Chromosomal

inversion

An inversion is a chromosome rearrangement in which a segment of a chromosome becomes

invert ed wit hin it s original posit ion. An inversion occurs when a chromosome undergoes a t wo
breaks wit hin t he chromosomal arm, and t he segment bet ween t he t wo breaks insert s it self in
t he opposit e direct ion in t he same chromosome arm. The breakpoint s of inversions oft en happen
in regions of repet it ive nucleot ides, and t he regions may be reused in ot her inversions.[1]
Chromosomal segment s in inversions can be as small as 100 kilobases or as large as 100
megabases. The number of genes capt ured by an inversion can range from a handful of genes t o
hundreds of genes.[2] Inversions can happen eit her t hrough ect opic recombinat ion, chromosomal
breakage and repair, or non-homologous end joining.[3]

Inversions are of t wo t ypes: paracent ric and pericent ric. Paracent ric inversions do not include t he
cent romere, and bot h breakpoint s occur in one arm of t he chromosome. Pericent ric inversions
span t he cent romere, and t here is a breakpoint in each arm[4].
 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 abnormalit ies in carriers, as long as t he rearrangement is
balanced, wit h no ext ra or missing DNA. However, in individuals which are het erozygous for an
inversion, t here is an increased product ion of abnormal chromat ids (t his occurs when crossing-
over occurs wit hin t he span of t he inversion). This leads t o lowered fert ilit y, due t o product ion of
unbalanced gamet es. Inversions do not involve eit her loss or gain of genet ic informat ion; t hey
simply rearrange t he linear DNA sequence.

Detection
Cyt ogenet ic t echniques may be able t o det ect inversions, or inversions may be inferred from
genet ic analysis. Nevert heless, in most species, small inversions go undet ect ed. More recent ly,
comparat ive genomics has been used t o det ect chromosomal inversions, by mapping t he
genome.[5][6] Populat ion genomics may also be used t o det ect inversions, using areas of high
linkage disequilibrium as indicat ors for possible inversion sit es. Human families t hat may be
carriers of inversions may be offered genet ic counseling and genet ic t est ing.[7]
History
The first evidence of a chromosomal inversion was found in 1921 by Alfred St urt evant in
Drosophila melanogast er.[8] Since t hen, inversions have been found in a all eukaryot es.[2] When
discovered by St urt evant , inversions were regarded as areas of recombinat ion suppression.

Originally, t hese inversions were not ed in polyt ene chromosomes wit hin t he salivary glands of
het erozygous Drosophila melanogaster larvae.[2] In 1970, Theodosius Dobzhansky not ed t hat
genes wit hin an inversion had higher fit ness t han t hose t hat are found out side of t he inversions,
alt hough t his is an area t hat needs furt her st udy.[9]

One of t he more recent models of inversions is t he Kirkpat rick and Bart on Model (2006), which
st at es t hat inversions are select ively advant ageous by linking t oget her adapt ive alleles. By
physically linking co-adapt ed variant s at mult iple genes int o dist inct versions (haplot ypes) of an
inversion, select ion should be more efficient in driving t hese variant s t o high frequency in a
populat ion. This is in cont rast t o non-invert ed regions, which may allow adapt ive and maladapt ive
alleles t o be carried.[10]

Also, inversions may be used as det ect ors for global climat e change. In Drosophila melanogaster
, a st udy done in 2015 showed t hat a specific inversion (3R) may cont ribut e t o adapt ions t o
climat e change.[11] In t he species Drosophila subobscura, researchers have been able t o t rack
global climat e change by measuring t he magnit ude and direct ional shift in chromosome inversion
frequencies, relat ive t o t emperat ures at specific global sit es.[12]

Effects on recombination
When an inversion carrying chromosome is paired wit h a non-invert ed homologous chromosome
(Inversion het erozygot es) during meiosis, t hey fail t o synapse properly and inversion loops are
formed. A crossing-over wit hin t he loop can produce unbalanced gamet es. In a paracent ric
inversion, recombinat ion result s in one dicent ric chromat id and one acent ric chromat id. During
Anaphase, bot h recombinant s are faced wit h problems. The acent ric chromat id is pulled t o one
pole or t he ot her, and t he dicent ric recombinant generat es dicent ric bridges as it is pulled in t wo
direct ions.[13]
In a pericent ric inversion, similar imbalanced chromosomes are produced. The recombinant
chromosomes result ing from t hese crosses include delet ions and duplicat ions. The offspring
produced by such gamet es are most ly inviable, and t herefore, recombinat ion is indirect ly
suppressed wit hin invert ed regions.[13]

