Increase in the number of the B-chromosomes and variants of their system in mouse <Emphasis Type="Italic">Apodemus peninsulae</Emphasis> in Mountain Altai population over 26 years

A new evolutionary genetic phenomenon—increase in the number of B-chromosomes (almost threefold) accompanied by change in B chromosome morphotypes—was discovered in the Artybash population of East Asian mouse Apodemus peninsulae (Mountain Altai) over a historically short period (22 years), comparable with a small number of mouse generations (ten generations). The process of increase in the number of A. peninsulae B-chromosomes in Mountain Altai has been monitored over 26 years (1980, 1986, 1988, 1990, 2002, and 2006). A concept of a new type of genomic mutations—mutations in the number of B-chromosomes—has been substantiated. The phenomenon of genomic mutations in pro-B-chromosomes has been discovered only in the Mountain Altai populations. This region differs from the remaining A. peninsulae dwelling localities by pollution with unspent liquid propellant (UDMH, heptyl). It is assumed that the variants of mouse B chromosome system in the studied A. peninsulae populations in the overall habitation area over long-term periods are likely to remain relatively stable and that their variation is controlled by homeostatic processes. Disturbance of these processes, in particular, due to anthropogenic load, can disrupt the cyclic pattern of increase and decrease in the number of A. peninsulae B-chromosomes in individual years.     

Supernumerary chromosomes, mosaicism, and dynamics in two populations of Eastern-Asian wood mouse (Apodemus peninsulae, Rodentia) from Primorski? Krai at various seasons of the year]

Karyotypes of 47 individuals of the wood mice Apodemus peninsulae from two reserves in Primorskii krai (Kedrovaya Pad' and Ussuriiskii) were studied during spring, summer, and autumn. In each population, variation in the number of B-chromosomes (2n = 48 + 0-5B) as well as the intratissue mosaicism determined by variation of the number, size and morphology of supernumerary chromosomes were described. Animals that have one dotlike B-chromosome were first described in both populations as rare variants. Individuals that have bone-marrow cell clones with two or three B-chromosomes were found to be predominant in the populations of Primorskii krai. The number of clones varied from one to three per animal. The frequency of mosaics showed seasonal variation. In the population of the Ussuriiskii reserve, a sharp increase in the frequency of animals with a stable karyotype was detected in autumn, at the phase of increased numbers. The variation for mosaicism was suggested to correlate with the population numbers in mice and to indicate the genetic differences between generations of the population.     

Instability of B-chromosomes in somatic and germline cells of Apodemus peninsulae

Instability of B-chromosomes was estimated in somatic and germline cells of samples Apodemus peninsulae from different localities of the species range. In 84 out of 188 animals (45%), in cells assessed for B-chromosome mosaicism, bone marrow cells with different B-chromosome number were observed. The numbers of B-chromosomes in spermatocytes at the pachytene stage were estimated in ten males. It was shown that the average number of B-chromosomes and the number of cell clones in germline cells was higher than the corresponding numbers in bone marrow cells. The higher number of B-chromosomes and their higher variability in germline cells than in somatic cells suggest the existence of a mechanism of premeiotic accumulation of B-chromosomes in spermatogenesis of A. peninsulae.     

Synaptonemal complex analysis of B-chromosome behavior in meiotic prophase I in the East-Asiatic mouse Apodemus peninsulae (Muridae, Rodentia)

The mitotic and meiotic chromosomes of four male East-Asiatic mice, Apodemus peninsulae, having three to seven chromosomes in addition to the standard karyotype (2n = 48), were investigated. B-chromosomes were represented by medium-sized metacentric and dotlike chromosomes. Mosaicism of bone marrow cells due to a numerical variation of accessory chromosomes was established for the males examined. Capacity of B-chromosomes to form axial elements and synaptonemal complexes in meiotic prophase I was revealed by electron microscopy. The occurrence of univalents of different morphology, bivalents, and multivalents, corresponding to B-chromosomes, was demonstrated. An increase in the number of B-chromosomes was found in spermatocytes at zygotene-pachytene relative to the number in bone marrow cells, which may be evidence of B-chromosome accumulation in the germ cell line of the East-Asiatic mouse.     

