Editing Disassortative mating

{{short description|Preferential mating pattern between individuals with dissimilar phenotypes (e.g., size, colour)}}
'''Disassortative mating''' (also known as '''negative assortative mating''' or '''heterogamy''') is a mating pattern in which individuals with dissimilar [[phenotypes]] mate with one another more frequently than would be expected under [[random mating]]. Disassortative mating reduces the mean genetic similarities within the population and produces a greater number of [[heterozygotes]]. The pattern is character specific, but does not affect allele frequencies.<ref>{{cite journal |last1=Lewontin |first1=Richard |last2=Kirk |first2=Dudley |last3=Crow |first3=James |title=Selective mating, assortative mating, and inbreeding: Definitions and implications |journal=Eugenics Quarterly |date=1963|volume=15 |issue=2 |pages=141–143 |doi=10.1080/19485565.1968.9987764 |pmid=5702329 }}</ref> This nonrandom mating pattern will result in deviation from the [[Hardy-Weinberg principle]] (which states that genotype frequencies in a population will remain constant from generation to generation in the absence of other evolutionary influences, such as "mate choice" in this case).

Disassortative mating is different from [[outbreeding]], which refers to mating patterns in relation to [[genotypes]] rather than phenotypes.

Due to homotypic preference (bias toward the same type), [[assortative mating]] occurs more frequently than disassortative mating.<ref>{{cite journal |last1=Thiessen |last2=Gregg |title=Human assortative mating and genetic equilibrium: An evolutionary perspective |journal=Ethology and Sociobiology |year=1980 |volume=1 |issue=2 |pages=111–140 |url=|doi=10.1016/0162-3095(80)90003-5}}</ref><ref>{{cite tech report |last1=Wallace |first1=B |title=The role of heterozygosity in drosophila populations |date=January 1958 |osti=4289507}}</ref> This is because homotypic preferences increase relatedness between mates and between parents and offspring that would promote cooperation and increases inclusive fitness. With disassortative mating, heterotypic preference (bias towards different types) in many cases has been shown to increase overall fitness.<ref>{{cite journal |last1=Burley |first1=Nancy |title=The meaning of assortative mating |journal=Ethology and Sociobiology |year=1983 |volume=4 |issue=4 |pages=191–203 |doi=10.1016/0162-3095(83)90009-2}}</ref> When this preference is favored, it allows a population to generate and/or maintain polymorphism (genetic variation within a population).

The fitness advantage aspect of disassortative mating seems straightforward, but the evolution of selective forces involved in disassortative mating are still largely unknown in natural populations.

== Types of disassortative mating ==
Imprinting is one example of disassortative mating. A model shows that individuals imprint on a genetically transmitted trait during early [[ontogeny]] and choosy females later use those parental images as a basis of mate choice. A viability-reducing trait may be maintained even without the fertility cost of same-type matings.<ref>{{cite journal |last1=Ihara |first1=Yasuo |last2=Feldman |first2=Marcus |title=Evolution of disassortative and assortative mating preferences based on imprinting |journal=Theoretical Population Biology |year=2003 |volume=64 |issue=2 |pages=193–200 |doi=10.1016/s0040-5809(03)00099-6 |pmid=12948680 |bibcode=2003TPBio..64..193I }}</ref> With imprinting, preference can be established even if it is initially rare, when there is a fertility cost of same-type matings.

One uncommon type of disassortative mating is the female preference on rare (or novel) male phenotypes. A study on guppies, ''Poecilia reticulata'', revealed that the female preference was sufficient to tightly maintain polymorphism in male traits.<ref>{{cite journal |last1=Kokko |first1=Hanna |last2=Jennions |first2=Michael |last3=Houde |first3=Anne |title=Evolution of frequency-dependent mate choice: keeping up with fashion trends |journal=Proceedings. Biological Sciences |year=2007 |volume=274 |issue=1615 |pages=1317–1324 |doi=10.1098/rspb.2007.0043 |pmid=17360285 |pmc=2176183 }}</ref> This type of mate choice shows that costly preferences can persist at higher frequencies if mate choice is hindered, which would allow the alleles to approach fixation.

== Effects ==
Disassortative mating may result in [[balancing selection]] and the maintenance of high genetic variation in the population. This is due to the excess heterozygotes that are produced from disassortative mating relative to a randomly mating population.

== In humans ==
The best-known example of disassortative mating in humans is preference for genes in the [[major histocompatibility complex]] (MHC) region on chromosome 6. Individuals feel more attracted to odors of individuals who are genetically different in this region.<ref>{{cite journal |last1=Wedekind |first1=Claus |title=MHC-dependent mate preferences in humans |journal=Proceedings of the Royal Society of London. Series B: Biological Sciences |year=1995 |volume=260 |issue=1359 |pages=245–249 |doi=10.1098/rspb.1995.0087 |pmid=7630893 |s2cid=34971350 |url=https://royalsocietypublishing.org/doi/abs/10.1098/rspb.1995.0087|url-access=subscription }}</ref> This promotes MHC heterozygosity in the children, making them less vulnerable to pathogens.

