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Corresponding author: Nina Bulatova ( ninbul@mail.ru ) Academic editor: Valentina G. Kuznetsova
© 2020 Nina Bulatova.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Bulatova N (2020) Notable homologous variation in chromosomal races of the common shrew. Comparative Cytogenetics 14(3): 313-318. https://doi.org/10.3897/CompCytogen.v14i3.54526
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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
Chromosome rearrangements (CRs), mammal karyotype, N.I. Vavilov heritage, Robertsonian fusions, Sorex araneus
Genetics started in the XXth century with rediscovery of G. Mendel’s hereditary laws, T.H. Morgan’s chromosome theory of heredity and prior evolutionary generalizations of W. Bateson, the author of the term “genetics”. Advances of the first two decades in the emerging field of plant genetics have been promptly consolidated into the law of homologous series in variation by Nikolai Vavilov, who was considered himself to be a student of William Bateson. A concise first presentation of the law idea (
The record of chromosomal variation within the common shrew was recently reviewed in essential details in the monograph “Shrews, Chromosomes and Speciation” which summarized more than the 30-year period of joined multidisciplinary studies of S. araneus chromosomal races in Eurasia initiated by the International Sorex araneus Cytogenetics Committee, ISACC (
The chromosomes of S. araneus are composed of 21 chromosomal arms that can be fused in a variety of combinations in different populations forming an astonishing array of chromosomal races. According to the nomenclature, 21 arms are designated by Latin letters (a to s) in correspondence with the arrangement in decreasing size from largest to smallest.
In the karyotype of S. araneus, chromosomes in pairs appear in either the bi-armed (metacentric) or one-armed (acrocentric) form. Among autosomes, three pairs are always bi-armed and demonstrate stable combination of chromosome arms (af, bc and tu). One other pair is always composed of arms j and l, but can display individual or population Robertsonian polymorphism appearing in acrocentric or/and metacentric forms (j, l / jl) (
Sex chromosomes of S. araneus have complex origin because of the ancient event of the autosome to sex chromosome translocation: a fusion between the original “true” X (arm e) and an autosome (arm d). Thus, in females, XX pair is represented by bi-armed (de) chromosomes, and in males, by a system of triple sex chromosomes – X(de)Y1(“true” Y)Y2(d).
Ten other chromosome arms (g to r, except j, l) are fused in a variety of combinations which show the remarkable intraspecies polytypic variation. Such arm reshuffling creates a variety of chromosomal races: 37 different combinations of chromosomal arms fused into metacentrics were detected in 76 described chromosomal races (see tables 5.2 and 5.3 in:
af , bc, de (XX)/ d (Y2), g*, h*, i *, j/l , k*, m*, n*, o*, p* , q*, r*, s (Y1), tu.
To analyse the peculiarities of the variable group of chromosome arms, the list of the synoptic table 5.3 from
In our analysis, each chromosome series begins with an acrocentric state (for instance, g) and accumulates varying fusion combinations with other elements of the variable group (in this case – gi, gk, gm, go, gp, gq, gr). That is, from nine possible combinations, two variants of the arm g fusions are absent from this series (gh, gn) – but probably could still be found in nature.
All nine possible fusion variants were realized in two cases, for the arms o and q (Table
Fusions predominate among evolutionary changes of karyotypes in the genus Sorex Linnaeus, 1758. Cascades of fusions have happened in the past karyotype evolution of this genus according to refined cytogenetic studies (
In this review, numerous series of fusion variations in race specific chromosomes of Sorex araneus were considered (Tables
The law of homologous series in chromosome variation may be considered a new notion of comparative cytogenetic studies not only of commemorative (
Serial presentation of chromosomal race specific metacentrics (monobrachial homologs) defined in Sorex araneus. Asterisks mark the fusions absent* in the list of chromosomal races, and potential** for the race/species karyotypes. A double letter designation is given in the alphabetical order following the standard nomenclature of chromosomes of S. araneus (
Arm | g | h | i | k | m | n | o | p | q | r | ** |
---|---|---|---|---|---|---|---|---|---|---|---|
g | * | gi | gk | gm | * | go | gp | gq | gr | gh** | |
h | * | hi | hk | * | hn | ho | * | hq | * | gh**, hm** | |
i | gi | hi | ik | im | * | io | ip | iq | * | ||
k | gk | hk | ik | km | * | ko | kp | kq | kr | ||
m | gm | * | im | km | mn | mo | mp | mq | mr | hm** | |
n | * | hn | * | * | mn | no | np | nq | nr | ||
o | go | ho | io | ko | mo | no | op | oq | or | NOR | |
p | gp | * | ip | kp | mp | np | op | pq | pr | ||
q | gq | hq | iq | kq | mq | nq | oq | pq | qr | NOR | |
r | gr | * | * | kr | mr | nr | or | pr | qr | ||
Total of 9 | 7 | 5 | 7 | 8 | 8 | 6 | 9 | 8 | 9 | 7 |
The chromosomal arms of the common shrew in alphabetical order (top line), without s (Y1). Homology to the elements of the Sorex ancestral karyotype, AKPS, is indicated in the rows below. Number of ancestral homologous elements that formed present-day arms through past fusions was identified by G-banding and chromosome painting by
a | b | c | d | e | f | g | h | i | j | k | l | m | n | o | p | q | r | tu |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
a1 | b1 | c1 | d1 | e | f1 | g1 | h | i | j1 | k1 | l1 | m | n | o | p | q | r | tu |
a2 | b2 | c2 | d2 | f2 | g2 | j2 | k2 | l2 | ||||||||||
a3 | b3 |
The author thanks colleagues and professionals for their kind linguistic and editorial input which has been very helpful in finishing this paper.