Changes: Phylogenetic tree

Revision as of 21:08, 8 December 2006

A phylogenetic tree, also called an evolutionary tree or a tree of life, is a tree showing the evolutionary interrelationships among various species or other entities that are believed to have a common ancestor. In a phylogenetic tree, each node with descendants represents the most recent common ancestor of the descendants, with edge lengths sometimes corresponding to time estimates. Each node in a phylogenetic tree is called a taxonomic unit. Internal nodes are generally referred to as Hypothetical Taxonomic Units (HTUs) as they cannot be directly observed.

Though the idea of a "tree" originally arose from earlier ideas of life as a progression from lower to higher forms (The Great Chain of Being), modern evolutionary biologists still use trees to depict evolution because it still effectively captures the idea speciation occurs through the splitting of lineages.

Types of phylogenetic trees

MyosinUnrootedTree — Fig. 2: Unrooted tree of the myosin supergene family^[1]

A rooted phylogenetic tree is a directed tree with a unique node corresponding to the (usually imputed) most recent common ancestor of all the entities at the leaves of the tree. Figure 1 depicts a rooted phylogenetic tree, which has been colored according to the three-domain system^[2]. The most common method for rooting trees is the use of an uncontroversial outgroup - close enough to allow inference from sequence or trait data, but far enough to be a clear outgroup.

Unrooted trees illustrate the relatedness of the leaf nodes without making assumptions about ancestry. While unrooted trees can always be generated from rooted ones by simply omitting the root, a root cannot be inferred from an unrooted tree without some means of identifying ancestry; this is normally done by including an outgroup in the input data or introducing additional assumptions about the relative rates of evolution on each branch, such as an application of the molecular clock hypothesis. Figure 2 depicts an unrooted phylogenetic tree^[3] for myosin, a superfamily of proteins.

Both rooted and unrooted phylogenetic trees can be either bifurcating or multifurcating, and either labeled or unlabeled. A bifurcating tree has a maximum of two descendants arising from each interior node, while a multifurcating tree may have more than two. A labeled tree has specific values assigned to its leaves, while an unlabeled tree, sometimes called a tree shape, only defines a topology. The number of possible trees for a given number of leaf nodes depends on the specific type of tree, but there are always more multifurcating than bifurcating trees, more labeled than unlabeled trees, and more rooted than unrooted trees. The last distinction is the most biologically relevant; it arises because there are many places on an unrooted tree to put the root. For labeled bifurcating trees, there are

{\frac {(2n-3)!}{2^{n-2}(n-2)!}}

total rooted trees and

{\frac {(2n-5)!}{2^{n-3}(n-3)!}}

total unrooted trees, where n represents the number of leaf nodes. The number of unrooted trees for n input sequences or species is equal to the number of rooted trees for n-1 sequences.^[4]

A Dendrogram is a broad term for the diagrammatic representation of a phylogenetic tree.

A Cladogram is a tree formed using cladistic methods. This type of tree only represents a branching pattern, i.e., its branch lengths do not represent time.

A Phylogram is a phylogenetic tree that explicitly represents number of character changes through its branch lengths.

A Chronogram is a phylogenetic tree that explicitly represents evolutionary time through its branch lengths.

Phylogenetic tree construction

Main article: Computational phylogenetics

Phylogenetic trees among a nontrivial number of input sequences are constructed using computational phylogenetics methods. Distance-matrix methods such as neighbor-joining, which require multiple sequence alignments to calculate genetic distance, are simplest to implement; many sequence alignment methods such as ClustalW produce both sequence alignments and phylogenetic trees. Other methods include maximum parsimony and probabilistic inference techniques such as maximum likelihood; Bayesian inference has also been applied to phylogenetics but has been controversial.^[4] Identifying the optimal tree using many of these techniques is NP-complete or NP-hard^[4], so heuristic search and optimization methods are used in combination with tree-scoring functions to identify a reasonably good tree that fits the data.

Tree-building methods can be assessed on the basis of several criteria:^[5]

efficiency (how long does it take to compute the answer, how much memory does it need?)
power (does it make good use of the data, or is information being wasted?)
consistency (will it converge on the same answer repeatedly, if each time given different data for the same model problem?)
robustness (does it cope well with violations of the assumptions of the underlying model?)
falsifiability (does it alert us when it is not good to use, i.e. when assumptions are violated?)

