2–4 supernumerary chromosomes were fertile and plants with 5–8, sterile. Plants with a given number of supernumerary chromosomes when used as seed or pollen parent gave gametes with a higher number of such chromosomes.     

SUPERNUMERARY CHROMOSOMES IN SAXIFRAGA VIRGINIENSIS (SAXIFRAGACEAE)

Acetocarmine squashes of root tips have demonstrated that 2n = 20 and 38 in Saxifraga virginiensis. These contrast with the earlier reported count of 2n = 28 for this species. In several populations supernumerary chromosomes were detected. Both intrapopulational and interpopulational variation in supernumerary chromosome number were detected, with the largest number of supernumerary chromosomes observed being six. Because these supernumerary chromosomes are equal in size to many of the smaller A chromosomes during mitotic metaphase, the presence of supernumerary chromosomes in this species could not be ascertained by analysis of mitotic metaphase preparations alone. During mitotic prophase, however, the supernumerary chromosomes of S. virginiensis are highly heterochromatic, appearing more densely coiled and darkly stained than the A chromosomes. This characteristic facilitated the recognition of supernumerary chromosomes in this species. The similarity in size of A and supernumerary chromosomes during mitotic metaphase and the observation of six supernumerary chromosomes in one population suggest that the count of 2n = 28 reported earlier for S. virginiensis may actually be a misinterpretation of 2n = 20 plus 8 supernumerary chromosomes. Furthermore, these findings and the observation of this same supernumerary chromosome phenomenon in other species of Saxifraga raise the possibility that some of the many disparate chromosome counts attributed to aneuploidy in the large genus Saxifraga may also be the result of misinterpretations of supernumerary chromosomes as A chromosomes.     

Karyotypic evolution and phylogeny of Mexican Passalidae (Coleoptera: Polyphaga: Scarabaeoidea)

The chromosomes of 26 taxa from Mexico of the tribes Passalini (three species) and Proculini (23 species) have been studied, increasing the karyotypically known species of the family Passalidae to 56. Karyotypic dynamism is high since the diploid number varies from 18 to 44 in the tribe Proculini. and from 25 to 31 in the tribe Passalini. In addition, supernumerary chromosomes, chromosome heteromorphism, translocations and possible sex multivalents have been found. These results contrast with the numerical conservatism found in related families of the superfamily Scarabaeoidea. However, both tribes are conservative with regard to sex determination, as all species of Proculini have male XY chromosomes whereas species of the Passalini have male XO chromosomes. It is postulated that differences in patterns shown by these two tribes are mainly due to population structure, because many species of Proculini are endemic to restricted areas of Meso and South America, favouring the settlement of karyotypic changes, whereas species of Passalini are distributed over large areas in the lowlands. It is also postulated that the ancestral karyotypic formula of the family is close to 12–14 pairs of autosomes although the ancestral male sex determination may be either XY or XO. At present only a weak relationship between morphological and karyotypic evolution has been found, which together with the marked numerical variability found within and between genera make it difficult to obtain phylogenetic conclusions from karyotypic results.     

六种麻蜥核型的研究

报道麻蜥属(Eremias ,Lacertidae) 6种15个不同居群的染色体核型及银分带核型。丽斑麻蜥（E. argus）、快步麻蜥(E. velox)、敏麻蜥(E. arguta)、密点麻蜥(E. multiocellata)、网纹麻蜥（E. grammica ）的核型一致：2n=38=36I+ 2m,NF=38;虫纹麻蜥（E. vermiculata） 2n=38=12V+2sI+22I+2m, NF=50。中国麻蜥属的核型可分为3个类型：（1）丽斑麻蜥型（2）山地麻蜥(E. brenchleyi)型（3）虫纹麻蜥型。虫纹麻蜥核型演化有两种可能性（1）经历三倍体阶段,并通过罗伯逊易位形成;（2）通过染色体臂间倒位形成,倒位成因可能和天山山脉以及青藏高原的隆起有关。密点麻蜥、快步麻蜥、敏麻蜥、网纹麻蜥、虫纹麻蜥均观察到一对NOR于一对较小染色体对上。雌雄个体中均未发现性异型染色体。Abstract: Based on the Giemsa-dyeing karyotypes and silver-staining bands of 15 populations from different localities in China belonging to 6 species of the genus Eremias , We found all species studied have 19 pairs of chromosomes, the size of chromosomes reduces gradually and there are no marked differences between the arranged pairs of macrochromosomes except the last pair of microchromosome. There are the same karyotype formula as 2n=38=36I+2m with NF=38 in E. argus、 E. multiocellata、 E. velox、 E. arguta and E. grammica; but the karyotype formula of E.vermiculata is different as 2n=38=12V+2sI+22I+2m with NF=50. The NOR are all located on one small pair in female of E. velox, and E. arguta , in male of E. grammica and E. vermiculata ,and in both male and female of E. multiocellata. We have not found two or more than two pairs of NOR. Having one pair of NOR may be common in Genus Eremias and also the trait of Eremias. We speculate that the derivation of the karyotype of E.vermiculata had two possible way: one experienced the stage of triploid, and later the Robertsonian transposal of chromosomes; the other way was through the inversions between the arms on the chromosome and the phenomenon of inversions might occur during or subsequently after the upheaval of the Tibet and Qinghai plateau and the founding of the Tianshan . With regard to the trend of the evolution of chromosomes in the lizards [1], the karyotype of E.vermiculata is more advanced. Making specialties of E. vermiculata will help in building the phylogenic tree of Eremias. In both male and female of the species studied, the heteromorphic sex-chromosomes were not found.     

