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A phylogenetic network is any
graph Graph may refer to: Mathematics *Graph (discrete mathematics), a structure made of vertices and edges **Graph theory, the study of such graphs and their properties *Graph (topology), a topological space resembling a graph in the sense of discre ...
used to visualize evolutionary relationships (either abstractly or explicitly) between
nucleotide sequence A nucleic acid sequence is a succession of bases signified by a series of a set of five different letters that indicate the order of nucleotides forming alleles within a DNA (using GACT) or RNA (GACU) molecule. By convention, sequences are usu ...
s,
gene In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...
s,
chromosome A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...
s,
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ge ...
s, or
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
. They are employed when reticulation events such as hybridization,
horizontal gene transfer Horizontal gene transfer (HGT) or lateral gene transfer (LGT) is the movement of genetic material between unicellular and/or multicellular organisms other than by the ("vertical") transmission of DNA from parent to offspring (reproduction). H ...
, recombination, or gene duplication and loss are believed to be involved. They differ from
phylogenetic trees A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
by the explicit modeling of richly linked networks, by means of the addition of hybrid nodes (nodes with two parents) instead of only tree nodes (a hierarchy of nodes, each with only one parent).
Phylogenetic tree A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
s are a subset of
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
networks. Phylogenetic networks can be inferred and visualised with software such as
SplitsTree SplitsTree is a popular freeware program for inferring phylogenetic trees, phylogenetic networks, or, more generally, splits graphs, from various types of data such as a sequence alignment, a distance matrix or a set of trees. SplitsTree impleme ...
, the R-package, phangorn, and, more recently,
Dendroscope Dendroscope is an interactive computer software program written in Java for viewing Phylogenetic trees. This program is designed to view trees of all sizes and is very useful for creating figures. Dendroscope can be used for a variety of analyse ...
. A standard format for representing phylogenetic networks is a variant of
Newick format In mathematics, Newick tree format (or Newick notation or New Hampshire tree format) is a way of representing graph-theoretical trees with edge lengths using parentheses and commas. It was adopted by James Archie, William H. E. Day, Joseph Fels ...
which is extended to support networks as well as trees. Many kinds and subclasses of phylogenetic networks have been defined based on the biological phenomenon they represent or which data they are built from (hybridization networks, usually built from rooted trees, ancestral recombination graphs (ARGs) from binary sequences, median networks from a set of
split Split(s) or The Split may refer to: Places * Split, Croatia, the largest coastal city in Croatia * Split Island, Canada, an island in the Hudson Bay * Split Island, Falkland Islands * Split Island, Fiji, better known as Hạfliua Arts, entertai ...
s, optimal realizations and reticulograms from a
distance matrix In mathematics, computer science and especially graph theory, a distance matrix is a square matrix (two-dimensional array) containing the distances, taken pairwise, between the elements of a set. Depending upon the application involved, the ''dist ...
), or restrictions to get computationally tractable problems (galled trees, and their generalizations level-k phylogenetic networks, tree-child or tree-sibling phylogenetic networks).


Microevolution

Phylogenetic trees also have trouble depicting
microevolution Microevolution is the change in allele frequencies that occurs over time within a population. This change is due to four different processes: mutation, selection (natural and artificial), gene flow and genetic drift. This change happens over a ...
ary events, for example the geographical distribution of muskrat or fish populations of a given species among river networks, because there is no
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
boundary to prevent gene flow between populations. Therefore, a more general phylogenetic network better depicts these situations.


Rooted vs unrooted

;Unrooted phylogenetic network :Let X be a set of
taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
. An unrooted phylogenetic network N on X is any undirected graph whose leaves are
bijective In mathematics, a bijection, also known as a bijective function, one-to-one correspondence, or invertible function, is a function between the elements of two sets, where each element of one set is paired with exactly one element of the other ...
ly labeled by the taxa in X. A number of different types of unrooted phylogenetic networks are in use like
split networks For a given set of taxa like X, and a set of splits S on X, usually together with a non-negative weighting, which may represent character changes distance, or may also have a more abstract interpretation, if the set of splits S is compatible, the ...
and
quasi-median networks The concept of a quasi-median network is a generalization of the concept of a median network that was introduced to represent multistate characters. Note that, unlike median networks, quasi-median networks are not split networks. A quasi-median n ...
. In most cases, such networks only depict relations between taxa, without giving information about the evolutionary history. Although some methods produce unrooted networks that can be interpreted as undirected versions of rooted networks, which do represent a phylogeny. ;Rooted phylogenetic network :Let X be a set of taxa. A rooted phylogenetic network N on X is a rooted
directed acyclic graph In mathematics, particularly graph theory, and computer science, a directed acyclic graph (DAG) is a directed graph with no directed cycles. That is, it consists of vertices and edges (also called ''arcs''), with each edge directed from one ve ...
where the set of leaves is bijectively labeled by the taxa in X. Rooted phylogenetic networks, like rooted phylogenetic trees, give explicit representations of evolutionary history. This means that they visualize the order in which the species diverged (speciated), converged (hybridized), and transferred genetic material (horizontal gene transfer).


Classes of networks

For computational purposes, studies often restrict their attention to classes of networks: subsets of all networks with certain properties. Although computational simplicity is the main goal, most of these classes have a biological justification as well. Some prominent classes currently used in the mathematical phylogenetics literature are tree-child networks, tree-based networks, and level-k networks


Software to compute phylogenetic networks


PhyloNet
A Java-based software package that builds phylogenetic networks taking ILS, HGT etc. into consideration.
PhyloNetworks

Julia
package for the manipulation, visualization, inference of phylogenetic networks, and their use for trait evolution.

Free Phylogenetic Network Software. Network generates evolutionary trees and networks from genetic, linguistic, and other data.

some of which compute phylogenetic networks
List of programs for phylogenetic network reconstruction, evaluation, visualization, etc.
*
SplitsTree SplitsTree is a popular freeware program for inferring phylogenetic trees, phylogenetic networks, or, more generally, splits graphs, from various types of data such as a sequence alignment, a distance matrix or a set of trees. SplitsTree impleme ...
*
Dendroscope Dendroscope is an interactive computer software program written in Java for viewing Phylogenetic trees. This program is designed to view trees of all sizes and is very useful for creating figures. Dendroscope can be used for a variety of analyse ...

Network inferring on the T-REX serverTCS
Phylogenetic networks from DNA sequences or nucleotide distances using statistical parsimony.
NetTest
Characterization of phylogenetic networks.


SimPlot++
Sequence similarity network analysis.


References


Further reading

* {{Phylogenetics Phylogenetics