Tree-building techniques have also gained the attention of mathematicians. Trees can also be built using T-theory. ^[6]

Limitations of phylogenetic trees

Although phylogenetic trees produced on the basis of sequenced genes or genomic data in different species can provide evolutionary insight, they do have important limitations. Phylogenetic trees do not necessarily (and likely do not) represent actual evolutionary history. The data on which they are based is noisy; horizontal gene transfer^[7], hybridisation between species that were not nearest neighbors on the tree before hybridisation takes place, convergent evolution, and conserved sequences can all confound the analysis. One method of analysis implemented in the program PhyloCode does not assume a tree structure to avoid these limitations.

Furthermore, basing the analysis on a single gene or protein taken from a group of species can be problematic because such trees constructed from another unrelated gene or protein sequence often differ from the first, and therefore great care is needed in inferring phylogenetic relationships amongst species. This is most true of genetic material that is subject to lateral gene transfer and recombination, where different haplotype blocks can have different histories.

When extinct species are included in a tree, they should always be terminal nodes, as it is unlikely that they are direct ancestors of any extant species. Scepticism must apply when extinct species are included in trees that are wholly or partly based on DNA sequence data, due to evidence that "ancient DNA" is not preserved intact for longer than 100,000 years.^{[How to reference and link to summary or text]}

References

↑ Hodge, T. & M. J. T. V. Cope. 2000. A Myosin Family Tree. Journal of Cell Science 113: 3353-3354. See also the Myosin external link below.
↑ Woese, C. R. 1998. The Universal Ancestor. Proceedings of the National Academy of Sciences 95: 6854-6859.
↑ Maher, B. A. 2002. Uprooting the Tree of Life. The Scientist 16: 18 (Sep. 16, 2002); subscription only
↑ ^4.0 ^4.1 ^4.2 Felsenstein J. (2004). Inferring Phylogenies Sinauer Associates: Sunderland, MA.
↑ Penny, D., Hendy, M. D. & M. A. Steel. 1992. Progress with methods for constructing evolutionary trees. Trends in Ecology and Evolution 7: 73-79.
↑ A. Dress, K. T. Huber, and V. Moulton. 2001. Metric Spaces in Pure and Applied Mathematics. Documenta Mathematica LSU 2001: 121-139
↑ Woese, C. R. 2002. On the evolution of cells. Proceedings of the National Academy of Sciences 99: 8742-8747.

External links

Pictures on the web

Phylogenetic Trees Based on 16s rDNA
A 3D View
Human Y-Chromosome 2002 Phylogenetic Tree
In 2003, the Science journal dedicated a special issue to the tree of life, including an online version of a tree of life.

General

PhyloCode
A Multiple Alignment of 139 Myosin Sequences and a Phylogenetic Tree
Tree of Life Web Project
http://www.aisee.com/graph_of_the_month/jura.htm The most detailed and comprehensive family tree of dinosaurs yet available
http://www.omne-vivum.com tree of life with lots of pictures

Topics in phylogenetics This box: view • talk • edit
Relevant fields: phylogenetics \| computational phylogenetics \| molecular phylogeny \| cladistics
Basic concepts: synapomorphy \| phylogenetic tree \| phylogenetic network \| long branch attraction
Phylogeny inference methods: maximum parsimony \| maximum likelihood \| neighbour joining \| UPGMA \| Bayesian inference
Current topics: PhyloCode \| DNA barcoding
List of evolutionary biology topics

ru:Филогенетическое дерево de:phylogenetischer Baum fr:Arbre phylogénétique no:Stamtre (evolusjon) oc:Classificacion filogenetica pt:Super reino vi:Cây phát sinh chủng loại zh:系統發生樹

This page uses Creative Commons Licensed content from Wikipedia (view authors).

[1] Hodge, T. & M. J. T. V. Cope. 2000. A Myosin Family Tree. Journal of Cell Science 113: 3353-3354. See also the Myosin external link below.

[2] Woese, C. R. 1998. The Universal Ancestor. Proceedings of the National Academy of Sciences 95: 6854-6859.