A complex B chromosome system in the Korean field mouse, Apodemus peninsulae

Information on B chromosomes of six subspecies of A. peninsulae Thomas, 1906, from 79 local populations of Russia (Siberia, Altai, Buryatia and the Far East), Mongolia, China, Korea and Japan (Hokkaido) is reviewed. The frequency of animals with B chromosomes is higher in this taxon than in other mammals and ranges from 0.4 up to 1.0, excluding two insular populations (Sakhalin Island and Stenin Island, Primorye) where Bs were not found. The B chromosome polymorphism shows four levels of variation in number (intraindividual mosaicism, intrapopulational and interpopulational), as well as variability in size, morphology and differential staining. Geographic variation was found among populations for these cytogenetic characteristics and, in some cases, it coincided with subspecies distribution. Comparative chromosome banding of micro and macro Bs illuminates possible pathways for their origin.     

Synaptonemal complexes of the A- and B-chromosomes of the spermatocytes from the East Asian mouse Apodemus peninsulae

The analysis of whole-mount preparations of synaptonemal complexes (SCs) from surface-spread spermatocytes of A. peninsulae (2n = 48A + 1, 2, ... 12 B) had revealed SCs of 23 autosomal bivalents, sex bivalent XY, axial cores and SCs of the B-chromosomes. The intercellular and interindividual variability of the number of B-chromosomes varied from 1 to 12 per cell. The SCs of autosomal bivalents were shown to have a typical structure. The structure and behaviour of SCs of sex bivalent throughout meiotic prophase I appeared to be similar to those observed in other species of this order. Mainly B-univalents and less frequently B-bivalents containing SCs were found to be formed in meiotic prophase I. The full homologues appear to be rarely seen among B-chromosomes of the East-Asiatic mouse. A tendency of forming clusters of B-univalents near the sex bivalent was found, in addition to B-bivalents with lateral elements, having the form of bi- and tri-stranded elements with rare synaptic fragments. Besides this, the SCs of the autosomes of pachytene cells were found to contain structures resembling the recombination nodules.     

The B-chromosome system--a marker of Apodemus peninsulae (Rodentia, Muridae) populations in the Ba?kal region

Karyological studies of 56 Apodemus peninsulae mice from 6 points in the neighbourhood of the Baikal lake revealed both considerable likeness and difference in the variants of the B-chromosomes system of this species. The mice from the regions situated at a distance of 150 kilometers from each other have similar karyotype variants, which allowed to consider these animals as one geographical population. The mice from the south-west, South Pribaikal and south-east regions considerably differ in the variants of the B-chromosomes system and may belong to 3 separate geographical populations.     

Polymorphism for supernumerary chromosomes in populations of Rana temporaria L

Occurrence of B-chromosomes in four populations of Rana temporaria L. in Minsk, its suburb and Berezinski? biosphere reserve has been studied. The percentage of individuals with B-chromosomes and the average number of B-chromosomes per genome are shown to multiply increase in the population inhabiting for 40 years in the region polluted with waste products. The results obtained are discussed from the standpoint of the action of mechanism for the accumulation of additional chromosomes due to environmental pollution.     

Presence and behaviour of B-chromosomes in Acanthophyllum laxiusculum (Caryophyllaceae)

Two karyotypic variants were recognized in populations of Acanthophyllum laxiusculum Schiman-Czeika. Variant A had 2N = 30 and variant B showed the presence of 0 to 3 B-chromosomes in addition to 2N = 30 chromosomes. Analysis of chromosome behaviour at meiosis showed that the presence of B-chromosomes increases chiasma frequency in A-chromosomes; this effect was higher for plants with odd numbers of B-chromosomes compared with plants with even numbers of B-chromosomes. Comparisons of variants A and B, suggests that B-chromosomes have an effect pollen stainability and seed production. It seems that the presence of B-chromosomes may increase pollen stainability and seed production in variant B.     

