Factors of maintaining chromosome polymorphism in common vole Microtus arvalis Pallas, 1779: reduced fertility and meiotic drive

The common vole Microtus arvalis (the form obscurus) exhibits polymorphism of a pericentric inversion in chromosome pair 5 throughout the species range. In the Urals populations, the frequency of an acrocentric variant of the heteromorphic chromosome is very low (on average 3.2%) and virtually does not change annually. The factors of maintaining stable chromosomal polymorphism in the common vole were studied under conditions of a laboratory colony. Heterozygous and homozygous for the acrocentric chromosome females showed a significant reduction of the reproductive output irrespective of the male karyotype. This effect was manifested mostly in litter size at birth. A number of cytogenetic and exophenotypic characteristics, as well as parent--offspring transmission of this chromosome in crosses of various types, were examined. We have found meiotic drive in favor of the acrocentric, as a result of which the frequency of the acrocentric (without taking into account the postnatal mortality) totaled over all cross variants (0.48) was significantly higher than that expected with random segregation (0.42). It is likely that meiotic drive of the acrocentric largely compensates for the reduced fertility of its carriers, being among the factors of maintaining it in natural populations.     

Chromosomal and ontogenetic instability in sibling species of common vole (Microtus arvalis group): comparative aspects

Populations of chromosomal sibling species Microtus arvalis and Microtus rossiaemeridionalis were studied in Ural region in habitats affected by high radiation and the control ones. Frequency of chromosome disturbances in the marrow cells and fluctuating asymmetry (FA) of 8 craniometric characters were investigated. In impact populations the frequency of chromosome aberration was very high. Such frequency was also maintained in the offspring of the first laboratory generation of M. arvalis. In natural and control populations of both species frequently occurred individuals with anomalies in sex chromosome. Individuals of M. rossiaemeridionalis from Totsky radioactive region (forest-steppe zone) were characterized by very high integrative FA in comparison with control populations (southern taiga). At the same time neighboring impact and control populations of M. arvalis from southern taiga did not differ in this character. Despite the high level of caryotype divergence M. arvalis and M. rossiaemeridionalis showed similarity in mutation process that causes chromosome disturbances in somatic and germinative cells. Probably the level of FA of measured characters in both species is connected rather with geographical location than with man influence.     

Chromosomal polymorphism in planarians (Turbellaria,Tricladida) and the plate tectonics of the western Mediterranean

In two populations of Dugesia (Schmidtea) mediterranea the third chromosome pair consists of two heteromorphous elements. This chromosomal polymorphism, which is common in other animal groups, was hitherto unknown in planarians.The evolutionary and biogeographical significance of this chromosomal polymorphism is discussed, for the geographical distribution of the two aberrant populations may have relevance for the theories of microplate dispersal involving the western Mediterrancan.     

A population analysis of Robertsonian and Ag-NOR polymorphisms in brown trout (Salmo trutta)

An analysis of Robertsonian polymorphism and variation in the number of active NORs has been carried out in several populations of brown trout (Salmo trutta) from Northwestern Spain. The karyotype of this species appears to be soundly established, and essentially no variation has been found in chromosome number. Interindividual and interpopulation variation in arm number was detected, with figures ranging between 100 and 102 among individuals, and between 100.10 and 100.80 among populations. This variation in arm number is solely attributable to the polymorphism of the short arm of the main NOR-bearing pair 11, which can appear from acrocentric to metacentric in different individuals. Most populations analyzed showed the standard distribution of active NORs previously observed in this species. The Miño drainage basin, and specially the Chamoso population, showed a multi-chromosomal distribution of active NORs, with several new locations, always telomeric. In most cases no concordance was observed between previously detected rDNA sites in S. trutta and the new Ag-NOR locations. This fact suggests a transposition mechanism rather than an activation of silent rDNA sites to explain this multichromosomal NOR pattern.     