[3] Maher, B. A. 2002. Uprooting the Tree of Life. The Scientist 16: 18 (Sep. 16, 2002); subscription only

[Felsenstein-4] 4.0 ^4.1 ^4.2 Felsenstein J. (2004). Inferring Phylogenies Sinauer Associates: Sunderland, MA.

[5] Penny, D., Hendy, M. D. & M. A. Steel. 1992. Progress with methods for constructing evolutionary trees. Trends in Ecology and Evolution 7: 73-79.

[6] A. Dress, K. T. Huber, and V. Moulton. 2001. Metric Spaces in Pure and Applied Mathematics. Documenta Mathematica LSU 2001: 121-139

[7] Woese, C. R. 2002. On the evolution of cells. Proceedings of the National Academy of Sciences 99: 8742-8747.

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[2]

[3]

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@@ Line 1: / Line 1: @@
+A '''phylogenetic tree''', also called an '''evolutionary tree''' or a '''[[Tree_of_Life#Charles_Darwin_on_the_Tree_of_Life|tree of life]]''', is a [[tree (graph theory)|tree]] showing the [[evolution]]ary interrelationships among various [[species]] or other entities that are believed to have a [[common descent|common ancestor]].  In a phylogenetic tree, each node with descendants represents the most recent common ancestor of the descendants, with edge lengths sometimes corresponding to [[time]] estimates. Each node in a phylogenetic tree is called a taxonomic unit. Internal nodes are generally referred to as Hypothetical Taxonomic Units (HTUs) as they cannot be directly observed.
-#redirect[[Phylogenetic tree]]
+Though the idea of a "tree" originally arose from earlier ideas of life as a progression from lower to higher forms ([[The Great Chain of Being]]), modern evolutionary biologists still use trees to depict evolution because it still effectively captures the idea speciation occurs through the splitting of lineages.
+==Types of phylogenetic trees==
+[[Image:Phylogenetic_tree.svg|thumb|340px|Fig. 1: A speculatively rooted tree for rRNA genes]]
+[[Image:MyosinUnrootedTree.jpg|thumb|340px|Fig. 2: Unrooted tree of the myosin supergene family<ref>Hodge, T. & M. J. T. V. Cope. 2000. A Myosin Family Tree. ''Journal of Cell Science'' '''113''': 3353-3354. See also the Myosin external link below.</ref> ]]
+A '''rooted''' phylogenetic tree is a [[directed graph|directed]] tree with a unique node corresponding to the (usually [[imputation (statistics)|imputed]]) most recent common ancestor of all the entities at the [[leaf node|leaves]] of the tree.  Figure 1 depicts a rooted phylogenetic tree, which has been colored according to the ''[[three-domain system]]''<ref>[[Carl Woese|Woese, C. R]]. 1998. The Universal Ancestor. ''[[Proceedings of the National Academy of Sciences]]'' '''95''': 6854-6859.</ref>. The most common method for rooting trees is the use of an uncontroversial outgroup - close enough to allow inference from sequence or trait data, but far enough to be a clear outgroup.
+'''Unrooted''' trees illustrate the relatedness of the leaf nodes without making assumptions about ancestry. While unrooted trees can always be generated from rooted ones by simply omitting the root, a root cannot be inferred from an unrooted tree without some means of identifying ancestry; this is normally done by including an outgroup in the input data or introducing additional assumptions about the relative rates of evolution on each branch, such as an application of the [[molecular clock]] hypothesis. Figure 2 depicts an unrooted phylogenetic tree<ref>Maher, B. A. 2002. Uprooting the Tree of Life. ''The Scientist'' '''16''': 18 (Sep. 16, 2002); [http://www.the-scientist.com/yr2002/sep/research1_020916.html subscription only]</ref> for [[myosin]], a [[gene family|superfamily]] of [[protein]]s.
+Both rooted and unrooted phylogenetic trees can be either '''bifurcating''' or '''multifurcating''', and either ''labeled'' or ''unlabeled''. A bifurcating tree has a maximum of two descendants arising from each [[interior node]], while a multifurcating tree may have more than two. A labeled tree has specific values assigned to its leaves, while an unlabeled tree, sometimes called a '''tree shape''', only defines a topology. The number of possible trees for a given number of leaf nodes depends on the specific type of tree, but there are always more multifurcating than bifurcating trees, more labeled than unlabeled trees, and more rooted than unrooted trees. The last distinction is the most biologically relevant; it arises because there are many places on an unrooted tree to put the root. For labeled bifurcating trees, there are