Populational chromosomal polymorphism the Asian forest mouse Apodemus peninsulae]

29 individuals of Apodemus peninsulae, an Asian forest mouse from the Suputin State Nature Reserve (Maritime Territory) were studied karyologically. Diploid chromosome sets of the animals studied contained from 48 to 52 chromosomes. The difference in the diploid chromosome numbers is caused by the presence in some individuals of supernumerary metacentric chromosomes (up to 4). The set is always represented by 48 acrocentric chromosomes, gradually decreasing in size. The supernumerary metacentric chromosomes vary in size from medium-sized to the smallest in the set. The character of intrapopulational chromosome polymorphism of continental and island populations of Apodemus peninsulae is discussed. Attempts are made to explain the polymorphism with respect to supernumerary chromosomes as the result of some viral diseases.     

Segregation distortion of mouse t haplotypes the molecular basis emerges

The t haplotype is an ancestral version of proximal mouse chromosome 17 that has evolved mechanisms to persist as an intact genomic variant in mouse populations. t haplotypes contain mutations that affect embryonic development, male fertility and male transmission ratio distortion (TRD). Collectively, these mutations drive the evolutionary success of t haplotypes, a phenomenon that remains one of the longstanding mysteries of mouse genetics. Molecular genetic analysis of TRD has been confounded by inversions that arose to lock together the various elements of this complex trait. Our first molecular glimpse of the TRD mechanism has finally been revealed with the cloning of the t complex responder (Tcr) locus, a chimeric kinase with a genetically cis active effect. Whereas + sperm in a +/t male have impaired flagellar function caused by the deleterious action of trans-active, t-haplotype-encoded 'distorters,' the mutant activity of Tcr counterbalances the distorter effects, maintaining the motility and fertilizing ability of t sperm.     

Nature and role of accessory chromosomes in silver-gray foxes. IV. The behavior of accessory chromosomes in meiosis

Comparison of chromosome number at somatic and spermatogonial mitoses has demonstrated the increase in the number of additional chromosomes in cells of germinal tissue. This may evidence a mechanism of B-chromosomes accumulation in foxes. B-chromosomes may lag as univalents, may form bivalent associations, or occasionally form trivalents at the stage of diakinesis-metaphase I, and they may associate with macrobivalents (A-chromosome bivalents). The analysis of metaphase II has shown that the distribution of B-chromosomes in the second metaphase is random resulting in gametes with various numbers of B-chromosomes.     

Molecular cloning of the t complex responder genetic locus

Although mouse t haplotypes carry recessive mutations causing male sterility and embryonic lethality, they persist in wild mouse populations via male transmission ratio distortion (TRD). Genetic evidence suggests that at least five t-haplotype-encoded loci combine to cause TRD. One of these loci, called the t complex responder (Tcr), is absolutely required for any deviation from Mendelian segregation to occur. A candidate for the Tcr gene has previously been identified. Evidence that this gene represents Tcr is its localization to the appropriate genomic subregion and testis-specific expression pattern. Here, we report the molecular cloning of the region between recombinant chromosome breakpoints defining the Tcr locus. These results circumscribe Tcr to a 150- to 220-kb region of DNA, including the 22-kb candidate responder gene. This gene and two other homologs were created by large genomic duplications, each involving segments of DNA 10-fold larger than the individual genes.     