O chromosome inversion polymorphism in Northern and Atlantic Europe and its implications in the American colonization by Drosophila subobscura

The Western Mediterranean populations of D. subobscura have a very similar chromosomal inversion polymorphism to the American colonizing populations, with the exception of the O₅ inversion. This inversion, which is found in appreciable frequencies in colonizing populations and is distributed according to a significant latitudinal cline, both in North and South America, has never been reported in that Palearctic region. Therefore, an analysis of the distribution of this inversion, along a latitudinal cline, has been carried out in Europe. The O Chromosome inversion polymorphism has been studied in seven populations: Gävle and Lilla Edet (Sweden), Gesten (Denmark), Ter-Apel (Holland) and Crézy, Aizenay and Taulé (France). The O₅ inversion was only detected in three populations and in low frequencies: Gävle (3.7%), Lilla Edet (1.8 %) and Taulé (1.2 %). However, none of these three populations has all the inversions found in the American populations. The clinal distribution of some O chromosomal arrangements has also been studied according to latitude and the temporal changes of this polymorphism.     

Numerical and structural variations of the X chromosomes and no. 2 autosomes in mandarin vole, Microtus mandarinus (Rodentia)

Here we describe our studies on Microtus mandarinus faeceus of Jiangyan in Jiangsu province of China. By karyotype and G-banding analysis we have found variation in chromosome number and polymorphisms of the X chromosome and the second pair of autosomes of the subspecies. Chromosome number of the subspecies is 2n=47-50. The subspecies has three kinds of chromosomal sex: XX, XO and XY, among which one of the X chromosomes is subtelocentric (X(ST)) and the other is metacentric (X(M)). After comparing karyotypes of different subspecies, we found the specific cytogenetic characteristics of Microtus mandarinus, that is they have three kinds of chromosomal sex: XX, XO and XY; X chromosomes are heteromorphic; the chromosome number of female individuals are one less than male individuals; chromosome number of XX individuals are equal to that of XO ones. We hypothesize that Robertsonian translocation is the main reason of the polymorphism of the second pair of autosomes and variety of chromosome number, and it also causes the chromosome number evolution in different subspecies of Microtus mandarinus.     

Contrasting signatures of population growth for mitochondrial DNA and Y chromosomes among human populations in Africa

A history of Pleistocene population expansion has been inferred from the frequency spectrum of polymorphism in the mitochondrial DNA (mtDNA) of many human populations. Similar patterns are not typically observed for autosomal and X-linked loci. One explanation for this discrepancy is a recent population bottleneck, with different rates of recovery for haploid and autosomal loci as a result of their different effective population sizes. This hypothesis predicts that mitochondrial and Y chromosomal DNA will show a similar skew in the frequency spectrum in populations that have experienced a recent increase in effective population size. We test this hypothesis by resequencing 6.6 kb of noncoding Y chromosomal DNA and 780 basepairs of the mtDNA cytochrome c oxidase subunit III (COIII) gene in 172 males from 5 African populations. Four tests of population expansion are employed for each locus in each population: Fu's Fs statistic, the R(2) statistic, coalescent simulations, and the mismatch distribution. Consistent with previous results, patterns of mtDNA polymorphism better fit a model of constant population size for food-gathering populations and a model of population expansion for food-producing populations. In contrast, none of the tests reveal evidence of Y chromosome growth for either food-gatherers or food-producers. The distinct mtDNA and Y chromosome polymorphism patterns most likely reflect sex-biased demographic processes in the recent history of African populations. We hypothesize that males experienced smaller effective population sizes and/or lower rates of migration during the Bantu expansion, which occurred over the last 5,000 years.     

Cytogenetic analysis of a self-fertilizing fish, Rivulus marmoratus: remarkable chromosomal constancy over a vast geographic range.

