Notable homologous variation in chromosomal races of the common shrew

This paper is a review of the rare phenomenon of chromosome intraspecies variation manifested in monobrachial homology series in the comprehensively investigated karyotype of the common shrew Sorex araneus Linnaeus, 1758 (Eulipotyphla, Mammalia). The detailed dataset on the account of this mammalian species was drawn from the recently published monograph by Searle et al. (2019) “Shrews, Chromosomes and Speciation”. The parallels to the law of homologous series in variation by Nikolai Vavilov are discussed.

Chromosome Translocations as a Driver of Diversification in Mole Voles Ellobius (Rodentia, Mammalia)

The involvement of chromosome changes in the initial steps of speciation is controversial. Here we examine diversification trends within the mole voles Ellobius, a group of subterranean rodents. The first description of their chromosome variability was published almost 40 years ago. Studying the G-band structure of chromosomes in numerous individuals revealed subsequent homologous, step-by-step, Robertsonian translocations, which changed diploid numbers from 54 to 30. Here we used a molecular cytogenetic strategy which demonstrates that chromosomal translocations are not always homologous; consequently, karyotypes with the same diploid number can carry different combinations of metacentrics. We further showed that at least three chromosomal forms with 2n = 34 and distinct metacentrics inhabit the Pamir-Alay mountains. Each of these forms independently hybridized with E. tancrei, 2n = 54, forming separate hybrid zones. The chromosomal variations correlate slightly with geographic barriers. Additionally, we confirmed that the emergence of partial or monobrachial homology appeared to be a strong barrier for hybridization in nature, in contradistinction to experiments which we reported earlier. We discuss the possibility of whole arm reciprocal translocations for mole voles. Our findings suggest that chromosomal translocations lead to diversification and speciation.

Chromosome Synapsis and Recombination in Male Hybrids Between Two Chromosome Races of the Common Shrew (Sorex araneus L., Soricidae, Eulipotyphla)

The hybrid zone between chromosome races of the common shrew (Sorex araneus) provides an exceptional model to study the potential role of chromosome rearrangements in the initial steps of speciation. The Novosibirsk and Tomsk races differ by a series of Robertsonian fusions with monobrachial homology. They form a narrow hybrid zone and generate hybrids with both simple (chain of three chromosomes) and complex (chain of eight or nine) synaptic configurations. Using immunolocalisation of the meiotic proteins, we examined chromosome pairing and recombination in males from the hybrid zone. Homozygotes and simple heterozygotes for Robertsonian fusions showed a low frequency of synaptic aberrations (< 10%). The carriers of complex synaptic configurations showed multiple pairing abnormalities, which might lead to reduced fertility. Recombination frequency in the proximal regions of most chromosomes of all karyotypes was much lower than in the other regions. The strong suppression of recombination in the pericentromeric regions and co-segregation of race specific chromosomes involved in the long chains would be expected to lead to linkage disequilibrium between genes located there. Genic differentiation, together with the high frequency of pairing aberrations in male carriers of the long chains, might contribute to maintenance of the narrow hybrid zone.

Spermatogenesis in house mouse in a Robertsonian polymorphism zone

Robertsonian (Rb) translocations can be important in speciation as a mechanism of postzygotic isolation between populations. Meiotic non-disjunction, gametogenic impairment, and association of impaired autosomal segments with sex chromosomes have been postulated as mechanisms responsible for reducing fertility in Rb mice. Quantitative histological studies needed to understand the role of Rb fusions in gametogenic impairment are scarce. Most research on Rb mice has analyzed meiotic non-disjunction of laboratory and wild-derived strains, which have complex or simple structural heterozygosity with large numbers of fusions. Using histological multilevel sampling, we examined spermatogenesis in mice from the Rb polymorphism area of Barcelona. We studied four chromosomal groups having: a) one Rb heterozygote fusion and 2n=39, b) one Rb heterozygote fusion and 2n=31, c) three Rb heterozygote fusions without monobrachial homology and with diploid number ranging from 2n=29 to 2n=37, and d) only Rb homozygote fusions with diploid number ranging from 2n=28 to 2n=30. Standard mice from the area surrounding the Rb zone were used as control. We analyzed morphological variables of the testes, relative frequency of stages in the seminiferous epithelium cycle, the ‘round spermatids:primary spermatocytes’ ratio, and other derived parameters. Our results reveal that structural homozygote mice and simple heterozygote mice having as few as one to three Rb fusions undergo greater germ cell death (GCD) than standard mice, suggesting that Rb fusions are related to increased GCD (in both the heterozygous and homozygous state) and may be the main cause of decreased gene flow between mice populations from this area.

Chromosomal fusion and meiotic behaviour in Delena cancerides (Araneae: Sparassidae). I. Chromosome pairing and X-chromosome segregation

Surface spreading of meiotic material in Delena cancerides indicates that pairing initiation among metacentric chromosomes with monobrachial homology differs from that of telocentric forms and free metacentric bivalents and results in a star-shaped structure at pachytene. Distance cosegregation of the three X chromosomes in ancestral, telocentric forms is prefaced by a centric association early in prophase I. This centric association of the X chromosomes is conserved in metacentric races despite the presence of an X-autosome fusion.Key words: synaptonemal complex, translocation heterozygote, X chromosome, spider.

Factors responsible for a karyotypic polymorphism in the common shrew, Sorex araneus

A Robertsonian karyotypic polymorphism in the common shrew in the Oxford area, first described in the 1950s, was re-examined. The polymorphism involves chromosome arm combinations no and pr (characteristic of the Oxford karyotypic race), ko (characteristic of the Hermitage karyotypic race) and jl (found in both races). The poly- morphism for jl was sporadic along a north-south transect through the Oxford area, with the frequency of the twin-acrocentric morph never exceeding 10%. The frequency of the Oxford race-specific metacentrics decreased and the frequency of the Hermitage race-specific metacentric ko increased from north to south along the transect. At a latitudinal grid reference of about 180 km, there was a high frequency of individuals with chromosome arms k, n, o and q in the ancestral acrocentric state. This was coincident with the area of occurrence of ko-kq and ko-no OxfordHermitage hybrids. Such hybrids are double Robertsonian heterozygotes with monobrachial homology and are likely to suffer reduced fertility in consequence. It is proposed that this is a source of selection against the monobrachial hybrids and hence results in an increase in frequency of the acrocentric morphs. This scheme goes some way to explain the dines of polymorphism for arm combinations kq, and ko, but it is suggested that other selective factors are involved. It cannot explain the cline of polymorphism for pr, which is in general terms similar to that for kq and no, but is more shallow and centred further north.