B chromosome morphotypes of Apodemus peninsulae (Rodentia) from the Russian Far East

Polymorphism of B chromosome morphotypes in the natural populations of the Korean field mouse Apodemus peninsulae (n = 367) has been studied in five regions of the Russian Far East: Magadan and Amur regions, the Jewish Autonomous region, Khabarovsk and Primorskii regions. Six groups of B chromosome morphotypes have been described on the size and morphological parameters. On the basis of their combinations 78 cell clones have been revealed, 23 of which are found in the animals with a stable karyotype and 76--in mosaics. The mosaics have also more diverse and unique combinations of B chromosome morphotypes. No differences between the geographic and local populations of mice on the studied characteristics have been found. Homogeneous structure of DNA in the B chromosomes of this species at the territory of the Russian Far East, as shown previously, allowed us to combine the numerical and dimensional data for each clone by introduction conditional "mass quantity" of B chromosomes (mB index). The leading role of natural selection in production of "critical" mass of supernumerary chromosomes in individuals with a stable karyotype and the weakening of its role in mosaics has been suggested.     

Origin and molecular organization of supernumerary chromosomes of Prochilodus lineatus (Characiformes, Prochilodontidae) obtained by DNA probes

In Prochilodus lineatus B-chromosomes are visualized as reduced size extra elements identified as microchromosomes and are variable in morphology and number. We describe the specific total probe (B-chromosome probe) in P. lineatus obtained by chromosome microdissection and a whole genomic probe (genomic probe) from an individual without B-chromosome. The specific B-chromosome was scraped and processed to obtain DNA with amplification by DOP-PCR, and so did the genomic probe DNA. Fluorescence in situ hybridization using the B-chromosome probe labeled with dUTP-Tetramethyl-rhodamine and the genomic probe labeled with digoxigenin-FITC permitted to establish that in this species supernumerary chromosomes with varying number and morphology had different structure of chromatin when compared to that of the regular chromosomes or A complement, since only these extra elements were labeled in the metaphases. The present findings suggest that modifications in the chromatin structure of B-chromosomes to differentiate them from the A chromosomes could occur along their dispersion in the individuals of the population.     

THE GEOGRAPHICAL DISTRIBUTION OF B-CHROMOSOMES OF XANTHISMA TEXANUM DC. (ASTERACEAE). I. SURVEY OF THE RANGE

The distribution of B-chromosomes throughout the range of Xanthisma texanum has been determined from approximately 750 individuals from 80 wild populations in Texas, Oklahoma, and New Mexico. Counts of meiotic chromosome figures of n = 4_II + 1 B_I and n = 5_II (4_II + IB_II) are reported for the first time for subsp. texanum var. texanum and for subsp. texanum var. orientalis. The wide occurrence of B-chromosomes in natural populations of subsp. drummondii is also documented. B-chromosomes are present in 22 (27.2%) randomly distributed populations whose characteristics are in no way distinct from populations without B-chromosomes. In populations with B-chromosomes present, 19.8% of the plants contained one or two B-chromosomes, but in total only 5.8 % of all plants contained one or two B-chromosomes. All populations in one isolated population group of subsp. drummondii lack B-chromosomes.     

Molecular characterization and expression analysis of Mtmr2, mouse homologue of MTMR2, the Myotubularin-related 2 gene, mutated in CMT4B

Charcot-Marie-Tooth type 4B (CMT4B) is caused by mutations in the myotubularin-related 2 gene, MTMR2, on chromosome 11q22. To date, six loss of function mutations and one missense mutation have been demonstrated in CMT4B patients. It remains to be determined how dysfunction of a ubiquitously expressed phosphatase causes a demyelinating neuropathy. An animal model for CMT4B would provide insights into the pathogenesis of this disorder. We have therefore characterized the mouse homologue of MTMR2 by reconstructing the full-length Mtmr2 cDNA as well as the genomic structure. The 1932 nucleotide open reading frame corresponds to 15 coding exons, spanning a genomic region of approximately 55 kilobases, on mouse chromosome 9 as demonstrated by fluorescence in situ hybridization analysis. A comparison between the mouse and human genes revealed a similar genomic structure, except for the number of alternatively spliced exons in the 5'-untranslated region, two in mouse and three in man. In situ hybridization analysis of mouse embryos showed that Mtmr2 was ubiquitously expressed during organogenesis at E9.5, with some areas of enriched expression. At E14.5, Mtmr2 mRNA was more abundant in the peripheral nervous system, including in dorsal root ganglia and spinal roots.     

