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Chromosomal inversion

January 01, 1970

An inversion is a chromosome rearrangement in which a segment of a chromosome becomes inverted within its original position. An inversion occurs when a chromosome undergoes a two breaks within the chromosomal arm, and the segment between the two breaks inserts itself in the opposite direction in the same chromosome arm. The breakpoints of inversions often happen in regions of repetitive nucleotides, and the regions may be reused in other inversions.[1] Chromosomal segments in inversions can be as small as 1 kilobases or as large as 100 megabases.[2] The number of genes captured by an inversion can range from a handful of genes to hundreds of genes.[3] Inversions can happen either through ectopic recombination between repetitive sequences, or through chromosomal breakage followed by non-homologous end joining.[4]

Inversions are of two types: paracentric and pericentric. Paracentric inversions do not include the centromere, and both breakpoints occur in one arm of the chromosome. Pericentric inversions span the centromere, and there is a breakpoint in each arm[5].

A clay model showing why heterozygous inversion loops are visible in polytene chromosome preparations
An inversion loop in the A arm of a chromosome from an Axarus species midge

Inversions usually do not cause any abnormalities in carriers, as long as the rearrangement is balanced, with no extra or missing DNA. However, in individuals which are heterozygous for an inversion, there is an increased production of abnormal chromatids (this occurs when crossing-over occurs within the span of the inversion). This leads to lowered fertility, due to production of unbalanced gametes. Inversions do not involve either loss or gain of genetic information; they simply rearrange the linear DNA sequence.

Detection edit

Cytogenetic techniques may be able to detect inversions, or inversions may be inferred from genetic analysis. Nevertheless, in most species, small inversions go undetected. More recently, comparative genomics has been used to detect chromosomal inversions, by mapping the genome.[6][7] Population genomics may also be used to detect inversions, using areas of high linkage disequilibrium as indicators for possible inversion sites. Human families that may be carriers of inversions may be offered genetic counseling and genetic testing.[8]

History edit

The first evidence of a chromosomal inversion was found in 1921 by Alfred Sturtevant in Drosophila melanogaster.[9] Since then, inversions have been found in a all eukaryotes.[3] When discovered by Sturtevant, inversions were regarded as areas of recombination suppression.

Originally, these inversions were noted in polytene chromosomes within the salivary glands of heterozygous Drosophila melanogaster larvae.[3] In 1970, Theodosius Dobzhansky noted that genes within an inversion had higher fitness than those that are found outside of the inversions, although this is an area that needs further study.[10]

One of the more recent models of inversions is the Kirkpatrick and Barton Model (2006), which states that inversions are selectively advantageous by linking together adaptive alleles. By physically linking co-adapted variants at multiple genes into distinct versions (haplotypes) of an inversion, selection should be more efficient in driving these variants to high frequency in a population. This is in contrast to non-inverted regions, which may allow adaptive and maladaptive alleles to be carried.[11]

Effects on recombination edit

When an inversion carrying chromosome is paired with a non-inverted homologous chromosome (Inversion heterozygotes) during meiosis, they fail to synapse properly and inversion loops are formed. A crossing-over within the loop can produce unbalanced gametes. In a paracentric inversion, recombination results in one dicentric chromatid and one acentric chromatid. During Anaphase, both recombinants are faced with problems. The acentric chromatid is pulled to one pole or the other, and the dicentric recombinant generates dicentric bridges as it is pulled in two directions.[12]

In a pericentric inversion, similar imbalanced chromosomes are produced. The recombinant chromosomes resulting from these crosses include deletions and duplications. The offspring produced by such gametes are mostly inviable, and therefore, recombination is indirectly suppressed within inverted regions.[12]

Evolutionary consequences edit

The suppressed recombination between inversion heterozygotes provides an opportunity for the independent evolution of the ancestral and inverted arrangements. At the beginning, the inverted arrangement lacks variation, while the ancestral one does not. If the inverted haplotype is not lost (eg. due to drift), the variation in the inverted arrangement can increase over time, and recombination rate in the inverted region is somewhat restored as more homozygotes are introduced.[13]

Chromosomal inversions have gained a lot of attention in evolutionary research due to their potential role in local adaptation and speciation. Because non-recombining inversion haplotypes may harbor multiple co-adapted gene variants, inversions are thought to facilitate local adaptation to different environments because natural selection is more efficient in driving such linked adaptive variants to high frequency within a population.[14] However, empirically demonstrating the presence of linked, co-adapted gene variants within inversions is difficult because inversion haplotypes do not recombine. Moreover, this possible positive effect of chromosomal inversions for adaptation to different environments rests on the assumption that adaptive gene variants linked into distinct inversion haplotypes are indeed co-adapted. This idea is, however, likely violated in situations where populations experience spatially or temporally varying selection. Because of fluctuating selection on inversion-linked variants, the absence of recombination between inversion haplotypes harboring distinct gene variants may then constrain rather than help adaptation to distinct environments.[15] The importance of chromosomal inversions in adaptation to different environments therefore remains an open empirical problem in evolutionary genetics.

