Хромосомы и видообразование у туко-туко (Ctenomys, Hystricognathi, Rodentia)

Tuco-tucos, South American rodents of the genus Ctenomys represent an interesting model of speciation. Their strict territorial and solitary life under the earth, vast but highly fragmented habitats, low migration activity were the causes of their very fast radiation. About 60 species of this genus have been described. They are found in a variety of habitats, from the mountains of the Andes to the coastal dunes of the Atlantic, from humid steppes of Pampas to dry deserts of Chaco. Tuco-tucos have a very high level of chromosomal polymorphism and polytypism based on Robertsonian and whole-arm reciprocal translocations and inversions, and can therefore be used to test different versions of chromosomal speciation hypothesis. The classic version of this hypothesis emphasizes the sterility of the hybrids, due to incorrect or incomplete chromosome synapsis in heterozygotes for rearrangements, germ cells death, chromosome nondisjunction and the generation of unbalanced gametes. The modern version of chromosomal speciation hypothesis suggests that the reduction of gene flow across chromosomal hybrid zones is due to the suppression of recombination in hybrids around the break points of rearrangements distinguishing the parental species. Field studies have not revealed strong negative effects of chromosomal heterozygosity on the fitness of the carriers. These results cast doubt on the validity of the classic version of the hypothesis. Analysis of chromosome behavior in the meiotic prophase in the chromosomal heterozygotes revealed significant changes in the frequency and distribution of recombination: crossingover suppression around the breakpoint and chiasma distalization. These changes can modulate the flow of genes between chromosomal races and amplify the divergence which has arisen due to spatial isolation. These data confirm the recombinational model of chromosomal speciation.Fil: Torgasheva, Anna A.. Novosibirsk State University; Rusia. Institute of Cytology and Genetics; RusiaFil: Savchenko, Ekaterina. Institute of Cytology and Genetics; RusiaFil: Gomez Fernandez, Maria Jimena. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; ArgentinaFil: Mirol, Patricia Monica. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; ArgentinaFil: Borodin, Pavel M.. Novosibirsk State University; Rusia. Institute of Cytology and Genetics; Rusi

Synapsis and recombination of autosomes and sex chromosomes in two terns (Sternidae, Charadriiformes, Aves)

The frequency of recombination and the patterns of crossover site distribution along the chromosomes vary considerably among animal species, including closely related species. Several hypotheses concerning the adaptive value and evolution of these variations were proposed. It was supposed that the recombination patterns of the species’ genomes are influenced by their phylogenetic history and ecology. However, most original data were obtained from mammals. The mammals show high karyological variability, which strongly influences the recombination patterns. Therefore it is important to study recombination rate and distribution in more karyologically stable taxa, such as reptiles and birds. We used immunolocalization of SYCP3, the protein of the lateral element of the synaptonemal complex (SC), centromere proteins and the mismatch-repair protein MLH1, which is associated with the recombination nodules, at the synaptonemal complex spreads of prophase oocytes of two tern species, black tern (Chlidonias niger) and common tern (Sterna hirundo). We first described the karyotype of Ch. niger (2n = 74, FN= 94) and identified suggestive rearrangements by which its karyotype differs from that of S. hirundo (2n = 68, FN = 90). We found that these species significantly differed by the numbers of the MLH1 foci per cell (Ch. niger: 53.0±4.2; S. hirundo: 44.1±5.0). We showed that the difference in the crossover numbers per cell was determined by the difference in the SC length (total and of individual bivalents) and by chromosomal rearrangements, which also influenced the distributions of crossover sites along the chromosomes. The difference in recombination patterns was higher between the rearranged homeologues than between the non-rearranged ones. We investigated the synaptic patterns of the heteromor phic Z and W chromosomes, localized the pseudoautosomal regions and estimated their lengths. In spite of several autosomal rearrangements, which differentiate these species, the structure and synaptic patterns of the sex chromosomes have not changed over 9 MY, which have passed since the moment of divergence between the genera Sterna and Chlidonias

Chromosome synapsis, recombination and epigenetic modification in rams heterozygous for metacentric chromosome 3 of the domestic sheep Ovis aries and acrocentric homologs of the argali Ovis ammon

Hybridization of domestic animal breeds with their wild relatives is a promising method for increasing the genetic diversity of farm animals. Resource populations derived from the hybridization of various breeds of domestic sheep with mouflon and argali are an important source of breeding material. The karyotypes of argali and domestic sheep differ for a Robertsonian translocation, which occurred in the common ancestor of mouflon and domestic sheep (Ovis aries) due to the centric fusion of chromosomes 5 and 11 of the argali (O. ammon) into chromosome 3 of sheep. It is known that heterozygosity for translocation can lead to synapsis, recombination and chromosome segregation abnormalities in meiosis. Meiosis in the heterozygotes for translocation that distinguishes the karyotypes of sheep and argali has not yet been studied. We examined synapsis, recombination, and epigenetic modification of chromosomes involved in this rearrangement in heterozygous rams using immunolocalization of key proteins of meiosis. In the majority of cells, we observed complete synapsis between the sheep metacentric chromosome and two argali acrocentric chromosomes with the formation of a trivalent. In a small proportion of cells at the early pachytene stage we observed delayed synapsis in pericentromeric regions of the trivalent. Unpaired sites were subjected to epigenetic modification, namely histone H2A.X phosphorylation. However, by the end of the pachytene, these abnormalities had been completely eliminated. Asynapsis was replaced by a nonhomologous synapsis between the centromeric regions of the acrocentric chromosomes. By the end of the pachytene, the γH2A.X signal had been preserved only at the XY bivalent and was absent from the trivalent. The translocation trivalent did not differ from the normal bivalents of metacentric chromosomes for the number and distribution of recombination sites as well as for the degree of centromeric and crossover interference. Thus, we found that heterozygosity for the domestic sheep chromosome 3 and argali chromosomes 5 and 11 does not cause significant alterations in key processes of prophase I meiosis and, therefore, should not lead to a decrease in fertility of the offspring from interspecific sheep hybridization