The aplocheiloid killifish Rivulus marmoratus is the only known self-fertilizing hermaphroditic vertebrate. Most natural populations consist almost entirely of hermaphrodites and comprise arrays of homozygous clones. However, in almost all populations thus far studied, clonal variation, as detected with molecular techniques, is very high. A karyological survey was carried out on specimens from Brazil, the Bahamas, Belize, and Florida (4 locales) by C-banding, silver staining, and fluorescent staining. The chromosome complement of R. marmoratus is surprisingly uniform over its vast geographic range, in terms of both chromosome number and morphology, heterochromatin distribution and composition, and nucleolar organizer region (NOR) distribution. The short arms of chromosome pair 15, where NORs are located, showed the only variation detected in this study: those of pattern A were consistently shorter than those of pattern B; moreover, the latter show positive heteropycnosis with Giemsa staining. The present data demonstrate that chromosomal variation is not a significant part of the clonal divergence in this species, even though its breeding system, by forming homozygotes for new rearrangements almost immediately, should make that variation easier to detect. The high chromosomal homogeneity is discussed in the light of the peculiar natural history of the species.     

A chromosomal analysis of some water beetle species recently transferred from Agabus Leach to Ilybius Erichson, with particular reference to the variation in chromosome number shown by I. montanus Stephens (Coleoptera: Dytiscidae)

The karyotypes of seven Ilybius species are described and illustrated. All except I. wasastjernae have a basic karyotype of 34 autosomes plus sex chromosomes which are X0 ( male symbol ), XX ( female symbol ), with the X chromosome among the largest in the nucleus. This karyotype appears to be the norm for Ilybius and supports the transfer of the species concerned from Agabus to Ilybius. I. wasastjernae has 36 autosomes and the X chromosome is the smallest in the nucleus and its karyotype is unlike any other known karyotype in either Ilybius or Agabus. In most of the species studied no intraspecific variation has been detected. Exceptions are I. chalconatus, where there is one inversion polymorphism in one of the autosomes, and I. montanus whose autosome number has been found to vary from 29 to 34. Such variation is highly unusual among Coleoptera. The variation results from fusion-fission polymorphisms involving three different pairs of autosomes. In each case the fusions may be homozygous, heterozygous or absent. All populations investigated were polymorphic for some of the fusions, but only one (La Salceda, Spain) included individuals lacking all fusions. The frequencies of fused and unfused chromosomes were analysed in three English populations. In only one case was there a departure from the values expected from the Hardy-Weinberg equilibrium, and this population also showed a significant difference from the other two. Meiosis in males heterozygous for fusions involves the production of trivalents in first division, but results in the production of abundant sperm, with no evidence of chromosomal abnormalities in second metaphase, or of degenerating cells as a result of failed meiosis. The three fusions sites are consistent in all the populations studied, and it is concluded that these fusions represent unique historical events rather than current chromosomal instability.     

Evidence for chromosome number reduction and chromosomal homosequentiality in the 24-chromosome Korean frog Rana dybowskii and related species

The karyotype of the Korean frog Rana dybowskii with its pattern of C-band heterochromatin distribution was numerically analyzed. There are 2n = 24 chromosomes in the karyotype representing a reduction in number from the typical 2n = 26 chromosome karyotype of Rana. The karyotype shows other evidence of reorganization relative to 26-chromosome species. The chromosomes grade smoothly in size from largest to smallest without the two size classes that are characteristic for 26-chromosome species. In contrast to many 26-chromosome species, there are few centromeric C-bands but many interstitial ones. C-bands for each homologous chromosome pair are distinctive. A prominent secondary constriction is located on one of the smallest chromosomes, chromosome 11, in a position similar to that seen in most 26-chromosome species. The karyotype of R. dybowskii is compared to those of other species of Rana known to have 2n = 24 chromosomes; it is most similar to that of R. chensinensis, less so that of R. ornativentris and less still to that of R. arvalis in terms of the positions of centromeres and secondary constrictions. C-bands as well as secondary constrictions in the karyotypes of these frogs show evidence of chromosomal homosequentiality. The process and possible consequences of chromosome number reduction from an ancestral 26-chromosome karyotype is also evident in the karyotypes of these closely allied palearctic frogs. Pericentric inversions followed by fusion of two small elements apparently produced a new chromosome, chromosome 6, occurring originally among northeast Asian populations.     

