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nary consequences 5 nomenclature 6 notable cases 7 references 8 further reading toggle the table of contents chromosomal inversion 27 languages العربية bosanski català deutsch ελληνικά español فارسی galego magyar italiano 日本語 ქართული қазақша 한국어 монгол nederlands polski português русский srpskohrvatski српскохрватски simple english тоҷикӣ türkçe українська oʻzbekcha ўзбекча tiếng việt 中文 edit links article talk english read edit view history tools tools move to sidebar hide actions read edit view history general what links here related changes upload file permanent link page information cite this page get shortened url switch to legacy parser print export download as pdf printable version in other projects wikimedia commons wikidata item appearance move to sidebar hide from wikipedia the free encyclopedia chromosome rearrangement in which a segment of a chromosome is reversed 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 two breaks within the same 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 kilobase 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 some of the techniques that are used to detect inversions are karyotype analysis g banding fluorescence in situ hybridization 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 linkage disequilibrium refers to the non random association of alleles at different genetic loci when alleles combinations occur more frequently than expected under random assortment the loci are said to be in linkage disequilibrium in regions containing chromosomal inversions suppressed recombination can result in linkage disequilibrium 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 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 e g 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 citation needed his height is caused by an inversion of chromosome 12 citation needed an example of chromosomal inversion in organisms is demonstrated in the insect coelopa frigida citation needed this particular species of coelopa has a variation of chromosomal inversions that allow the species to create a series of physical differences citation needed individual c frigida that are larger do not undergo a chromosomal inversion whereas individuals that are smaller do undergo a chromosomal inversion citation needed 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 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 1 2 3 wellenreuther m bernatchez l june 2018 eco evolutionary genomics of chromosomal inversions trends in ecology evolution 33 6 427 440 bibcode 2018tecoe 33 427w 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 bibcode 2020frps 11 296h 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 bibcode 2020frps 11 296h 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 bibcode 2020frps 11 296h 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 bibcode 2020frps 11 296h doi 10 3389 fpls 2020 00296 pmc 7093584 pmid 32256515 1 2 concepts of genetics www pearson com archived from the original on 2022 12 05 retrieved 2022 12 05 1 2 faria r johannesson k butlin rk westram am march 2019 evolving inversions trends in ecology evolution 34 3 239 248 bibcode 2019tecoe 34 239f doi 10 1016 j tree 2018 12 005 pmid 30691998 s2cid 59339762 rieseberg l h 2001 chromosomal rearrangements and speciation trends in ecology evolution 16 351 358 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 1 2 3 4 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 term...