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Stabilizing selection (not to be confused with negative or purifying selection) is a type of
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 heritable traits characteristic of a population over generations. Cha ...
in which the
population Population typically refers to the number of people in a single area, whether it be a city or town, region, country, continent, or the world. Governments typically quantify the size of the resident population within their jurisdiction usi ...
mean stabilizes on a particular non-extreme trait value. This is thought to be the most common mechanism of action for natural selection because most traits do not appear to change drastically over time. Stabilizing selection commonly uses negative selection (a.k.a. purifying selection) to select against extreme values of the character. Stabilizing selection is the opposite of
disruptive selection 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 increases and the population ...
. Instead of favoring individuals with extreme phenotypes, it favors the intermediate variants. Stabilizing selection tends to remove the more severe phenotypes, resulting in the reproductive success of the norm or average phenotypes. This means that most common phenotype in the population is selected for and continues to dominate in
future generations Future generations are cohorts of hypothetical people not yet born. Future generations are contrasted with current and past generations, and evoked in order to encourage thinking about intergenerational equity. The moral patienthood of future g ...
.


History

The Ukrainian evolutionary biologist
Ivan Schmalhausen Ivan Ivanovich Schmalhausen (russian: Ива́н Ива́нович Шмальга́узен; April 23, 1884 – October 7, 1963) was a Ukrainian, Russian and later Soviet zoologist and evolutionary biologist of German descent. He developed the th ...
founded the theory of stabilizing selection, publishing a paper in Russian titled "Stabilizing selection and its place among factors of evolution" in 1941 and a monograph "Factors of Evolution: The Theory of Stabilizing Selection" in 1945.


Influence on population structure

Stabilizing selection causes the narrowing of the phenotypes seen in a population. This is because the extreme phenotypes are selected against, causing reduced survival in organisms with those traits. This results in a population consisting of fewer phenotypes, with most traits representing the mean value of the population. This narrowing of phenotypes causes a reduction in genetic diversity in a population. Maintaining genetic variation is essential for the survival of a population because it is what allows them to evolve over time. In order for a population to adapt to changing environmental conditions they must have enough genetic diversity to select for new traits as they become favorable.


Analyzing stabilizing selection

There are four primary types of data used to quantify stabilizing selection in a population. The first type of data is an estimation of fitness of different phenotypes within a single generation. Quantifying fitness in a single generation creates predictions for the expected fate of selection. The second type of data is changes in allelic frequencies or phenotypes across different generations. This allows quantification of change in prevalence of a certain phenotype, indicating the type of selection. The third type of data is differences in allelic frequencies across space. This compares selection occurring in different populations and environmental conditions. The fourth type of data is DNA sequences from the genes contributing to observes phenotypic differences. The combination of these four types of data allow population studies that can identify the type of selection occurring and quantify the extent of selection. However, a meta-analysis of studies that measured selection in the wild failed to find an overall trend for stabilizing selection. The reason can be that methods for detecting stabilizing selection are complex. They can involve studying the changes that causes natural selection in the mean and variance of the trait, or measuring fitness for a range of different
phenotype In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology (biology), morphology or physical form and structure, its Developmental biology, developmental proc ...
s under natural conditions and examining the relationship between these fitness measurements and the trait value, but analysis and interpretation of the results is not straightforward.


Examples

The most common form of stabilizing selection is based on phenotypes of a population. In phenotype based stabilizing selection, the mean value of a phenotype is selected for, resulting a decrease in the phenotypic variation found in a population.


Humans

Stabilizing selection is the most common form of nonlinear selection (non-directional) in humans. There are few examples of genes with direct evidence of stabilizing selection in humans. However, most quantitative traits (height, birthweight, schizophrenia) are thought to be under stabilizing selection, due to their polygenicity and the distribution of the phenotypes throughout human populations. * Birth Weight − A classic example of this is human birth weight. Babies of low weight lose heat more quickly and get ill from infectious diseases more easily, whereas babies of large body weight are more difficult to deliver through the pelvis. Infants of a more medium weight survive much more often. For the larger or smaller babies, the baby mortality rate is much higher. The bell curve of the human population peaks at a birth weight that the newly born babies exhibit the minimum death rate.


