Underdominance
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genetics Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinians, Augustinian ...
, underdominance, also known as homozygote advantage, heterozygote disadvantage, or negative overdominance," is the opposite of overdominance. It is the selection against the heterozygote, causing
disruptive selection In evolutionary biology, disruptive selection, also called diversifying selection, describes changes in population genetics in which extreme values for a trait are favored over intermediate values. In this case, the variance of the trait in ...
and divergent
genotype The genotype of an organism is its complete set of genetic material. Genotype can also be used to refer to the alleles or variants an individual carries in a particular gene or genetic location. The number of alleles an individual can have in a ...
s. Underdominance exists in situations where the heterozygotic
genotype The genotype of an organism is its complete set of genetic material. Genotype can also be used to refer to the alleles or variants an individual carries in a particular gene or genetic location. The number of alleles an individual can have in a ...
is inferior in fitness to either the dominant or recessive homozygotic genotype. Compared to examples of overdominance in actual populations, underdominance is considered more unstable and may lead to the fixation of either allele. An example of stable underdominance may occur in individuals who are heterozygotic for polymorphisms that would make them better suited for one of two niches. Consider a situation in which a population is completely homozygotic for an "A"
allele An allele is a variant of the sequence of nucleotides at a particular location, or Locus (genetics), locus, on a DNA molecule. Alleles can differ at a single position through Single-nucleotide polymorphism, single nucleotide polymorphisms (SNP), ...
, allowing exploitation of a particular resource. Eventually, a polymorphic "a" allele may be introduced into the population, resulting in an individual who is capable of exploiting a different resource. This would result in an "aa" homozygotic invasion of the population due to nonexistent competition of the unexploited resource. The frequency of "aa" individuals would increase until the abundance of the "a" resource begins to decline. Eventually, the "AA" and "aa" genotypes would reach equilibrium with each other, with "Aa" heterozygotic individuals potentially experiencing a reduced fitness compared to those individuals who are homozygotic for utilization of either resource. This example of underdominance is stable because any shift in equilibrium would result in selection for the rare allele due to increased resource abundance. This compensatory selection would ultimately return the dimorphic system to underdominant equilibrium.


Incidence in butterfly populations

An example of stable underdominance can be found in the African butterfly species '' Pseudacraea eurytus'', which utilizes
Batesian mimicry Batesian mimicry is a form of mimicry where a harmless species has evolved to imitate the warning signals of a harmful species directed at a predator of them both. It is named after the English naturalist Henry Walter Bates, who worked on butt ...
to escape
predation Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
. This species possesses two alleles which each confer an appearance similar to that of another local butterfly species that is toxic to its predator. Individuals who are heterozygous for this trait appear to be intermediate in appearance and thus experience increased predation and lowered overall fitness.


Uses in pest control

Models of stable underdominance have shown potential in driving the introduction of refractory genes into pest populations that are responsible for the spread of infective diseases such as
malaria Malaria is a Mosquito-borne disease, mosquito-borne infectious disease that affects vertebrates and ''Anopheles'' mosquitoes. Human malaria causes Signs and symptoms, symptoms that typically include fever, Fatigue (medical), fatigue, vomitin ...
and
dengue fever Dengue fever is a mosquito-borne disease caused by dengue virus, prevalent in tropical and subtropical areas. Asymptomatic infections are uncommon, mild cases happen frequently; if symptoms appear, they typically begin 3 to 14 days after i ...
. A refractory gene alone would not have higher fitness than the native genes, but engineered underdominance may prove effective as a mechanism to spread such a gene. In this model, two genetics constructs are introduced into two non-
homologous chromosome Homologous chromosomes or homologs are a set of one maternal and one paternal chromosome that pair up with each other inside a cell during meiosis. Homologs have the same genes in the same locus (genetics), loci, where they provide points along e ...
s. Each construct is lethal when expressed individually but can be suppressed by the other construct. In this way, individuals with only one of the two constructed genes (heterozygotes) are selected against, but homozygotes with both or neither construct are genetically healthy. Analysis of this model using simple population genetics shows that successful spread of refractory genes using this engineered underdominance is possible with relatively small release of the constructed genotype into the population. A similar system of manipulation of pest populations was achieved in a population of ''
Drosophila melanogaster ''Drosophila melanogaster'' is a species of fly (an insect of the Order (biology), order Diptera) in the family Drosophilidae. The species is often referred to as the fruit fly or lesser fruit fly, or less commonly the "vinegar fly", "pomace fly" ...
'' by using a knock-down/rescue system.{{Cite journal, last1=Reeves, first1=R. Guy, last2=Bryk, first2=Jaroslaw, last3=Altrock, first3=Philipp, last4=Denton, first4=Jai, last5=Reed, first5=Floyd, year=2014, title=First Steps towards Underdominant Genetic Transformation of Insect Populations, journal=PLOS ONE, volume=9, issue=5, pages=e97557, doi=10.1371/journal.pone.0097557, pmid=24844466, pmc=4028297, bibcode=2014PLoSO...997557R, doi-access=free The genetic construct in this system employs a dsRNAi knockdown of a C-reactive protein, RpL14, as well as a rescue element (a complete copy of the wild type RpL14 gene). Individuals that are heterozygous for this construct experience lowered fitness due to limited restoration of the RpL14 gene, which results in reduced female fertility and delayed development, along with various other mutations that ultimately lower fitness by 70-80%. This system of underdominance allowed manipulation of the population and ultimate fixation of the constructed genotype and has potential applications in a number of settings, including agriculture and the reduction of various pest-carried diseases.


See also

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Natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the Heredity, heritable traits characteristic of a population over generation ...
*
Disruptive selection In evolutionary biology, disruptive selection, also called diversifying selection, describes changes in population genetics in which extreme values for a trait are favored over intermediate values. In this case, the variance of the trait in ...
*
Heterozygote advantage A heterozygote advantage describes the case in which the heterozygous genotype has a higher relative fitness (biology), fitness than either the homozygous Dominance (genetics), dominant or homozygous recessive gene, recessive genotype. Loci exhib ...
*
Population genetics Population genetics is a subfield of genetics that deals with genetic differences within and among populations, and is a part of evolutionary biology. Studies in this branch of biology examine such phenomena as Adaptation (biology), adaptation, s ...


References


External links


Underdominance- ResearchGate



Population Genetics

The Stability of An Equilibrium and the Average Fitness of a Population

Chromosome Inversions, Local Adaptations and Speciation

Chromosomal Rearrangements cause Underdominant Pollen Sterility
Classical genetics