Factors of maintaining chromosome polymorphism in common vole Microtus arvalis pallas, 1779: Reduced fertility and meiotic drive

The common vole Microtus arvalis (the form obscurus) exhibits polymorphism of a pericentric inversion in chromosome pair 5 throughout the species range. In the Urals populations, the frequency of an acrocentric variant of the heteromorphic chromosome is very low (on average 3.2%) and virtually does not change annually. The factors of maintaining stable chromosomal polymorphism in the common vole were studied under conditions of a laboratory colony. Heterozygous and homozygous for the acrocentric chromosome females showed a significant reduction of the reproductive output irrespective of the male karyotype. This effect was manifested mostly in litter size at birth. A number of cytogenetic and exophenotypic characteristics, as well as parent-offspring transmission of this chromosome in crosses of various types, were examined. We have found meiotic drive in favor of the acrocentric, as a result of which the frequency of the acrocentric (without taking into account the postnatal mortality) totaled over all cross variants (0.48) was significantly higher than that expected with random segregation (0.42). It is likely that meiotic drive of the acrocentric largely compensates for the reduced fertility of its carriers, being among the factors of maintaining it in natural populations.     

Patterns of inversion polymorphism in three species of the Drosophila melanogaster species group

In Drosophila, chromosomal polymorphism due to paracentric inversions is very common and constitutes an adaptive character. The degree of chromosomal variability varies in different species and also in different populations of the same species. Chromosomal polymorphism in Indian natural populations of three species, D. melaonogaster, D. ananassae and D. bipectinata which belong to the melanogaster species group has been studied and the quantitative data on frequency of inversions have been reported. Behaviour of chromosome inversions has also been studied in laboratory conditions. The present review summarises the work done on inversion polymorphism in Indian populations of three species which clearly demonstrates that these three species vary in their patterns of inversion polymorphism and have evolved different mechanisms for adjustment to their environments although they belong to the same species group.     

Autosomal and sex-chromosomal polymorphism in a wild population of the Norway rat,Rattus norvegicus

The karyotypes of 33 specimens of Rattus norvegicus captured in a wild population, were studied by means of conventional Giemsa staining and G- and C-banding methods. A chromosomal polymorphism for the presence or absence of extra segments in autosomes 3 and 12 and the X chromosome was found, beside a polymorphism for centromeric C-bands in pair no. 6.The frequency of different chromosome types found in this population was compared with that found in a wild population in Japan as well as with that found in some inbred strains.     

Human chromosomal polymorphism. VII. The distribution of chromosomal Q-polymorphic bands in different human populations

The distribution of chromosomal Q-polymorphic bands was studied in different human populations. The populations studied showed no differences in the relative amount of Q bands in all the 12 polymorphic loci of seven autosomes, but interpopulation differences did exist in the absolute amount of Q bands in all the 12 potentially polymorphic loci of seven autosomes, these differences consisting of uniform increases or decreases in this absolute amount. Comparisons of the mean number of Q-heterochromatin bands with fluorescence levels 4 and 5 per individual showed a consistent prevalence of this quantitative parameter of chromosomal Q polymorphism in females as compared to males in all the national groups. It is suggested that there is some dosage compensation of chromosomal Q-heterochromatin material in females due to the absence of a chromosome in their genome, which is able to "compensate" for the large Q band in chromosome Y which is present only in the karyotype of males.     

Lethal genes in O5, chromosomes of Drosophila subobscura from Europe and America

In populations of D. subobscura , a species that is know for its high chromosomal polymorphism, the O₅ inversion has a rather erratic frequency distribution in the Palearctic region. An O₅ lethal chromosomal line obtained from a Balkan population near Zanjic (South Adriatic, Montenegro, Yugoslavia) was tested for lethal allelism with other O₅ lethal chromosomal lines derived from American (USA and Chile) colonizing populations, and from the French population of Taulé. No allelism was found between the Balkan lethal gene and those from America and France. Thus, the lethal genes of the O₅ inversions are not of the same origin and it is most probable that the American colonizations did not start from the Zanjic population. The general difference in the chromosomal inversion polymorphism corroborates this conclusion. The cytological analysis confirms the assumption that all O₅ chromosomes studied are identical with respect to breakage points.     