Inversion polymorphism can be established in two ways. Genetic drift or selection can result in fixation of an inversion in a local population. Inversion polymorphism can result from gene flow between this population and a population without the inversion. Balancing selection can also result in inversion polymorphism by frequency dependence or overdominance.[13] The fitness differences between the inverted and the ancestral chromosome can either produce a stable polymorphism or can result in the fixation of one or the other chromosome.[16]

Inversions have been essential to sex chromosome evolution. In mammals, the Y chromosome is unable to recombine with the X chromosome, almost along its entire length. This non-recombining portion results from a series of inversions that overlap. Decreased recombination rate between sex determining loci and sex-anatagonistic genes is favored by selection. This causes linkage disequilibrium between the male determining locus and an allele at another locus that is beneficial to males. This can happen through inversions resulting in a non-recombining block including both loci, as is the case in the mammalian Y chromosome.[16]

Inversions can also be essential in the origination of new sex chromosomes. They can cause linkage disequilibrium between a sex-determining mutation and sex-antagonistic loci and create a new sex chromosome from an autosome.[17]

Inversions can be involved in speciation in multiple ways. Since heterozygote inversions can be underdominant, they can cause hybrid fitness loss, resulting in post-zygotic isolation. They can also accumulate selected differences between species, causing both pre- and post-zygotic isolation.[16]

Inversions often form geographical clines in frequency which can hint to their role in local adaptation. A prominent instance of such a cline is inversion 3RP in Drosophila melanogaster that can be observed in three different continents.[16] When an inversion contains two or more locally adaptive alleles, it can be selected and spread. For example; in the butterfly Heliconius numata, 18 genes controlling colors are linked together by inversions as together they confer higher fitness.[18]

Nomenclature edit

 
Three chromosomal abnormalities with ISCN nomenclature, with increasing complexity: (A) A tumour karyotype in a male with loss of the Y chromosome, (B) Prader–Willi Syndrome i.e. deletion in the 15q11-q12 region and (C) an arbitrary karyotype that involves a variety of autosomal and allosomal abnormalities including inversions (abbreviated as inv).[19]
 
Human karyotype with annotated bands and sub-bands as used for the nomenclature of chromosome abnormalities. It shows dark and white regions as seen on G banding. Each row is vertically aligned at centromere level. It shows 22 homologous autosomal chromosome pairs as well as both the female (XX) and male (XY) versions of the two sex chromosomes.
Further information: Karyotype

The International System for Human Cytogenomic Nomenclature (ISCN) is an international standard for human chromosome nomenclature, which includes band names, symbols, and abbreviated terms used in the description of human chromosome and chromosome abnormalities. Abbreviations include inv for inversions.[20]

Notable cases edit

  • Brenden Adams: former holder of the Guinness World Record for tallest teenager. His height is caused by an inversion of chromosome 12.
  • An example of chromosomal Inversion in organisms is demonstrated in the insect, Coelopa frigida. This particular species of Coelopa have a variation of chromosomal inversions that allow the species to create a series of physical differences. Individual C. frigida that are larger do not undergo a chromosomal inversion, whereas individuals that are smaller do undergo a chromosomal inversion.

