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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 discret ...
used to visualize evolutionary relationships (either abstractly or explicitly) between
nucleotide sequence A nucleic acid sequence is a succession of bases within the nucleotides forming alleles within a DNA (using GACT) or RNA (GACU) molecule. This succession is denoted by a series of a set of five different letters that indicate the order of the nu ...
s,
gene In biology, the word gene has two meanings. The Mendelian gene is a basic unit of heredity. The molecular gene is a sequence of nucleotides in DNA that is transcribed to produce a functional RNA. There are two types of molecular genes: protei ...
s,
chromosome A chromosome is a package of DNA containing part or all of the genetic material of an organism. In most chromosomes, the very long thin DNA fibers are coated with nucleosome-forming packaging proteins; in eukaryotic cells, the most import ...
s,
genome 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 genes, other functional regions of the genome such as ...
s, or
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
. 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 organisms other than by the ("vertical") transmission of DNA from parent to offspring (reproduction). HGT is an important factor in the e ...
, recombination, or
gene duplication Gene duplication (or chromosomal duplication or gene amplification) is a major mechanism through which new genetic material is generated during molecular evolution. It can be defined as any duplication of a region of DNA that contains a gene ...
and loss are believed to be involved. They differ from
phylogenetic trees A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA. In o ...
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 or phylogeny is a graphical representation which shows the evolutionary history between a set of species or taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA. In ...
s are a subset of
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
networks. Phylogenetic networks can be inferred and visualised with software such as SplitsTree, the R-package, phangorn, and, more recently, Dendroscope. A standard format for representing phylogenetic networks is a variant of Newick format 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, enter ...
s, optimal realizations and reticulograms from a distance matrix), 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 ...
ary events, for example the geographical distribution of muskrat or fish populations of a given species among river networks, because there is no
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
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''; : 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 name and ...
. An unrooted phylogenetic network N on X is any undirected graph whose leaves are
bijective In mathematics, a bijection, bijective function, or one-to-one correspondence is a function between two sets such that each element of the second set (the codomain) is the image of exactly one element of the first set (the domain). Equival ...
ly labeled by the taxa in X. A number of different types of unrooted phylogenetic networks are in use like split networks and quasi-median networks. 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 ...
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 * Dendroscope
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