Corresponding author: VeronikaV. Golygina (
Academic editor: P. Michailova
A revision of the main and alternative banding sequences in chromosome II (CD) has been made for all 14 species of the
The
Revision of chromosome II (CD) mapping was conducted for 14
Mapping of arms C and D was done according to the Keyl-Devai mapping system (
Each banding sequence in each chromosomal arm is given a short designation as followes: three-letter abbreviation of the species name (for example, agi – for
Equipment in the Centre of Microscopical analysis of biological objects SB RAS in the Institute of Cytology and Genetics (Novosibirsk) was used for this work: microscope “Axioskop” 2 Plus, CCD-camera AxioCam HRc, software package AxioVision 4 (Zeiss, Germany).
Mapping of banding sequences of
Mapping of arm C main and alternative banding sequences in
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h’agiC1*† | 1a-2c 6c-f 7a-d 16a-17a 6hg 11d-15e 8a-11c 6b-2d 17b-22g C<br/> ( |
p’agi2C1* | 1a-2c 17a-16a 7d-a 6f-c 5a-6b 11c-8a 15e-11d 6gh 4i-2d 17b-22g C<br/> ( |
p’balC1* | 1a-2d 6c-e 7a-d 16a-17a 6h-f 11e-12d 4a-6b 11d-8a 15e-13a 3c-2e 17b-22g [24a-e 23a-c 23g-d 24fg] C§ ( |
p’balC2 | not mapped in Keyl-Devai system |
p’bonC1* | 1a-2c 6c-f 7a-d 16a-17a 6hg 11d-12d 4a-6b 11c-8a 15e-13a 3c-2d 17b-22g C<br/> ( |
p’borC1* | 1a-2c 6c-f 7a-d 16a-17a 6hg 11d-12d 4a-6b 11c-8a 15e-13a 3c-2d 17b-22g C<br/> ( |
p’entC1 | 1a 14a-11d 6gh 17a-16a 7a-d 6f-c 2c-1b 14b-15e 8a-11c 6b-2d 17b-22g C<br/> ( |
p’entC2* | 1a 11h-d 6gh 17a-16a 7a-d 6f-c 2c-1b 12a-15e 8a-11c 6b-2d 17b-22g C<br/> ( |
n’entC3 | 1a 11h-d 6gh 17a-16a 7a-d 6f-c 2c 5a-6b 11c-8a 15e-12a 1b-2b 4i-2d 17b-22g C ( |
p’murC1* | 1a-2c 15e-a 8a-11c 6b-4a 6c-f 7a-d 16a-17a 6gh 11d-12d 14e-13a 3c-2d 17b-22g C ( |
p’nudC1* | 1a-2c 11d-15e 8a-11c 6b-2d 6c-f 7a-d 16a-17a 6gh 17b-22g C<br/> ( |
p’pluC1* | 1a-2c 6c-f 7a-d 16a-17a 6hg 11d-12d 4a-6b 11c-8a 15e-13a 3c-2d 17b-22g C<br/> ( |
h’pluC2 | 1a-2c 6c-f 7a-d 16a-17a 6hg 11d-15e 8a-11c 6b-2d 17b-22g C<br/> ( |
p’sinC1* | 1a-c 12d-11d 6gh 17a-16a 7d-a 6f-c 2c-1d 13a-15e 8a-11c 6b-2d 17b-22g C<br/> ( |
p’spJC1* | not mapped in Keyl-Devai system |
p’spKC1* | 1a-2c 6c-f 7a-d 5c-6b 11c-8a 15e-11d 6gh 17a-16a 5b-2d 17b-22 C<br/> ( |
h’suwC1* | 1a-2c 6c-f 7a-d 16a-17a 6hg 11d-12d 4a-6b 11c-8a 15e-13a 3c-2d 17b-22g C<br/> ( |
p’useC1* | not mapped in Keyl-Devai system |
Phylogenetic relationship of main and alternative banding sequences in arms C and D before (
According to our analysis, 11 banding sequences that belong to 8 species require a revision of mapping in this arm.