References edit

  1. ^ Corbett-Detig RB, Said I, Calzetta M, Genetti M, McBroome J, Maurer NW, et al. (December 2019). "Fine-Mapping Complex Inversion Breakpoints and Investigating Somatic Pairing in the Anopheles gambiae Species Complex Using Proximity-Ligation Sequencing". Genetics. 213 (4): 1495–1511. doi:10.1534/genetics.119.302385. PMC 6893396. PMID 31666292.
  2. ^ Porubsky D, Sanders AD, Höps W, Hsieh P, Sulovari A, Li R, et al. (August 2020). "Recurrent inversion toggling and great ape genome evolution". Nature Genetics. 52 (8): 849–858. doi:10.1038/s41588-020-0646-x. PMC 7415573. PMID 32541924.
  3. ^ a b c Wellenreuther M, Bernatchez L (June 2018). "Eco-Evolutionary Genomics of Chromosomal Inversions". Trends in Ecology & Evolution. 33 (6): 427–440. doi:10.1016/j.tree.2018.04.002. PMID 29731154. S2CID 22051290.
  4. ^ Huang K, Rieseberg LH (2020). "Frequency, Origins, and Evolutionary Role of Chromosomal Inversions in Plants". Frontiers in Plant Science. 11: 296. doi:10.3389/fpls.2020.00296. PMC 7093584. PMID 32256515.
  5. ^ Kirkpatrick M (September 2010). "How and why chromosome inversions evolve". PLOS Biology. 8 (9): e1000501. doi:10.1371/journal.pbio.1000501. PMC 2946949. PMID 20927412.
  6. ^ Huang K, Rieseberg LH (2020). "Frequency, Origins, and Evolutionary Role of Chromosomal Inversions in Plants". Frontiers in Plant Science. 11: 296. doi:10.3389/fpls.2020.00296. PMC 7093584. PMID 32256515.
  7. ^ Kirkpatrick M (September 2010). "How and why chromosome inversions evolve". PLOS Biology. 8 (9): e1000501. doi:10.1371/journal.pbio.1000501. PMC 2946949. PMID 20927412.
  8. ^ Gardner R, Sutherland GR, Shaffer LG (2011). "9 Inversions". Chromosome Abnormalities and Genetic Counseling (4th ed.). Oxford University Press. pp. 161–182. ISBN 978-0-19-974915-7.
  9. ^ Kirkpatrick M (September 2010). "How and why chromosome inversions evolve". PLOS Biology. 8 (9): e1000501. doi:10.1371/journal.pbio.1000501. PMC 2946949. PMID 20927412.
  10. ^ Huang K, Rieseberg LH (2020). "Frequency, Origins, and Evolutionary Role of Chromosomal Inversions in Plants". Frontiers in Plant Science. 11: 296. doi:10.3389/fpls.2020.00296. PMC 7093584. PMID 32256515.
  11. ^ Huang K, Rieseberg LH (2020). "Frequency, Origins, and Evolutionary Role of Chromosomal Inversions in Plants". Frontiers in Plant Science. 11: 296. doi:10.3389/fpls.2020.00296. PMC 7093584. PMID 32256515.
  12. ^ a b "Concepts of Genetics". www.pearson.com. Retrieved 2022-12-05.
  13. ^ a b Faria R, Johannesson K, Butlin RK, Westram AM (March 2019). "Evolving Inversions". Trends in Ecology & Evolution. 34 (3): 239–248. doi:10.1016/j.tree.2018.12.005. PMID 30691998. S2CID 59339762.
  14. ^ Rieseberg, L.H. (2001). Chromosomal rearrangements and speciation. Trends in Ecology & Evolution, 16, 351-358.
  15. ^ Roesti, M., Gilbert, K. J., & Samuk, K. (2022). Chromosomal inversions can limit adaptation to new environments. Molecular Ecology, 31, 4435–4439. https://doi.org/10.1111/mec.16609
  16. ^ a b c d Kirkpatrick M (September 2010). "How and why chromosome inversions evolve". PLOS Biology. 8 (9): e1000501. doi:10.1371/journal.pbio.1000501. PMC 2946949. PMID 20927412.
  17. ^ van Doorn GS, Kirkpatrick M (October 2007). "Turnover of sex chromosomes induced by sexual conflict". Nature. 449 (7164): 909–912. Bibcode:2007Natur.449..909V. doi:10.1038/nature06178. PMID 17943130. S2CID 4301225.
  18. ^ Joron M, Frezal L, Jones RT, Chamberlain NL, Lee SF, Haag CR, et al. (August 2011). "Chromosomal rearrangements maintain a polymorphic supergene controlling butterfly mimicry". Nature. 477 (7363): 203–206. Bibcode:2011Natur.477..203J. doi:10.1038/nature10341. PMC 3717454. PMID 21841803.
  19. ^ Warrender JD, Moorman AV, Lord P (December 2019). "A fully computational and reasonable representation for karyotypes". Bioinformatics. 35 (24): 5264–5270. doi:10.1093/bioinformatics/btz440. PMC 6954653. PMID 31228194.
    - "This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0/)"
  20. ^ "ISCN Symbols and Abbreviated Terms". Coriell Institute for Medical Research. Retrieved 2022-10-27.