纳特竖蟾(无尾目:滑背蟾科) 的细胞遗传、与相关属物种的核型相似性并兼论其分类地位

本文报道纳特竖蟾(Eupemphix nattereri Steindachner,1863)巴西3个产地标本的核型,均为2n=22,由1对端部和10对中或亚中部着丝点染色体组成.仅一对NORs位于No.11染色体对长臂端部,而其位置有别于常见的具有标志性意义的近着丝点位置.该NORs的位置为rDNA为探针的荧光原位杂交(FISH)所确认.各染色体对中,着丝点C-带明显,插入或端带偶见.某些标本的No.8同源染色体对间C-带的大小随异染色质化的程度不同而变化.居于较小的实验标本量,这种在3个产地的雌或雄性标本之一中观察到的C-带异型现象可能为种下细胞地理学变异,亦或为细胞学意义的性染体分化.3产地之一的标本核型的No.11着丝点C-带异染色质化的程度较高.CMA3染色检测到部分GC-rich区域,DAPI染色未显示任何AT-rich区.成功获得BrdU复制带,并将其与滑背蟾类动物(leiuperid)中近缘属及物种进行对比分析.比较结果表明,纳特竖蟾的核型与斑符泡蟾种组(Physalaemus signifer group)难以相互区别,而与肿肋蟾属(Pleurodema Tschudi,1838)极为相似.核型数据支持形态学上将纳特竖蟾、二光肿肋蟾[Pleurodema diplolister (Peters,1870)]和短头肿肋蟾[P.brachyops(Cope,1869)]划为同一分支的观点[动物学报 53(2):285-293,2007].     

Supernumerary chromosome variation and heterochromatin distribution in the endemic New Zealand frog Leiopelma hochstetteri

Specimens of the endemic New Zealand frog Leiopelma hochstetteri from Tapu on North Island were found to have six, nine or ten supernumerary chromosomes in their karyotypes. In comparison with previously published data, these results further indicate probable geographic variation in supernumerary chromosome number between populations. Increased numbers of supernumeraries in these frogs is correlated with apparent decrease of centromeric heterochromatin in the five large metacentric chromosomes of the karyotype, as detected by C-banding. Meiosis was abnormal in a male with a high number of supernumeraries. In lampbrush preparations from a single female with one supernumerary univalent, the supernumerary often had a denser, beaded appearance in comparison with the regular bivalents. Evidence is consistent with the notion that these supernumerary chromosomes may have arisen from centromeric fragments.     

Parallel polymorphism for supernumerary segments inChorthippus parallelus (zetterstedt)

A complex and parallel pattern of polymorphism for heterochromatic supernumerary segments in the M₇ and S₈ chromosomes has been found in 14 populations of the meadow grasshopperChorthippus parallelus. Nine distinct karyotype classes for these two chromosome pairs occur though they are not equally represented in different populations. Populations differ also with respect to the frequency of supernumerary segments they contain. In all populations the presence of supernumerary segments leads to a significant elevation of mean cell chiasma frequency compared to individuals from the same population lacking such segments. The extent of the effect appears to differ in different populations. The observed frequencies of S₈ karyotypes conform to the expectations of a Hardy-Weinberg distribution. Those of the M₇, however, do not, and in all but one of the 14 populations there is a significant excess of homokaryotypes. In the Ashurst population 26% of the individuals sampled were characterised by germ-line polysomy for the M₄ chromosome, either in the form of entire tetrasomics or more usually as mosaics ranging from tri- to hepta-somics. In all these polysomics the M₄ chromosomes in excess of two were regularly heteropycnotic at first meiotic prophase from zygotene to diakinesis. As a consequence of this multivalents are rare. Extra M₄ chromosomes do not modify the chiasma characteristics of the other chromosomes in the complement. Nor do they modify the action of the supernumerary segments in any way.