Plants

* Height − Another example of a trait, that might be acted on by stabilizing selection, is plant height. A plant that is too short may not be able to compete with other plants for sunlight. However, extremely tall plants may be more susceptible to wind damage. Combined, these two selection pressures select to maintain plants of medium height. The number of plants of medium height will increase while the numbers of short and tall plants will decrease. * Cacti Spine Number − Desert populations of spiny cacti experience predation by
peccaries A peccary (also javelina or skunk pig) is a medium-sized, pig-like hoofed mammal of the family Tayassuidae (New World pigs). They are found throughout Central and South America, Trinidad in the Caribbean, and in the southwestern area of North ...
, which consume the fleshy part of the cactus. This can be prevented by increasing the number of spines on the cactus. However, there is also a selection pressure in the opposite direction because there is a parasitic insect that will lay its eggs in spines if they are densely populated. This means that in order to manage both of these selection pressures the cacti experiences stabilizing selection to balance the appropriate number of spines to survive these different threats.


Insects

*Butterfly's Winged Eyespots − The African butterfly ''
Bicyclus anynana ''Bicyclus anynana'' (squinting bush brown) is a small brown butterfly in the family Nymphalidae, the most globally diverse family of butterflies. It is primarily found in eastern Africa from southern Sudan to Eswatini.eyespots. It has been suggested that the circular eyespots positioned on the wings are favoured functionally compared to other shapes and sizes. * Gall Size − The ''
Eurosta solidaginis The goldenrod gall fly (''Eurosta solidaginis''), also known as the goldenrod ball gallmaker, is a species of fly native to North America. The species is best known for the characteristic galls it forms on several species in the ''Solidago'', or ...
'' fly lays its eggs on the tip of plants, which then encase the larvae in a protective
gall Galls (from the Latin , 'oak-apple') or ''cecidia'' (from the Greek , anything gushing out) are a kind of swelling growth on the external tissues of plants, fungi, or animals. Plant galls are abnormal outgrowths of plant tissues, similar to be ...
. The size of this gall is under stabilizing selection, as determined by predation. These larvae are under threat from parasitic wasps, which lay a single egg in galls containing the flies. The single wasp offspring then consumes the fly larvae to survive. Therefore, a larger gall is favored to allow more places for larvae to hide from the wasp. However, larger galls attract a different type of predation from birds, as they can penetrate large galls with their beak. Therefore, the optimal gall is moderately sized in order to avoid predation from both birds and wasps.


Birds

* Clutch Size − The number of eggs laid by a female bird (clutch size) is typically under stabilizing selection. This is because the female must lay as many eggs as possible to maximize the number of offspring. However, they can only lay as many eggs as they can support with their own resources. Laying too many eggs could expend all of the energy of the mother bird causing her to die and the death of the chicks. Additionally, once the eggs hatch the mother must be able to obtain enough resources to keep all of the chicks alive. Therefore, the mother typically lays a moderate amount of eggs in order to increase offspring survival and maximize the number of offspring.


Mammals

* *The Siberian husky experiences stabilizing selection in terms of their leg muscles. These dogs have to have enough muscle in order to pull sleds and move quickly. However, they also must be light enough to stay on top of the snow. This means that the leg muscles of the husky are most fit when they are moderately sized, to balance their strength and their weight.


See also

*
Ambidirectional dominance Ambidirectional dominance occurs in a situation where multiple genes influence a phenotype and dominance is in different directions depending on the gene. For example, for gene A increased height is dominant, while for gene B decreased height is do ...
*
Assortative mating Assortative mating (also referred to as positive assortative mating or homogamy) is a mating pattern and a form of sexual selection in which individuals with similar phenotypes or genotypes mate with one another more frequently than would be ...
* Balancing selection *
Directional selection In population genetics, directional selection, is a mode of negative natural selection in which an extreme phenotype is favored over other phenotypes, causing the allele frequency to shift over time in the direction of that phenotype. Under dir ...
*
Disruptive selection 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 increases and the population ...
*
Frequency-dependent foraging by pollinators Frequency-dependent foraging is defined as the tendency of an individual to selectively forage on a certain species or morph based on its relative frequency within a population.Levin 1972. Low frequency disadvantage in the exploitation of pollin ...
*
Fluctuating selection Fluctuating selection is a mode of natural selection characterized by the fluctuation of the direction of selection on a given phenotype over a relatively brief period of evolutionary time. For example, a species of plant may come in two varieties ...
*
Hypergamy Hypergamy (colloquially referred to as "marrying up") is a term used in social science for the act or practice of a person marrying a spouse of higher caste or social status than themselves. The antonym "hypogamy" refers to the inverse: marryin ...
*
Koinophilia Koinophilia is an evolutionary hypothesis proposing that during sexual selection, animals preferentially seek mates with a minimum of unusual or mutant features, including functionality, appearance and behavior. Koinophilia intends to explain t ...


References

{{MolecularEvolution Selection Evolutionary biology tr:Dengelenmiş seçilim