In our opinion, the most important changes should be made in mapping of banding sequences of
The banding sequences of
Mapping of arm C main and alternative banding sequences in
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h’agiC1*† | =h’pluC2 |
p’agi2C1* | |
p’balC1* | 1a-2c 6c-f 7a-d 16a-17a 6hg 11d-12d 4a-6b 11c-8a 15e-13a 3c-2d 17b-22g |
p’balC2 |
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p’bonC1* | =p’pluC1 |
p’borC1* | =p’pluC1 |
p’entC1 | |
p’entC2* | |
n’entC3 | |
p’murC1* | |
p’nudC1* | |
p’pluC1* | 1a-2c 6c-f 7a-d 16a-17a 6hg 11d-12d 4a-6b 11c-8a 15e-13a 3c-2d 17b-22g C |
h’pluC2 | 1a-2c 6c-f 7a-d 16a-17a 6hg 11d-15e 8a-11c 6b-2d 17b-22g C |
p’sinC1* | 1a- |
p’spJC1* | =p’nudC1 |
p’spKC1* | 1a-2c 6c-f 7a-c 5c-6b 11c-8a 15e-11d 6gh 17a-16a 7d 5b-2d 17b-22g C |
h’suwC1* | =p’pluC1 |
p’useC1* |
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According to the previous mapping agi2C1 differs from agiC1 by two non-overlapping paracentric inversions (
The banding sequence p’balC2 was previously mapped according to Maximova’s system only. It originated from p’balC1 by one simple inversion and its up to date mapping in the Keyl-Devai system is shown in
It was presumed earlier that the main banding sequences of
As was mentioned above, the mapping of banding sequence p’entC2 is crucial for the mapping of n’entC3 and all banding sequences of
Mapping of banding sequences of
Mapping of some inversion breakpoints in species of
The mapping of n’entC3 should be corrected in accordance with mapping of p’entC1 (
Thorough analysis of the main banding sequences of
1a-d 11f-d 6gh 17a-16a 7a-d 6f-c 2c
Underline indicates simple inversion that distinguishes this banding sequence from p’entC2.
Both p’murC1 and p’nudC1 differ from this hypothetical banding sequence by simple inversions. Their revised mapping is shown in
Mapping of p’sinC1 require only a minor revision. According to the previous version of the mapping, region 1 was divided by an inversion between bands 1c and 1d. However, we had not been able to locate band 1d near band 13a so we suggest that the left breakpoint of this inversion situated between band 1d and 1e (
Until now the banding sequence h’useC1 has been mapped only partially (
Phylogenetic relationships of revised banding sequences in arm C of the
Mapping for banding sequences in this arm that has been published so far is shown in
Mapping of arm D main and alternative banding sequences in
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p’agiD1*† | 1a-d 4a-7g 18a-d 8a-10a 13a-11a 3g-1e 10e-b 13b-14f 20b-18e 17f-14g 20c-24g C( |
p’agi2D1* | 1a-d 4a-7g 18a-d 8a-10a 13a-11a 3g-1e 10e-b 13b-14f 20b-18e 17f-14g 20c-24g C ( |
p’balD1 | 1a-3g 10b-e 4a-7g 18a-d 8a-10a 13a-11a 13b-17f 18e-22e C<br/> ( |
p’balD2* | 1a-3g 10b-e 4a-7g 18a-d 8a-9e 15e-13b 11a-13a 10a 16a-17f 18e-22e C<br/> ( |
p’balD8 | 1a-3g 11a-13a 10a-8a 18d-a 7g-4a 10e-b 13b-17f 18e-22e C<br/> ( |
p’bonD1* | 1a-3g 11a-13a 10a-8a 18d-a 7g-4a 10e-b 13b-17f 18e-24g C<br/> ( |
p’borD1* | 1a-3g 11a-13a 10a-8a 18d-a 7g-4a 10e-b 13b-17f 18e-24g C<br/> ( |
h’borD2 | =h’pluD2 (Kerkis et al. 1988), not mapped according to Keyl-Devai system |
h’entD1* | 1a-2d 15e-16c 18d 8a-10a 13a-12a 18c-a 7g-4a 10e-b 13b-15d 2e-3g 11a-c 16d-17f 18e-24g C ( |
n’entD4 | 1a-2d 15e-16c 18d 8a-d 19h-18e 17f-16d 11c-a 3g-2e 15d-13b 10b-e 4a-7g 18a-c 12a-13a 10a-9a 20a-24g C ( |
p’murD1 | 1a-i 11c-a 3g-2e 15d-13b 10b-e 4a-7g 18a-c 12a-13a 10a-8a 18d 16c-15e 2d-a 16d-17f 18e-24g C( |
p’murD2* | not mapped according to Keyl-Devai system |
h’nudD1* | 1a-2d 15e-16c 18d 8a-10a 13a-12a 18c-a 7g-4a 10e-b 13b-15d 2e-3g 11a-c 16d-17f 18e-24g C ( |
p’nudD2 | not mapped according to Keyl-Devai system |
p’pluD1* | 1a-3g 11a-13a 10a-8a 18d-a 7g-4a 10e-b 13b-17f 18e-24g C ( |
h’pluD2 | 1a-3g 10b-e 4a-7g 18a-d 8a-10a 13a-11a 13b-17f 18e-24g C<br/> ( |