The chromosomes of Nycticebus coucang (Boddaert, 1785) (Primates: Prosimii)

The high-quality karyotype of a specimen of Nycticebus coucang is described and illustrated. The X chromosome is found to be indistinguishable from that of the greater galagos, and may represent a synapomorphic trait. The Y chromosome is a medium to small submetacentric (3.2% TCL) and constitutes one of the larger Y chromosomes known in primates. N. coucang is found to have multiple NOR-bearing chromosomes in contrast to the single pair found in galagine and catarrhine monkeys. Since a single NOR-bearing pair is often considered ancestral for primates, this new finding may have important implications for the evolution of these cistrons. One of the chromosomal polymorphisms in this specimen is a pericentric inversion, involving a NOR-bearing autosomal pair (no. 6), that alters the position of the active site. Further, homologues 2p differ by aparacentric inversion. These results confirm that lorisiforms are characterized by considerable chromosomal polymorphism.     

A populational survey of 45S rDNA polymorphism in the Jefferson salamander Ambystoma jeffersonianum revealed by fluorescence in situ hybridization （FISH）

The chromosomal localization of 45S ribosomal RNA genes in Ambystoma jeffersonianum was determined by fluorescence in situ hybridization with 18S rDNA fragment as a probe (FISH-rDNA). Our results revealed the presence of rDNA polymorphism among A.jeffersonianum populations in terms of number, location and FISH signal intensity on the chromosomes. Nine rDNA cytotypes were found in ten geographically isolated populations and most of them contained derivative rDNA sites. Our preliminary study provides strong indication of karyotypic diversification of A.jeffersonianum that is demonstrated by intraspecific variation of 45S rDNA cytotypes. rDNA cytotype polymorphism has been described in many other caudate amphibians. We predict that habitat isolation, low dispersal ability and decline of effective population size could facilitate the fixation and accumulation of variable rDNA cytotypes during their chromosome evolution [Current Zoology 55(2):145-149,2009].     

Karyological characteristics of Siberian fir (<Emphasis Type="Italic">Abies sibirica</Emphasis> Ledeb.) in Central Siberia

Data are presented on the karyotype structure of the Siberian fir (Abies sibirica Ledeb.) from five populations of the Central Siberia. The chromosome set (2n = 24) contains seven pairs of metacentric (I–VII), four pairs of submetacentric (VIII and X–XII), and one pair of intercentric (IX) chromosomes. The morphometric parameters of the identified chromosome groups had close values in the studied populations. The variation coefficients of chromosomal parameters of the Siberian fir correspond to very low and low variability. The intraspecific chromosome polymorphism of the Siberian fir is connected mainly to variations in the number and peculiarities of the nucleolar organizing regions of chromosomes. In the territory with technogenic loading, a wide spectrum of genomic mutations of the mixoploidy type was observed in the seed off-spring of the Siberian fir; triploid seedlings, as well as rare cases of somatic reduction of chromosomes, were revealed.     

Cytogenetic analysis of three Italian populations of Coregonus lavaretus (Pisces, Salmoniformes) with chromosomal localization of major and minor ribosomal genes, and telomeric repeats

The European whitefish, Coregonus lavaretus, widely distributed in freshwater of northern Europe and introduced into the major lakes of northern Italy, has been restocked in central Italian lakes. In accordance with current managing practices, a reduced number of spawners contribute to reproduction within each lake and a certain degree of isolation is to be expected between populations from different lakes, resulting in the rapid fixing of chromosomal changes. A detailed survey of three populations from different lakes was carried out using classical and molecular cytogenetic techniques, to verify if specific chromosomal markers are present in the distinct populations. The comparative analysis revealed intraspecific variability of NORs and fixed differences in their number in the three populations. A co-localization of major and minor rRNA genes on one chromosome site was also observed. The original data regarding the chromosome mapping of the (TTAGGG)(n) telomeric repeat obtained in this study, demonstrated their exclusively terminal distribution, and a conspicuous inter-chromosomal variation in the number of repeats. The results are compared with data available for populations from native geographic ranges.