Further reading edit

  • Lehtonen S, Myllys L, Huttunen S (2009). "Phylogenetic analysis of non-coding plastid DNAthtjtdjj in the presence of short inversions" (PDF-preview). Phytotaxa. 1: 3–20. doi:10.11646/phytotaxa.1.1.2.
 
Wikimedia Commons has media related to Chromosome inversion.

January 01, 1970
chromosomal, inversion, inversion, chromosome, rearrangement, which, segment, chromosome, becomes, inverted, within, original, position, inversion, occurs, when, chromosome, undergoes, breaks, within, chromosomal, segment, between, breaks, inserts, itself, opp. An inversion is a chromosome rearrangement in which a segment of a chromosome becomes inverted within its original position An inversion occurs when a chromosome undergoes a two breaks within the chromosomal arm and the segment between the two breaks inserts itself in the opposite direction in the same chromosome arm The breakpoints of inversions often happen in regions of repetitive nucleotides and the regions may be reused in other inversions 1 Chromosomal segments in inversions can be as small as 1 kilobases or as large as 100 megabases 2 The number of genes captured by an inversion can range from a handful of genes to hundreds of genes 3 Inversions can happen either through ectopic recombination between repetitive sequences or through chromosomal breakage followed by non homologous end joining 4 Inversions are of two types paracentric and pericentric Paracentric inversions do not include the centromere and both breakpoints occur in one arm of the chromosome Pericentric inversions span the centromere and there is a breakpoint in each arm 5 A clay model showing why heterozygous inversion loops are visible in polytene chromosome preparations An inversion loop in the A arm of a chromosome from an Axarus species midge Inversions usually do not cause any abnormalities in carriers as long as the rearrangement is balanced with no extra or missing DNA However in individuals which are heterozygous for an inversion there is an increased production of abnormal chromatids this occurs when crossing over occurs within the span of the inversion This leads to lowered fertility due to production of unbalanced gametes Inversions do not involve either loss or gain of genetic information they simply rearrange the linear DNA sequence Contents 1 Detection 2 History 3 Effects on recombination 4 Evolutionary consequences 5 Nomenclature 6 Notable cases 7 References 8 Further readingDetection editCytogenetic techniques may be able to detect inversions or inversions may be inferred from genetic analysis Nevertheless in most species small inversions go undetected More recently comparative genomics has been used to detect chromosomal inversions by mapping the genome 6 7 Population genomics may also be used to detect inversions using areas of high linkage disequilibrium as indicators for possible inversion sites Human families that may be carriers of inversions may be offered genetic counseling and genetic testing 8 History editThe first evidence of a chromosomal inversion was found in 1921 by Alfred Sturtevant in Drosophila melanogaster 9 Since then inversions have been found in a all eukaryotes 3 When discovered by Sturtevant inversions were regarded as areas of recombination suppression Originally these inversions were noted in polytene chromosomes within the salivary glands of heterozygous Drosophila melanogaster larvae 3 In 1970 Theodosius Dobzhansky noted that genes within an inversion had higher fitness than those that are found outside of the inversions although this is an area that needs further study 10 One of the more recent models of inversions is the Kirkpatrick and Barton Model 2006 which states that inversions are selectively advantageous by linking together adaptive alleles By physically linking co adapted variants at multiple genes into distinct versions haplotypes of an inversion selection should be more efficient in driving these variants to high frequency in a population This is in contrast to non inverted regions which may allow adaptive and maladaptive alleles to be carried 11 Effects on recombination editWhen an inversion carrying chromosome is paired with a non inverted homologous chromosome Inversion heterozygotes during meiosis they fail to synapse properly and inversion loops are formed A crossing over within the loop can produce unbalanced gametes In a paracentric inversion recombination results in one dicentric chromatid and one acentric chromatid During Anaphase both recombinants are faced with problems The acentric chromatid is pulled to one pole or the other and the dicentric recombinant generates dicentric bridges as it is pulled in two directions 12 In a pericentric inversion similar imbalanced chromosomes are produced The recombinant chromosomes resulting from these crosses include deletions and duplications The offspring produced by such gametes are mostly