Directional selection

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Three different types of genetic selection. On each graph, the x-axis variable is the type of phenotypic trait and the y-axis variable is the amount of organisms. Group A is the original population and Group B is the population after selection. Top (Graph 1) represents directional selection with one extreme favored. Middle (Graph 2) represents stabilizing selection with the moderate trait favored. Bottom (Graph 3) represents disruptive selection with both extremes being favored. Genetic Distribution.svg
Three different types of genetic selection. On each graph, the x-axis variable is the type of phenotypic trait and the y-axis variable is the amount of organisms. Group A is the original population and Group B is the population after selection. Top (Graph 1) represents directional selection with one extreme favored. Middle (Graph 2) represents stabilizing selection with the moderate trait favored. Bottom (Graph 3) represents disruptive selection with both extremes being favored.

In population genetics, directional selection is a type of natural selection in which one extreme phenotype is favored over both the other extreme and moderate phenotypes. This genetic selection causes the allele frequency to shift toward the chosen extreme over time as allele ratios change from generation to generation. The advantageous extreme allele will increase as a consequence of survival and reproduction differences among the different present phenotypes in the population. The allele fluctuations as a result of directional selection can be independent of the dominance of the allele, and in some cases if the allele is recessive, it can eventually become fixed in the population. [1] [2]

Contents

Directional selection was first identified and described by naturalist Charles Darwin in his book On the Origin of Species published in 1859. [3] He identified it as a type of natural selection along with stabilizing selection and disruptive selection. [4] These types of selection also operate by favoring a specific allele and influencing the population's future phenotypic ratio. Disruptive selection favors both extreme phenotypes while the moderate trait will be selected against. The frequency of both extreme alleles will increase while the frequency of the moderate allele will decrease, differing from the trend in directional selection when only one extreme allele is favored. Stabilizing selection favors the moderate phenotype and will select against both extreme phenotypes. [5] Directional selection can be observed in finch beak size, peppered moth color, African cichlid mouth types, and sockeye salmon migration periods.

If there is continuous allele frequencies changes as a result of directional selection generation to generation, there will be observable changes in the phenotypes of the entire population over time. Directional selection can change the genotypic and phenotypic variation of a population and cause a trend toward one specific phenotype. [6] This selection is an important mechanism in the selection of complex and diversifying traits, and is also a primary force of speciation. [7] Changes in a genotype and consequently a phenotype can either be advantageous, harmful, or neutral and depend on the environment in which the phenotypic shift is happening. [8]

Evidence

Detection methods

Directional selection most often occurs during environmental changes or population migrations to new areas with different environmental pressures. Directional selection allows for swift changes in allele frequency that can accompany rapidly changing environmental factors and plays a major role in speciation. [7] Analysis on quantitative trait locus (QTL) effects has been used to examine the impact of directional selection in phenotypic diversification. QTL is a region of a gene that corresponds to a specific phenotypic trait, and the measuring the statistical frequencies of the traits can be helpful in analyzing phenotypic trends. [9] In one study, the analysis showed that directional changes in QTLs affecting various traits were more common than expected by chance among diverse species. This was an indication that directional selection is a primary cause of the phenotypic diversification that can eventually result in speciation. [10]

There are different statistical tests that can be run to test for the presence of directional selection in a population. A highly indicative test of changes in allele frequencies is the QTL sign test, and other tests include the Ka/Ks ratio test and the relative rate test. The QTL sign test compares the number of antagonistic QTL to a neutral model, and allows for testing of directional selection against genetic drift. [11] The Ka/Ks ratio test compares the number of non-synonymous to synonymous substitutions, and a ratio that is greater than 1 indicates directional selection. [12] The relative ratio test looks at the accumulation of advantageous traits against a neutral model, but needs a phylogenetic tree for comparison. This can prove difficult if the full phylogenic history is not known or is not specific enough for the test comparison. [13]