Karyological features of the Siberan spruce Obovata ledeb. from various locations

Results of karyological study of the Siberian spruce from different provenances are presented. A comparative idiogramm for the Siberian spruce from Krasnoyarsk Territory and the Khakass Autonomous Region was constructed. In addition to the typical chromosome number (2n = 24), the Siberian spruce has one or two B-chromosomes (2n = 24 + 1B; 2n = 24 + 2B). Hereditary transmission of B-chromosomes and their behaviour in mitosis have been studied. It is supposed that B-chromosomes of the Siberian spruce have a nucleolar-organizing ability.     

Geographical distribution of cytotypes and genomic structures in natural populations of the<Emphasis Type="Italic">Scilla scilloides</Emphasis> complex in Korea

Cytotype distribution, B-chromosome frequency, and genomic constitution in the natural populations of theScilla scilloides complex in Korea were analyzed. Plants with various cytotypes were found: AA (2n = 16), BB (18), AAB (25), ABB (26), ABB (34), ABBB (35), BBBB (36), AABBB (43), AAABBB (51) and AAAABBBB (68). Allotetraploid AABB plants predominated with a frequency of 68.3%, and were found to distribute all over the Korean peninsula and Cheju-do. In diploids, the type AA plants distributed throughout the Korean midwest, while the type BB plants were limited to Cheju-do. Two other cytotypes, ABBB and AABBB, were found only in the southern part of the Korean peninsula including Cheju-do. Chromosomal variations, aneuploidy, and centromeric shifts were also found in the natural populations. The cytotypes AAB and AAAABBBB are reported here for the first time. B-chromosomes were found in 149 (85.6%) of 174 populations, the highest frequency being 81.8% in BB populations. The number of B-chromosomes per plant ranged from 1 to 31, and IB plants predominated (21.0%). Subtypes, with respect to the number and composition of B-chromosomes, indicated that sexual reproduction is still prevalent in AABB populations.     

Enrichment and elimination of mutY mutators in Escherichia coli populations

The kinetics of mutator sweeps was followed in two independent populations of Escherichia coli grown for up to 350 generations in glucose-limited continuous culture. A rapid elevation of mutation rates was observed in both populations within 120-150 generations, as was apparent from major increases in the proportion of the populations with unselected mutations in fhuA. The increase in mutation rates was due to sweeps by mutY mutators. In both cultures, the enrichment of mutators resulted from hitchhiking with identified beneficial mutations increasing fitness under glucose limitation; mutY hitchhiked with mgl mutations in one culture and ptsG in the other. In both cases, mutators were enriched to constitute close to 100% of the population before a periodic selection event reduced the frequency of unselected mutations and mutators in the cultures. The high proportion of mutators persisted for 150 generations in one population but began to be eliminated within 50 generations in the other. The persistence of mutator, as well as experimental data showing that mutY bacteria were as fit as near-isogenic mutY(+) bacteria in competition experiments, suggest that mutator load by deleterious mutations did not explain the rapidly diminishing proportion of mutators in the populations. The nonmutators sweeping out mutators were also unlikely to have arisen by reversion or antimutator mutations; the mutY mutations were major deletions in each case and the bacteria sweeping out mutators contained intact mutY. By following mgl allele frequencies in one population, we discovered that mutators were outcompeted by bacteria that had rare mgl mutations previously as well as additional beneficial mutation(s). The pattern of appearance of mutY, but not its elimination, conforms to current models of mutator sweeps in bacterial populations. A mutator with a narrow mutational spectrum like mutY may be lost if the requirement for beneficial mutations is for changes other than GC --> TA transversions. Alternatively, epistatic interactions between mutator mutation and beneficial mutations need to be postulated to explain mutator elimination.