Heterotachy
Encyclopedia
Heterotachy refers to shifts in site-specific evolutionary rates over time. In the field of molecular evolution
Molecular evolution
Molecular evolution is in part a process of evolution at the scale of DNA, RNA, and proteins. Molecular evolution emerged as a scientific field in the 1960s as researchers from molecular biology, evolutionary biology and population genetics sought to understand recent discoveries on the structure...

, the principle of heterotachy states that the substitution rate of sites in a gene
Gene
A gene is a molecular unit of heredity of a living organism. It is a name given to some stretches of DNA and RNA that code for a type of protein or for an RNA chain that has a function in the organism. Living beings depend on genes, as they specify all proteins and functional RNA chains...

 can change through time. It has been proposed that the positions that show switches in substitution rate over time (that is, heterotachous sites) are good indicators of functional divergence. However, it appears that heterotachy is a much more general process, since most variable sites of homologous proteins with no evidence of functional shift are heterotachous.

The covarion
Covarion
The method of covarions, or concomitantly variable codons, is a technique in computational phylogenetics that allows the hypothesized rate of molecular evolution at individual codons in a set of nucleotide sequences to vary in an autocorrelated manner...

 hypothesis is a specific form of heterotachy. Some studies have proposed functional divergence models that are also heterotachous. Additionally, some mixture models that do not explicitly account for rate-shift, but site-partitions evolving at different relative substitution rates across lineages
Lineage (evolution)
An evolutionary lineage is a sequence of species, that form a line of descent, each new species the direct result of speciation from an immediate ancestral species. Lineages are subsets of the evolutionary tree of life. Lineages are often determined by the techniques of molecular systematics.-...

 are mathematically heterotachous.

Failure to take heterotachy into account in phylogenetic reconstructions may lead to incorrect phylogenetic tree
Phylogenetic tree
A phylogenetic tree or evolutionary tree is a branching diagram or "tree" showing the inferred evolutionary relationships among various biological species or other entities based upon similarities and differences in their physical and/or genetic characteristics...

s. Thus Zhong et al. (2011) say that heterotachy is one of the reasons for variability in reconstructions of the origin of gnetophytes.
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