Examples

Finch beak size

Another example of directional selection is the beak size in a specific population of finches. Darwin first observed this in the publication of his book, On the Origin of Species, and he details how the size of the finches beak differs based on environmental factors. On the Galápagos Islands west of the coast of Ecuador, there were groups of finches displaying different beak phenotypes. [14] In one group, the beaks ranged from large and tough to small and smooth. Throughout the wet years, small seeds were more common than large seeds, and because of the large supply of small seeds the finches rarely ate large seeds. During the dry years, neither the small or large seeds were in great abundance, and the birds trended towards eating larger seeds. The changes in diet of the finches based off the environmental wet and dry seasons affected the depth of the birds’ beaks in future generations. [15] The beaks most beneficial to the more plentiful type of seed would be selected for because the birds were able to feed themselves and reproduce.

Peppered moths

Peppered moth with dark phenotype that was positively selected for during the Industrial Revolution. Biston betularia (4777708667).jpg
Peppered moth with dark phenotype that was positively selected for during the Industrial Revolution.

A significant example of directional selection in populations is the fluctuations of light and dark phenotypes in peppered moths in the 1800s. [16] During the industrial revolution, environmental conditions were rapidly changing with the newfound emission of dark, black smoke from factories that would change the color of trees, rocks, and other niches of moths. [17] Before the industrial revolution, the most prominent phenotype in the peppered moth population was the lighter, speckled moths. They thrived on the light birch trees and their phenotype would provide them with better camouflage from predators. After the Industrial Revolution as the trees become darker with soot, the moths with the darker phenotype were able to blend in and avoid predators better than their white counterparts. As time went on, the darker moths were positively, directional selected for and the allele frequencies start to shift with the increase in number of the darker moths. [18]

African cichlids

African cichlids are known to be a diverse fish species and evidence indicates that they evolved extremely quickly. These fish evolved within the same habitat, but have a variety of morphologies, especially pertaining to the mouth and jaw. Experiments pertaining the cichlid jaw phenotypes was tested by Albertson and others in 2003 by crossing two species of African cichlids with very different mouth morphologies. The cross between Labeotropheus fuelleborni (subterminal mouth for biting algae off rocks) and Metriaclima zebra (terminal mouth for suction feeding) allowed for mapping of QTLs affecting feeding morphology. Using the QTL sign test, definitive evidence was used to support the existence of directional selection in the oral jaw apparatus in African cichlids. However, this was not the case for the suspensorium or skull QTLs, suggesting genetic drift or stabilizing selection as mechanisms for the speciation. [19]

Sockeye salmon

Sockeye salmon are one of the many species of fish that are anadromous, in which individuals migrate to the same rivers in which they were born to reproduce. These migrations happen around the same time every year, but a 2007 study shows that sockeye salmon found in the waters of the Bristol Bay in Alaska have recently undergone directional selection on the timing of migration. [20] In this study, two populations of sockeye salmon, Egegik and Ugashik, were observed. Data from 1969–2003 provided by the Alaska Department of Fish and Game were divided into five sets of seven years and plotted for average arrival to the fishery. After analyzing the data, it was determined that in both populations average migration date was earlier and the populations were undergoing directional selection as a result of changing ecological conditions. The Egegik population experienced stronger selection and the migration date shifted four days. The paper suggests that fisheries can be a factor driving this selection because fishing occurs more in the later periods of migration (especially in the Egegik district), preventing those fish from reproducing. [21] This discovery also goes to show that in addition to environmental changes, human behaviors can also have massive effects on the selection of species around them.

Ecological impact

Directional selection can quickly lead to vast changes in allele frequencies in a population because of the cumulative nature of reproduction of the fittest. Because the main cause for directional selection is different and changing environmental pressures, rapidly changing environments, such as climate change, can cause drastic changes within populations.

