population genetics Population genetics is a subfield of genetics that deals with genetic differences within and between populations, and is a part of evolutionary biology. Studies in this branch of biology examine such phenomena as adaptation, speciation, and po ...

, directional selection, is a mode of negative natural selection in which an extreme

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 ...

is favored over other phenotypes, causing the

allele frequency Allele frequency, or gene frequency, is the relative frequency of an allele (variant of a gene) at a particular locus in a population, expressed as a fraction or percentage. Specifically, it is the fraction of all chromosomes in the population tha ...

to shift over time in the direction of that phenotype. Under directional selection, the advantageous allele increases as a consequence of differences in survival and reproduction among different phenotypes. The increases are independent of the dominance of the allele, and even if the allele is

recessive In genetics, dominance is the phenomenon of one variant ( allele) of a gene on a chromosome masking or overriding the effect of a different variant of the same gene on the other copy of the chromosome. The first variant is termed dominant an ...

, it will eventually become

fixed Fixed may refer to: * ''Fixed'' (EP), EP by Nine Inch Nails * ''Fixed'', an upcoming 2D adult animated film directed by Genndy Tartakovsky * Fixed (typeface), a collection of monospace bitmap fonts that is distributed with the X Window System * F ...

. Directional selection was first described by

Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended ...

in the book ''On the Origin of Species'' as a form of natural selection. Other types of natural selection include stabilizing and

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 ...

. Each type of selection contains the same principles, but is slightly different. Disruptive selection favors both extreme phenotypes, different from one extreme in directional selection. Stabilizing selection favors the middle phenotype, causing the decline in variation in a population over time.

Evidence

Directional selection occurs most often under environmental changes and when populations migrate to new areas with different environmental pressures. Directional selection allows for fast changes in allele frequency, and plays a major role in speciation. Analysis on QTL effects has been used to examine the impact of directional selection in phenotypic diversification. This analysis showed that the genetic loci correlating to directional selection was higher than expected; meaning directional selection is a primary cause of phenotypic diversification, which leads to speciation.

Detection methods

There are different statistical tests that can be run to test for the presence of directional selection in a population. A few of the tests include the QTL sign test, 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 Genetic drift, also known as allelic drift or the Wright effect, is the change in the frequency of an existing gene variant (allele) in a population due to random chance. Genetic drift may cause gene variants to disappear completely and there ...

. The

Ka/Ks ratio In genetics, the Ka/Ks ratio, also known as ω or ''d''N/''d''S ratio, is used to estimate the balance between neutral mutations, purifying selection and beneficial mutations acting on a set of homologous protein-coding genes. It is calculated as ...

test compares the number of non-synonymous to synonymous substitutions, and a ratio that is greater than 1 indicates directional selection. The relative ratio test looks at the accumulation of advantageous against a neutral model, but needs a phylogenetic tree for comparison.

Examples

An example of directional selection is fossil records that show that the size of the black bears in Europe decreased during interglacial periods of the ice ages, but increased during each glacial period. Another example is the beak size in a population of finches. Throughout the wet years, small seeds were more common and there was such a large supply of the small seeds that the finches rarely ate large seeds. During the dry years, none of the seeds were in great abundance, but the birds usually ate more large seeds. The change in diet of the finches affected the depth of the birds’ beaks in future generations. Their beaks range from large and tough to small and smooth.

African cichlids

African cichlids are known to be some of the most diverse fish and evolved extremely quickly. These fish evolved within the same habitat, but have a variety of morphologies, especially pertaining to the mouth and jaw. Albertson et al. 2003 tested this 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 feed) allowed for mapping of QTLs affecting feeding morphology. Using the QTL sign test definitive evidence was shown to prove that directional selection was occurring in the oral jaw apparatus. However, this was not the case for the suspensorium or skull (suggesting genetic drift or stabilizing selection).

Sockeye salmon

Sockeye salmon The sockeye salmon (''Oncorhynchus nerka''), also called red salmon, kokanee salmon, blueback salmon, or simply sockeye, is an anadromous species of salmon found in the Northern Pacific Ocean and rivers discharging into it. This species is a ...

are one of the many species of fish that are

anadromous Fish migration is mass relocation by fish from one area or body of water to another. Many types of fish migrate on a regular basis, on time scales ranging from daily to annually or longer, and over distances ranging from a few metres to thousa ...

. Individuals migrate to the same rivers in which they were born to reproduce. These migrations happen around the same time every year, but Quinn et al. 2007 shows that sockeye salmon found in the waters of the

Bristol Bay Bristol Bay ( esu, Iilgayaq, russian: Залив Бристольский) is the easternmost arm of the Bering Sea, at 57° to 59° North 157° to 162° West in Southwest Alaska. Bristol Bay is 400 km (250 mi) long and 290 km, ...

Alaska Alaska ( ; russian: Аляска, Alyaska; ale, Alax̂sxax̂; ; ems, Alas'kaaq; Yup'ik: ''Alaskaq''; tli, Anáaski) is a state located in the Western United States on the northwest extremity of North America. A semi-exclave of the U ...

have recently undergone directional selection on the timing of migration. In this study two populations of sockeye salmon were observed ( Egegik and Ugashik). Data from 1969-2003 provided by the

Alaska Department of Fish and Game The Alaska Department of Fish and Game (ADF&G) is a department within the government of Alaska. ADF&G's mission is to protect, maintain, and improve the fish, game, and aquatic plant resources of the state, and manage their use and development in ...

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 was undergoing directional selection. The Egegik population experienced stronger selection and shifted 4 days. Water temperature is thought to cause earlier migration date, but in this study there was no statistically significant correlation. 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.

Ecological impact

Directional selection can quickly lead to vast changes in allele frequencies in a population. 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.

Timescale

Typically directional selection acts strongly for short bursts and is not sustained over long periods of time. If it did, 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 even a

local optimum In applied mathematics and computer science, a local optimum of an optimization problem is a solution that is optimal (either maximal or minimal) within a neighboring set of candidate solutions. This is in contrast to a global optimum, which ...

on a

fitness landscape Fitness may refer to: * Physical fitness, a state of health and well-being of the body * Fitness (biology), an individual's ability to propagate its genes * Fitness (cereal), a brand of breakfast cereals and granola bars * ''Fitness'' (magazine), ...

. A possible example of long-term directional selection is the tendency of

protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, res ...

s to become more

hydrophobic In chemistry, hydrophobicity is the physical property of a molecule that is seemingly repelled from a mass of water (known as a hydrophobe). In contrast, hydrophiles are attracted to water. Hydrophobic molecules tend to be nonpolar and, ...

over time, and to have their hydrophobic amino acids more interspersed along the

sequence In mathematics, a sequence is an enumerated collection of objects in which repetitions are allowed and order matters. Like a set, it contains members (also called ''elements'', or ''terms''). The number of elements (possibly infinite) is called ...