Evolutionary consequences
The suppressed recombinat ion bet ween inversion het erozygot es provides an opport unit y for t he
independent evolut ion of t he ancest ral and invert ed arrangement s. At t he beginning, t he invert ed
arrangement lacks variat ion, while t he ancest ral one does not . If t he invert ed haplot ype is not
lost (eg. due t o drift ), t he variat ion in t he invert ed arrangement can increase over t ime, and
recombinat ion rat e in t he invert ed region is somewhat rest ored as more homozygot es are
int roduced.[14]

Chromosomal inversions have gained a lot of at t ent ion in evolut ionary research due t o t heir
pot ent ial role in local adapt at ion and speciat ion. Because non-recombining inversion haplot ypes
may harbor mult iple co-adapt ed gene variant s, inversions are t hought t o facilit at e local
adapt at ion t o different environment s because nat ural select ion is more efficient in driving such
linked adapt ive variant s t o high frequency wit hin a populat ion.[15] However, empirically
demonst rat ing t he presence of linked, co-adapt ed gene variant s wit hin inversions is difficult
because inversion haplot ypes do not recombine. Moreover, t his possible posit ive effect of
chromosomal inversions for adapt at ion t o different environment s rest s on t he assumpt ion t hat
adapt ive gene variant s linked int o dist inct inversion haplot ypes are indeed co-adapt ed. This idea
is, however, likely violat ed in sit uat ions where populat ions experience spat ially or t emporally
varying select ion. Because of fluct uat ing select ion on inversion-linked variant s, t he absence of
recombinat ion bet ween inversion haplot ypes harboring dist inct gene variant s may t hen const rain
rat her t han help adapt at ion t o dist inct environment s.[16] The import ance of chromosomal
inversions in adapt at ion t o different environment s t herefore remains an open empirical problem in
evolut ionary genet ics.

Inversion polymorphism can be est ablished in t wo ways. Genet ic drift or select ion can result in
fixat ion of an inversion in a local populat ion. Inversion polymorphism can result from gene flow
bet ween t his populat ion and a populat ion wit hout t he inversion. Balancing select ion can also
result in inversion polymorphism by frequency dependence or overdominance.[14] The fit ness
differences bet ween t he invert ed and t he ancest ral chromosome can eit her produce a st able
polymorphism or can result in t he fixat ion of one or t he ot her chromosome.[17]
Inversions have been essent ial t o sex chromosome evolut ion. In mammals, t he Y chromosome is
unable t o recombine wit h t he X chromosome, almost along it s ent ire lengt h. This non-
recombining port ion result s from a series of inversions t hat overlap. Decreased recombinat ion
rat e bet ween sex det ermining loci and sex-anat agonist ic genes is favored by select ion. This
causes linkage disequilibrium bet ween t he male det ermining locus and an allele at anot her locus
t hat is beneficial t o males. This can happen t hrough inversions result ing in a non-recombining
block including bot h loci, as is t he case in t he mammalian Y chromosome.[17]

Inversions can also be essent ial in t he originat ion of new sex chromosomes. They can cause
linkage disequilibrium bet ween a sex-det ermining mut at ion and sex-ant agonist ic loci and creat e a
new sex chromosome from an aut osome.[18]

Inversions can be involved in speciat ion in mult iple ways. Since het erozygot e inversions can be
underdominant , t hey can cause hybrid fit ness loss, result ing in post -zygot ic isolat ion. They can
also accumulat e select ed differences bet ween species, causing bot h pre- and post -zygot ic
isolat ion.[17]

Inversions oft en form geographical clines in frequency which can hint t o t heir role in local
adapt at ion. A prominent inst ance of such a cline is inversion 3RP in Drosophila melanogaster t hat
can be observed in t hree different cont inent s.[17] When an inversion cont ains t wo or more locally
adapt ive alleles, it can be select ed and spread. For example; in t he but t erfly Heliconius numata,
18 genes cont rolling colors are linked t oget her by inversions as t oget her t hey confer higher
fit ness.[19]
Nomenclature

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).[20]
 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.

The Int ernat ional Syst em for Human Cyt ogenomic Nomenclat ure (ISCN) is an int ernat ional
st andard for human chromosome nomenclat ure, which includes band names, symbols, and
abbreviat ed t erms used in t he descript ion of human chromosome and chromosome
abnormalit ies. Abbreviat ions include inv for inversions.[21]
Notable cases

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.

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Further reading

Lehtonen S, Myllys L, Huttunen S (2009).

"Phylogenetic analysis of non-coding plastid
DNAthtjtdjj in the presence of short
inversions" (https://www.mapress.com/phyto
taxa/content/2009/f/p00001p020f.pdf)
(PDF-preview). Phytotaxa. 1: 3–20.
doi:10.11646/phytotaxa.1.1.2 (https://doi.or
g/10.11646%2Fphytotaxa.1.1.2) .

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