Limiting the number of genotypes in a certain population can be deleterious to the ecosystem as a whole by shrink the potential genetic gene pool. [22] Low amount of genetic variation can lead to mass extinctions and endangered species because of the large impact one mutation can have on the entire population if there are only a few specific genes present throughout. It is important to note the impact that humans have on genetic diversity as well, and be aware of the ways to reduce harmful impacts on natural environments. Major roads, waterway pollution, and urbanization all cause environmental selection and could potentially result in changes in allele frequencies. [23]

Timescale

Typically directional selection acts strongly for short bursts and is not sustained over long periods of time. [24] If it was sustained, a population might hit biological constraints such that it no longer responds to selection. However, it is possible for directional selection to take a very long time to find a local optimum on a fitness landscape. [25] A possible example of long-term directional selection is the tendency of proteins to become more hydrophobic over time, [26] and to have their hydrophobic amino acids more interspersed along the sequence. [27]

See also

Related Research Articles

An allele, or allelomorph, is a variant of the sequence of nucleotides at a particular location, or locus, on a DNA molecule.

<span class="mw-page-title-main">Natural selection</span> Mechanism of evolution by differential survival and reproduction of individuals

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. Charles Darwin popularised the term "natural selection", contrasting it with artificial selection, which is intentional, whereas natural selection is not.

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, speciation, and population structure.

<span class="mw-page-title-main">Polymorphism (biology)</span> Occurrence of two or more clearly different morphs or forms in the population of a species

In biology, polymorphism is the occurrence of two or more clearly different morphs or forms, also referred to as alternative phenotypes, in the population of a species. To be classified as such, morphs must occupy the same habitat at the same time and belong to a panmictic population.

A quantitative trait locus (QTL) is a locus that correlates with variation of a quantitative trait in the phenotype of a population of organisms. QTLs are mapped by identifying which molecular markers correlate with an observed trait. This is often an early step in identifying the actual genes that cause the trait variation.

<span class="mw-page-title-main">Stabilizing selection</span> Type of selection in evolution where a trait stabilizes around the average value

Stabilizing selection is a type of natural selection in which the population 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 to select against extreme values of the character. Stabilizing selection is the opposite of disruptive selection. 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.

<span class="mw-page-title-main">Disruptive selection</span> Natural selection for extreme trait values over intermediate ones

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 increases and the population is divided into two distinct groups. In this more individuals acquire peripheral character value at both ends of the distribution curve.

Genetic architecture is the underlying genetic basis of a phenotypic trait and its variational properties. Phenotypic variation for quantitative traits is, at the most basic level, the result of the segregation of alleles at quantitative trait loci (QTL). Environmental factors and other external influences can also play a role in phenotypic variation. Genetic architecture is a broad term that can be described for any given individual based on information regarding gene and allele number, the distribution of allelic and mutational effects, and patterns of pleiotropy, dominance, and epistasis.

<span class="mw-page-title-main">Pleiotropy</span> Influence of a single gene on multiple phenotypic traits

Pleiotropy occurs when one gene influences two or more seemingly unrelated phenotypic traits. Such a gene that exhibits multiple phenotypic expression is called a pleiotropic gene. Mutation in a pleiotropic gene may have an effect on several traits simultaneously, due to the gene coding for a product used by a myriad of cells or different targets that have the same signaling function.

A polygene is a member of a group of non-epistatic genes that interact additively to influence a phenotypic trait, thus contributing to multiple-gene inheritance, a type of non-Mendelian inheritance, as opposed to single-gene inheritance, which is the core notion of Mendelian inheritance. The term "monozygous" is usually used to refer to a hypothetical gene as it is often difficult to distinguish the effect of an individual gene from the effects of other genes and the environment on a particular phenotype. Advances in statistical methodology and high throughput sequencing are, however, allowing researchers to locate candidate genes for the trait. In the case that such a gene is identified, it is referred to as a quantitative trait locus (QTL). These genes are generally pleiotropic as well. The genes that contribute to type 2 diabetes are thought to be mostly polygenes. In July 2016, scientists reported identifying a set of 355 genes from the last universal common ancestor (LUCA) of all organisms living on Earth.