p’sinD1* | 1a-2g 13a 10a-8a 18d-a 7g-4a 10e-b 13b-14h 3g-2h 12d-11a 15a-17f 18e-24g C ( |
h’spJD1* | not mapped according to Keyl-Devai system |
p’spKD1* | 1a-3g 11a-13a 10a-8a 16d-13b 10b-e 4a-7g 18a-d 16e-17f 18e-24 C<br/> ( |
h’suwD1* | 1a-3g 10b-e 4a-7g 18a-d 8a-10a 13a-11a 13b-17f 18e-24g C<br/> ( |
p’useD1* | 1a-3g 11a-13part 9a-e 18part 8d-4a 10ba 13part-17f 18part-24g C<br/> ( |
h’useD2 | =h’pluD2 ( |
Mapping of banding sequences of
Mapping of banding sequences of
Mapping of arm D main and alternative banding sequences in
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p’agiD1*† | 1a-d 4a-7g 18a-d 8a-10a 13a-11a 3g-1e 10e-b 13b |
p’agi2D1* | =p’agiD1 |
p’balD1 | 1a-3g 10b-e 4a-7g 18a-d 8a-10a 13a-11a 13b-17f 18e-22e C |
p’balD2* | 1a-3g 10b-e 4a-7g 18a-d 8a- |
p’balD8 | 1a-3g |
p’bonD1* | =p’pluD1 |
p’borD1* | =p’pluD1 |
h’borD2 | =h’pluD2 |
h’entD1* | 1a- |
n’entD4 | 1a- |
p’murD1 | |
p’murD2* |
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h’nudD1* | =h’entD1 |
p’nudD2 |
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p’pluD1* | 1a-3g 11a-13a 10a-8a 18d-a 7g-4a 10e-b 13b-17f 18e-24g C |
h’pluD2 | 1a-3g 10b-e 4a-7g 18a-d 8a-10a 13a-11a 13b-17f 18e-24g C |
p’sinD1* | 1a-2g 13a 10a-8a 18d-a 7g-4a 10e-b 13b-14h 3g-2h 12d-11a 15a-17f 18e-24g C |
h’spJD1* | =h’entD1 |
p’spKD1* | 1a-3g 11a-13a 10a-8a 18d 16c-13b 10b-e 4a-7g 18a-c 16d-17f 18e-24g C |
h’suwD1* | 1a-3g 10b-e 4a-7g 18a-d 8a-10a 13a-11a 13b-17f 18e-24g C |
p’useD1* | =p’pluD1 |
h’useD2 | =h’pluD2 |
The banding sequences in arm D of both species are identical. They differ from p’pluC1 by two non-overlapping inversions. A correction should be made for breakpoints of the inversion in the proximal part of the arm: the left breakpoint falls between bands 14a and 14b instead of 14f and 14g, whereas the right breakpoint falls between regions 20 and 21 instead of bands 20b and 20c (
As was mentioned previously,
The banding sequence p’balD8 was considered previously as identical to p’pluD1 for all the arm length except the part affected by the pericentric inversion. However, our analysis has shown that this is not the case and p’pluD8 in fact originated from p’balD1 by a simple inversion. As a result, region 11 was broken into two parts and band 11a stayed between regions 10e-b and 13b-17f (
On the basis of our study we suggest that all banding sequences of these species have a common origin (
Mapping of banding sequences of
Thorough analysis of p’nudD2 allowed us to conclude that it differs from h’nudD1, not by one, but by two inversions (
Phylogenetic relationships of the revised banding sequences in arm D of the
The revision of banding sequences in chromosome CD of
We have shown that banding sequences of
The most ancient banding sequence in arm C should be considered h’pluC2 and the identical banding sequences of
In general, analysis of the phylogeny of banding sequences in arm C has shown that this arm has the highest level of divergence in comparison to arms A, B, and D, as only three clusters of homologous banding sequences exist in this arm, whereas there are four such clusters in arm A and D and seven in arm B (
The revision in arm D mostly provided minor changes in the mapping of inversion breakpoints without affecting phylogenetic relationships of banding sequences in general. The only significant change has come from the correction of the inversion breakpoint of p’spKD1 which has made it the ancestor for all banding sequences of
Considering the high level of banding sequence divergence in both arms, it can be stated that chromosome CD is the most divergent among the three big chromosomes of
Financial support was provided by grant RFBR 09-04-01440 and grants from the Presidium of Russian Academy of Sciences: “Biodiversity and dynamics of gene pools” and “Origin and evolution of biosphere”.