== In non-human species ==
Evidence from research regarding coloration in ''[[Heliconius]]'' butterflies suggests that disassortative mating is more likely to emerge when phenotypic variation is based on self-referencing (mate preference depends on phenotype of the choosing individual, therefore dominance in relationships influence the evolution of disassortative mating).<ref>{{cite journal |last1=Maisonneuve |first1=Ludovic |last2=Joron |first2=Mathieu |last3=Chouteau |first3=Mathieu |last4=Llaurens |first4=Violaine |title=Evolution and genetic architecture of disassortative mating at a locus under heterozygote advantage |journal=Evolution |year=2020 |volume=75 |issue=1 |pages=149–165 |doi=10.1111/evo.14129|biorxiv=10.1101/616409 |pmid=33210282|s2cid=227063195 }}</ref>

Disassortative mating has been found with traits such as body symmetry in [[Amphidromus inversus|''Amphridromus'' inversus]] snails. Normally in snails, rarely are individuals of the opposite coil able to mate with individuals of a normal coil pattern. However, it has been discovered that this species of snail frequents mating between individuals of opposing coils. It is said that the chirality of the spermatophore and the females reproductive tract have a greater chance of producing offspring.<ref>{{cite journal |last1=Schilthuizen |first1=M. |title=Sexual selection maintains whole-body chiral dimorphism in snails |year=2007 |journal=Journal of Evolutionary Biology |volume=20 |issue=5 |pages=1941–1949 |doi=10.1111/j.1420-9101.2007.01370.x |pmid=17714311 |pmc=2121153 }}</ref> This example of disassortative mating promotes polymorphism within the population.

In the scale eating predator fish, ''[[Perissodus microlepis]]'', disassortative mating allows the individuals with the rare phenotype of mouth-opening direction to have better success as predators.<ref>{{cite journal |last1=Hori |first1=Michio |title=Frequency-Dependent Natural Selection in the Handedness of Scale-Eating Cichlid Fish |year=1993 |volume=260 |issue=5105 |pages=216–219 |url=https://www.science.org/doi/10.1126/science.260.5105.216 |journal=Science |doi=10.1126/science.260.5105.216 |pmid=17807183 |bibcode=1993Sci...260..216H |s2cid=33113282 |url-access=subscription }}</ref>

House mice conduct disassortative mating as they prefer mates genetically dissimilar to themselves. Specifically, odor profiles in mice are strongly linked to genotypes at the MHC loci controlling changes in the immune response. When MHC-heterozygous offspring are produced, it enhances their immunocompetence because of their ability to recognize a large range of pathogens.<ref>{{cite journal |last1=Penn |first1=Dustin |last2=Potts |first2=Wayne |title=The Evolution of Mating Preferences and Major Histocompatibility Complex Genes |year=1999 |url=https://www.journals.uchicago.edu/doi/full/10.1086/303166 |journal=The American Naturalist|volume=153 |issue=2 |pages=145–164 |doi=10.1086/303166 |pmid=29578757 |bibcode=1999ANat..153..145P |s2cid=4398891 |url-access=subscription }}</ref> Thus, the mice tend to prefer providing "good genes" to their offspring so they will mate with individuals with differences at the MHC loci.

In the seaweed fly, ''[[Coelopa frigida]]'', heterozygotes at the locus [[alcohol dehydrogenase]] (Adh) have been shown to express better fitness by having higher larval density and relative viability.<ref>{{cite journal |last1=Butlin |first1=R |last2=Collins |first2=P |last3=Day |first3=T |title=The effect of larval density on an inversion polymorphism in the seaweed fly Coelopa frigida |year=1984 |journal=Heredity |volume=52 |issue=3 |pages=415–423 |doi=10.1038/hdy.1984.49 |s2cid=20675225 |doi-access=free }}</ref> Females displayed disassortative mating in respect to the Adh locus because they would only mate with males of the opposite Adh genotype.<ref>{{cite journal |last1=Day |first1=T |last2=Butlin |first2=R |title=Non-random mating in natural populations of the seaweed fly, Coelopa frigida |year=1987 |journal=Heredity |volume=58 |issue=2 |pages=213–220 |doi=10.1038/hdy.1987.35 |s2cid=24811609 |doi-access=free }}</ref> It is suspected that they do this to maintain genetic variation in the population. 

[[White-throated sparrow|White-throated sparrows]], ''Zonotrichia albicollis'', prefer strong disassortative mating behaviors regarding the color of their head stripe. The single locus that controls this expression is only observed in heterozygotes. Additionally, the heterozygote arrangement of chromosome 2 from disassortative mating produced offspring of high aggression which is shown to be a social behavior that allows them to dominate their opponents.<ref>{{cite journal |last1=Horton |first1=Brent |title=Behavioral Characterization of a White-Throated Sparrow Homozygous for the ZAL2m Chromosomal Rearrangement |year=2013 |journal=Behavior Genetics|volume=43 |issue=1 |pages=60–70 |doi=10.1007/s10519-012-9574-6 |pmid=23264208 |pmc=3552124 }}</ref>

== References ==
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[[Category:Mating]]
[[Category:Mating systems]]
[[Category:Population genetics]]
[[Category:Ecology]]