Genetic association is when one or more genotypes within a population co-occur with a phenotypic trait more often than would be expected by chance occurrence.

Genetic divergence is the process in which two or more populations of an ancestral species accumulate independent genetic changes (mutations) through time, often leading to reproductive isolation and continued mutation even after the populations have become reproductively isolated for some period of time, as there is not any genetic exchange anymore. In some cases, subpopulations cover living in ecologically distinct peripheral environments can exhibit genetic divergence from the remainder of a population, especially where the range of a population is very large. The genetic differences among divergent populations can involve silent mutations or give rise to significant morphological and/or physiological changes. Genetic divergence will always accompany reproductive isolation, either due to novel adaptations via selection and/or due to genetic drift, and is the principal mechanism underlying speciation.

Genetic assimilation is a process described by Conrad H. Waddington by which a phenotype originally produced in response to an environmental condition, such as exposure to a teratogen, later becomes genetically encoded via artificial selection or natural selection. Despite superficial appearances, this does not require the (Lamarckian) inheritance of acquired characters, although epigenetic inheritance could potentially influence the result. Waddington stated that genetic assimilation overcomes the barrier to selection imposed by what he called canalization of developmental pathways; he supposed that the organism's genetics evolved to ensure that development proceeded in a certain way regardless of normal environmental variations.

In biology, a cline is a measurable gradient in a single characteristic of a species across its geographical range. Clines usually have a genetic, or phenotypic character. They can show either smooth, continuous gradation in a character, or more abrupt changes in the trait from one geographic region to the next.

In genetics, transgressive segregation is the formation of extreme phenotypes, or transgressive phenotypes, observed in segregated hybrid populations compared to phenotypes observed in the parental lines. The appearance of these transgressive (extreme) phenotypes can be either positive or negative in terms of fitness. If both parents' favorable alleles come together, it will result in a hybrid having a higher fitness than the two parents. The hybrid species will show more genetic variation and variation in gene expression than their parents. As a result, the hybrid species will have some traits that are transgressive (extreme) in nature. Transgressive segregation can allow a hybrid species to populate different environments/niches in which the parent species do not reside, or compete in the existing environment with the parental species.

Marker assisted selection or marker aided selection (MAS) is an indirect selection process where a trait of interest is selected based on a marker linked to a trait of interest, rather than on the trait itself. This process has been extensively researched and proposed for plant- and animal- breeding.

In genetics, association mapping, also known as "linkage disequilibrium mapping", is a method of mapping quantitative trait loci (QTLs) that takes advantage of historic linkage disequilibrium to link phenotypes to genotypes, uncovering genetic associations.

Quantitative trait loci mapping or QTL mapping is the process of identifying genomic regions that potentially contain genes responsible for important economic, health or environmental characters. Mapping QTLs is an important activity that plant breeders and geneticists routinely use to associate potential causal genes with phenotypes of interest. Family-based QTL mapping is a variant of QTL mapping where multiple-families are used.

<span class="mw-page-title-main">Genetic variance</span> Biological concept

Genetic variance is a concept outlined by the English biologist and statistician Ronald Fisher in his fundamental theorem of natural selection. In his 1930 book The Genetical Theory of Natural Selection, Fisher postulates that the rate of change of biological fitness can be calculated by the genetic variance of the fitness itself. Fisher tried to give a statistical formula about how the change of fitness in a population can be attributed to changes in the allele frequency. Fisher made no restrictive assumptions in his formula concerning fitness parameters, mate choices or the number of alleles and loci involved.

This glossary of genetics and evolutionary biology is a list of definitions of terms and concepts used in the study of genetics and evolutionary biology, as well as sub-disciplines and related fields, with an emphasis on classical genetics, quantitative genetics, population biology, phylogenetics, speciation, and systematics. Overlapping and related terms can be found in Glossary of cellular and molecular biology, Glossary of ecology, and Glossary of biology.

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Further reading