inviable and therefore recombination is indirectly suppressed within inverted regions 12 Evolutionary consequences editThe suppressed recombination between inversion heterozygotes provides an opportunity for the independent evolution of the ancestral and inverted arrangements At the beginning the inverted arrangement lacks variation while the ancestral one does not If the inverted haplotype is not lost eg due to drift the variation in the inverted arrangement can increase over time and recombination rate in the inverted region is somewhat restored as more homozygotes are introduced 13 Chromosomal inversions have gained a lot of attention in evolutionary research due to their potential role in local adaptation and speciation Because non recombining inversion haplotypes may harbor multiple co adapted gene variants inversions are thought to facilitate local adaptation to different environments because natural selection is more efficient in driving such linked adaptive variants to high frequency within a population 14 However empirically demonstrating the presence of linked co adapted gene variants within inversions is difficult because inversion haplotypes do not recombine Moreover this possible positive effect of chromosomal inversions for adaptation to different environments rests on the assumption that adaptive gene variants linked into distinct inversion haplotypes are indeed co adapted This idea is however likely violated in situations where populations experience spatially or temporally varying selection Because of fluctuating selection on inversion linked variants the absence of recombination between inversion haplotypes harboring distinct gene variants may then constrain rather than help adaptation to distinct environments 15 The importance of chromosomal inversions in adaptation to different environments therefore remains an open empirical problem in evolutionary genetics Inversion polymorphism can be established in two ways Genetic drift or selection can result in fixation of an inversion in a local population Inversion polymorphism can result from gene flow between this population and a population without the inversion Balancing selection can also result in inversion polymorphism by frequency dependence or overdominance 13 The fitness differences between the inverted and the ancestral chromosome can either produce a stable polymorphism or can result in the fixation of one or the other chromosome 16 Inversions have been essential to sex chromosome evolution In mammals the Y chromosome is unable to recombine with the X chromosome almost along its entire length This non recombining portion results from a series of inversions that overlap Decreased recombination rate between sex determining loci and sex anatagonistic genes is favored by selection This causes linkage disequilibrium between the male determining locus and an allele at another locus that is beneficial to males This can happen through inversions resulting in a non recombining block including both loci as is the case in the mammalian Y chromosome 16 Inversions can also be essential in the origination of new sex chromosomes They can cause linkage disequilibrium between a sex determining mutation and sex antagonistic loci and create a new sex chromosome from an autosome 17 Inversions can be involved in speciation in multiple ways Since heterozygote inversions can be underdominant they can cause hybrid fitness loss resulting in post zygotic isolation They can also accumulate selected differences between species causing both pre and post zygotic isolation 16 Inversions often form geographical clines in frequency which can hint to their role in local adaptation A prominent instance of such a cline is inversion 3RP in Drosophila melanogaster that can be observed in three different continents 16 When an inversion contains two or more locally adaptive alleles it can be selected and spread For example in the butterfly Heliconius numata 18 genes controlling colors are linked together by inversions as together they confer higher fitness 18 Nomenclature edit nbsp Three chromosomal abnormalities with ISCN nomenclature with increasing complexity A A tumour karyotype in a male with loss of the Y chromosome B Prader Willi Syndrome i e deletion in the 15q11 q12 region and C an arbitrary karyotype that involves a variety of autosomal and allosomal abnormalities including inversions abbreviated as inv 19 nbsp Human karyotype with annotated bands and sub bands as used for the nomenclature of chromosome abnormalities It shows dark and white regions as seen on G banding Each row is vertically aligned at centromere level It shows 22 homologous autosomal chromosome pairs as well as both the female XX and male XY versions of the two sex chromosomes Further information Karyotype The International System for Human Cytogenomic Nomenclature ISCN is an international standard for human chromosome