QST (genetics)
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In
quantitative genetics Quantitative genetics is the study of quantitative traits, which are phenotypes that vary continuously—such as height or mass—as opposed to phenotypes and gene-products that are Categorical variable, discretely identifiable—such as eye-col ...
, QST is a statistic intended to measure the degree of genetic differentiation among populations with regard to a
quantitative trait Complex traits are phenotypes that are controlled by two or more genes and do not follow Mendelian inheritance, Mendel's Law of Dominance. They may have a range of Gene expression, expression which is typically continuous. Both environmental and g ...
. It was developed by Ken Spitze in 1993. Its name reflects that QST was intended to be analogous to the
fixation index The fixation index (FST) is a measure of population differentiation due to genetic structure. It is frequently estimated from Polymorphism (biology), genetic polymorphism data, such as single-nucleotide polymorphisms (SNP) or Microsatellite (genet ...
for a single genetic locus (FST). QST is often compared with FST of neutral loci to test if variation in a quantitative trait is a result of divergent selection or genetic drift, an analysis known as QST–FST comparisons.


Calculation of QST


Equations

QST represents the proportion of variance among subpopulations, and its calculation is synonymous to FST developed by
Sewall Wright Sewall Green Wright ForMemRS HonFRSE (December 21, 1889March 3, 1988) was an American geneticist known for his influential work on evolutionary theory and also for his work on path analysis. He was a founder of population genetics alongside ...
. However, instead of using genetic differentiation, QST is calculated by finding the variance of a quantitative trait within and among subpopulations, and for the total population. Variance of a quantitative trait among populations (σ2GB) is described as: \sigma_^2 = (1-Q_)\sigma_T^2 And the variance of a quantitative trait within populations (σ2GW) is described as: \sigma _^2 = 2Q_\sigma_T^2 Where σ2T is the total genetic variance in all populations. Therefore, QST can be calculated with the following equation: Q_ = \frac


Assumptions

Calculation of QST is subject to several assumptions: populations must be in Hardy–Weinberg equilibrium, observed variation is assumed to be due to additive genetic effects only, selection and
linkage disequilibrium Linkage disequilibrium, often abbreviated to LD, is a term in population genetics referring to the association of genes, usually linked genes, in a population. It has become an important tool in medical genetics and other fields In defining LD, it ...
are not present, and the subpopulations exist within an island model.


QST–FST comparisons

QST–FST analyses often involve culturing organisms in consistent environmental conditions, known as common garden experiments, and comparing the
phenotypic In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology (physical form and structure), its developmental processes, its biochemical and physiological propert ...
variance to genetic variance. If QST is found to exceed FST, this is interpreted as evidence of divergent selection, because it indicates more differentiation in the trait than could be produced solely by
genetic drift Genetic drift, also known as random genetic drift, allelic drift or the Wright effect, is the change in the Allele frequency, frequency of an existing gene variant (allele) in a population due to random chance. Genetic drift may cause gene va ...
. If QST is less than FST, balancing selection is expected to be present. If the values of QST and FSTare equivalent, the observed trait differentiation could be due to genetic drift. Suitable comparison of QST and FST is subject to multiple ecological and evolutionary assumptions, and since the development of QST, multiple studies have examined the limitations and constrictions of QST–FST analyses. Leinonen et al. notes FST must be calculated with neutral loci, however over filtering of non-neutral loci can artificially reduce FSTvalues. Cubry et al. found QST is reduced in the presence of dominance, resulting in conservative estimates of divergent selection when QST is high, and inconclusive results of balancing selection when QST is low. Additionally, population structure can significantly impact QST–FST ratios. Stepping stone models, which can generate more evolutionary noise than island models, are more likely to experience type 1 errors. If a subset of populations act as sources, such as during
invasion An invasion is a Offensive (military), military offensive of combatants of one geopolitics, geopolitical Legal entity, entity, usually in large numbers, entering territory (country subdivision), territory controlled by another similar entity, ...
, weighting the genetic contributions of each population can increase detection of adaptation. In order to improve precision of QST analyses, more populations (>20) should be included in analyses.


QST applications in literature

Multiple studies have incorporated QST to separate effects 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 Heredity, heritable traits characteristic of a population over generation ...
and genetic drift, and QST is often observed to exceed FST, indicating local adaptation. In an
ecological restoration Ecological restoration, or ecosystem restoration, is the process of assisting the recovery of an ecosystem that has been degraded, damaged, destroyed or transformed. It is distinct from conservation in that it attempts to retroactively repair ...
study, Bower and Aitken used QST to evaluate suitable populations for seed transfer of whitebark pine. They found high QST values in many populations, suggesting local adaptation for cold-adapted characteristics. During an assessment of the invasive species, '' Brachypodium sylvaticum'', Marchini et al. found divergence between native and invasive populations during initial establishment in the invaded range, but minimal divergence during range expansion. In an examination of the common snapdragon (''
Antirrhinum majus ''Antirrhinum majus'', the common snapdragon (often – especially in horticulture – simply "snapdragon"), is a species of flowering plant belonging to the genus ''Antirrhinum''. The plant was placed in the family Plantaginaceae following a r ...
'') along an elevation gradient, QST–FST analyses revealed different adaptation trends between two
subspecies In Taxonomy (biology), biological classification, subspecies (: subspecies) is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (Morphology (biology), morpholog ...
(''A. m. pseudomajus'' and ''A. m. striatum''). While both subspecies occur at all elevations, ''A. m. striatum'' had high QST values for traits associated with altitude adaptation: plant height, number of branches, and internode length. ''A. m. pseudomajus'' had lower QST than FST values for germination time.


See also

*
F-statistics In population genetics, ''F''-statistics (also known as fixation indices) describe the statistically expected level of heterozygosity in a population; more specifically the expected degree of (usually) a reduction in heterozygosity when compared ...
*
Quantitative genetics Quantitative genetics is the study of quantitative traits, which are phenotypes that vary continuously—such as height or mass—as opposed to phenotypes and gene-products that are Categorical variable, discretely identifiable—such as eye-col ...
*
Conservation genetics Conservation genetics is an interdisciplinary subfield of population genetics that aims to understand the dynamics of genes in a population for the purpose of natural resource management, conservation of genetic diversity, and the prevention of ...
* Divergent selection *
Genetic diversity Genetic diversity is the total number of genetic characteristics in the genetic makeup of a species. It ranges widely, from the number of species to differences within species, and can be correlated to the span of survival for a species. It is d ...


References

{{Reflist, 30em Genetics terminology Population genetics Statistical tests