Germline-restricted chromosome (GRC) is widespread among songbirds

An unusual supernumerary chromosome has been reported for two related avian species, the zebra and Bengalese finches. This large, germline-restricted chromosome (GRC) is eliminated from somatic cells and spermatids and transmitted via oocytes only. Its origin, distribution among avian lineages, and function were mostly unknown so far. Using immunolocalization of key meiotic proteins, we found that GRCs of varying size and genetic content are present in all 16 songbird species investigated and absent from germline genomes of all eight examined bird species from other avian orders. Results of fluorescent in situ hybridization of microdissected GRC probes and their sequencing indicate that GRCs show little homology between songbird species and contain a variety of repetitive elements and unique sequences with paralogs in the somatic genome. Our data suggest that the GRC evolved in the common ancestor of all songbirds and underwent significant changes in the extant descendant lineages

Negative heterosis for meiotic recombination rate in spermatocytes of the domestic chicken Gallus gallus

Benefits and costs of meiotic recombination are a matter of discussion. Because recombination breaks allele combinations already tested by natural selection and generates new ones of unpredictable fitness, a high recombination rate is generally beneficial for the populations living in a fluctuating or a rapidly changing environment and costly in a stable environment. Besides genetic benefits and costs, there are cytological effects of recombination, both positive and negative. Recombination is necessary for chromosome synapsis and segregation. However, it involves a massive generation of double-strand DNA breaks, erroneous repair of which may lead to germ cell death or various mutations and chromosome rearrangements. Thus, the benefits of recombination (generation of new allele combinations) would prevail over its costs (occurrence of deleterious mutations) as long as the population remains sufficiently heterogeneous. Using immunolocalization of MLH1, a mismatch repair protein, at the synaptonemal complexes, we examined the number and distribution of recombination nodules in spermatocytes of two chicken breeds with high (Pervomai) and low (Russian Crested) recombination rates and their F1 hybrids and backcrosses. We detected negative heterosis for recombination rate in the F1 hybrids. Backcrosses to the Pervomai breed were rather homogenous and showed an intermediate recombination rate. The differences in overall recombination rate between the breeds, hybrids and backcrosses were mainly determined by the differences in the crossing over number in the seven largest macrochromosomes. The decrease in recombination rate in F1 is probably determined by difficulties in homology matching between the DNA sequences of genetically divergent breeds. The suppression of recombination in the hybrids may impede gene flow between parapatric populations and therefore accelerate their genetic divergence

Mendelian nightmares: The germline-restricted chromosome of songbirds

Germline-restricted chromosomes (GRCs) are accessory chromosomes that occur only in germ cells. They are eliminated from somatic cells through programmed DNA elimination during embryo development. GRCs have been observed in several unrelated animal taxa and show peculiar modes of non-Mendelian inheritance and within-individual elimination. Recent cytogenetic and phylogenomic evidence suggests that a GRC is present across the species-rich songbirds, but absent in non-passerine birds, implying that over half of all 10,500 bird species have extensive germline/soma genome differences. Here, we review recent insights gained from genomic, transcriptomic, and cytogenetic approaches with regard to the genetic content, phylogenetic distribution, and inheritance of the songbird GRC. While many questions remain unsolved in terms of GRC inheritance, elimination, and function, we discuss plausible scenarios and future directions for understanding this widespread form of programmed DNA elimination

Programmed DNA elimination of germline development genes in songbirds

In some eukaryotes, germline and somatic genomes differ dramatically in their composition. Here we characterise a major germline–soma dissimilarity caused by a germline-restricted chromosome (GRC) in songbirds. We show that the zebra finch GRC contains >115 genes paralogous to single-copy genes on 18 autosomes and the Z chromosome, and is enriched in genes involved in female gonad development. Many genes are likely functional, evidenced by expression in testes and ovaries at the RNA and protein level. Using comparative genomics, we show that genes have been added to the GRC over millions of years of evolution, with embryonic development genes bicc1 and trim71 dating to the ancestor of songbirds and dozens of other genes added very recently. The somatic elimination of this evolutionarily dynamic chromosome in songbirds implies a unique mechanism to minimise genetic conflict between germline and soma, relevant to antagonistic pleiotropy, an evolutionary process underlying ageing and sexual traits

MEIOSIS: HOW TO HALVE THE CHROMOSOME NUMBER

The notion of meiosis has been changed and refined for over a century since the discovery of this complicated way of cell division. Its success depends on precise time and space orchestration of many processes, such as chromosome replication, packaging, exchange of homologous regions, alignment in the plane of division, and disjunction. The development of molecular and immunocytochemical methods in recent decades cast light on the details of these processes and brought scientists closer to the understanding of mechanisms regulating them. This review presents the current notion of the major meiotic events by examples of yeast and mammals. Particular attention is paid to processes underlying chromosome synapsis and recombination, as well as monoorientation of sister kinetochores in the first division, the key features distinguishing meiosis from mitosis and ensuring chromosome number reduction