nomenclature which includes band names symbols and abbreviated terms used in the description of human chromosome and chromosome abnormalities Abbreviations include inv for inversions 20 Notable cases editBrenden Adams former holder of the Guinness World Record for tallest teenager His height is caused by an inversion of chromosome 12 An example of chromosomal Inversion in organisms is demonstrated in the insect Coelopa frigida This particular species of Coelopa have a variation of chromosomal inversions that allow the species to create a series of physical differences Individual C frigida that are larger do not undergo a chromosomal inversion whereas individuals that are smaller do undergo a chromosomal inversion References edit Corbett Detig RB Said I Calzetta M Genetti M McBroome J Maurer NW et al December 2019 Fine Mapping Complex Inversion Breakpoints and Investigating Somatic Pairing in the Anopheles gambiae Species Complex Using Proximity Ligation Sequencing Genetics 213 4 1495 1511 doi 10 1534 genetics 119 302385 PMC 6893396 PMID 31666292 Porubsky D Sanders AD Hops W Hsieh P Sulovari A Li R et al August 2020 Recurrent inversion toggling and great ape genome evolution Nature Genetics 52 8 849 858 doi 10 1038 s41588 020 0646 x PMC 7415573 PMID 32541924 a b c Wellenreuther M Bernatchez L June 2018 Eco Evolutionary Genomics of Chromosomal Inversions Trends in Ecology amp Evolution 33 6 427 440 doi 10 1016 j tree 2018 04 002 PMID 29731154 S2CID 22051290 Huang K Rieseberg LH 2020 Frequency Origins and Evolutionary Role of Chromosomal Inversions in Plants Frontiers in Plant Science 11 296 doi 10 3389 fpls 2020 00296 PMC 7093584 PMID 32256515 Kirkpatrick M September 2010 How and why chromosome inversions evolve PLOS Biology 8 9 e1000501 doi 10 1371 journal pbio 1000501 PMC 2946949 PMID 20927412 Huang K Rieseberg LH 2020 Frequency Origins and Evolutionary Role of Chromosomal Inversions in Plants Frontiers in Plant Science 11 296 doi 10 3389 fpls 2020 00296 PMC 7093584 PMID 32256515 Kirkpatrick M September 2010 How and why chromosome inversions evolve PLOS Biology 8 9 e1000501 doi 10 1371 journal pbio 1000501 PMC 2946949 PMID 20927412 Gardner R Sutherland GR Shaffer LG 2011 9 Inversions Chromosome Abnormalities and Genetic Counseling 4th ed Oxford University Press pp 161 182 ISBN 978 0 19 974915 7 Kirkpatrick M September 2010 How and why chromosome inversions evolve PLOS Biology 8 9 e1000501 doi 10 1371 journal pbio 1000501 PMC 2946949 PMID 20927412 Huang K Rieseberg LH 2020 Frequency Origins and Evolutionary Role of Chromosomal Inversions in Plants Frontiers in Plant Science 11 296 doi 10 3389 fpls 2020 00296 PMC 7093584 PMID 32256515 Huang K Rieseberg LH 2020 Frequency Origins and Evolutionary Role of Chromosomal Inversions in Plants Frontiers in Plant Science 11 296 doi 10 3389 fpls 2020 00296 PMC 7093584 PMID 32256515 a b Concepts of Genetics www pearson com Retrieved 2022 12 05 a b Faria R Johannesson K Butlin RK Westram AM March 2019 Evolving Inversions Trends in Ecology amp Evolution 34 3 239 248 doi 10 1016 j tree 2018 12 005 PMID 30691998 S2CID 59339762 Rieseberg L H 2001 Chromosomal rearrangements and speciation Trends in Ecology amp Evolution 16 351 358 Roesti M Gilbert K J amp Samuk K 2022 Chromosomal inversions can limit adaptation to new environments Molecular Ecology 31 4435 4439 https doi org 10 1111 mec 16609 a b c d Kirkpatrick M September 2010 How and why chromosome inversions evolve PLOS Biology 8 9 e1000501 doi 10 1371 journal pbio 1000501 PMC 2946949 PMID 20927412 van Doorn GS Kirkpatrick M October 2007 Turnover of sex chromosomes induced by sexual conflict Nature 449 7164 909 912 Bibcode 2007Natur 449 909V doi 10 1038 nature06178 PMID 17943130 S2CID 4301225 Joron M Frezal L Jones RT Chamberlain NL Lee SF Haag CR et al August 2011 Chromosomal rearrangements maintain a polymorphic supergene controlling butterfly mimicry Nature 477 7363 203 206 Bibcode 2011Natur 477 203J doi 10 1038 nature10341 PMC 3717454 PMID 21841803 Warrender JD Moorman AV Lord P December 2019 A fully computational and reasonable representation for karyotypes Bioinformatics 35 24 5264 5270 doi 10 1093 bioinformatics btz440 PMC 6954653 PMID 31228194 This is an Open Access article distributed under the terms of the Creative Commons Attribution License http creativecommons org licenses by 4 0 ISCN Symbols and Abbreviated Terms Coriell Institute for Medical Research Retrieved 2022 10 27 Further reading editLehtonen S Myllys L Huttunen S 2009 Phylogenetic analysis of non coding plastid DNAthtjtdjj in the presence of short inversions PDF preview Phytotaxa 1 3 20 doi 10 11646 phytotaxa 1 1 2 nbsp Wikimedia Commons has media related to Chromosome inversion Retrieved from https en wikipedia org w index php title Chromosomal inversion amp oldid 1213903391, wikipedia, wiki, book, books, library,

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