+:<math>
+\frac{(2n-3)!}{2^{n-2}(n-2)!}
+</math>
+total rooted trees and
+:<math>
+\frac{(2n-5)!}{2^{n-3}(n-3)!}
+</math>
+total unrooted trees, where ''n'' represents the number of leaf nodes. The number of unrooted trees for ''n'' input sequences or species is equal to the number of rooted trees for ''n-1'' sequences.<ref name="Felsenstein">Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.</ref>
+A '''Dendrogram''' is a broad term for the diagrammatic representation of a phylogenetic tree.
+A '''Cladogram''' is a tree formed using [[Cladistics|cladistic]] methods.  This type of tree only represents a branching pattern, i.e., its branch lengths do not represent time.
+A '''Phylogram''' is a phylogenetic tree that explicitly represents number of character changes through its branch lengths.
+A '''Chronogram''' is a phylogenetic tree that explicitly represents evolutionary time through its branch lengths.
+==Phylogenetic tree construction==
+{{main|Computational phylogenetics}}
+Phylogenetic trees among a nontrivial number of input sequences are constructed using [[computational phylogenetics]] methods. Distance-matrix methods such as [[neighbor-joining]], which require [[multiple sequence alignment]]s to calculate [[genetic distance]], are simplest to implement; many sequence alignment methods such as [[ClustalW]] produce both sequence alignments and phylogenetic trees. Other methods include [[maximum parsimony]] and probabilistic inference techniques such as [[maximum likelihood]]; [[Bayesian inference]] has also been applied to phylogenetics but has been controversial.<ref name="Felsenstein" /> Identifying the optimal tree using many of these techniques is [[NP-complete]] or [[NP-hard]]<ref name="Felsenstein" />, so [[heuristic]] search and [[optimization]] methods are used in combination with tree-scoring functions to identify a reasonably good tree that fits the data.
+Tree-building methods can be assessed on the basis of several criteria:<ref>Penny, D., Hendy, M. D. & M. A. Steel. 1992. Progress with methods for constructing evolutionary trees. ''Trends in Ecology and Evolution'' '''7''': 73-79.</ref>
+*efficiency (how long does it take to compute the answer, how much memory does it need?)
+*power (does it make good use of the data, or is information being wasted?)
+*consistency (will it converge on the same answer repeatedly, if each time given different data for the same model problem?)
+*robustness (does it cope well with violations of the assumptions of the underlying model?)
+*falsifiability (does it alert us when it is not good to use, i.e. when assumptions are violated?)
+Tree-building techniques have also gained the attention of mathematicians. Trees can also be built using [[T-theory]]. <ref>A. Dress, K. T. Huber, and V. Moulton. 2001. Metric Spaces in Pure and Applied Mathematics. ''Documenta Mathematica'' ''LSU 2001'': 121-139</ref>
+==Limitations of phylogenetic trees==
+Although phylogenetic trees produced on the basis of sequenced [[gene]]s or [[genome|genomic]] data in different species can provide evolutionary insight, they do have important limitations. Phylogenetic trees do not necessarily (and likely do not) represent actual evolutionary history. The data on which they are based is [[signal noise|noisy]]; [[horizontal gene transfer]]<ref>Woese, C. R. 2002. On the evolution of cells. ''Proceedings of the National Academy of Sciences'' '''99''': 8742-8747.</ref>, [[hybrid]]isation between species that were not nearest neighbors on the tree before hybridisation takes place, [[convergent evolution]], and [[conservation (genetics)|conserved]] sequences can all confound the analysis. One method of analysis implemented in the program [[PhyloCode]] does not assume a tree structure to avoid these limitations.
+Furthermore, basing the analysis on a single [[gene]] or [[protein]] taken from a group of species can be problematic because such trees constructed from another unrelated gene or protein sequence often differ from the first, and therefore great care is needed in inferring phylogenetic relationships amongst species. This is most true of genetic material that is subject to lateral gene transfer and [[recombination]], where different [[haplotype]] blocks can have different histories.
+When extinct species are included in a tree, they should always be [[leaf node|terminal node]]s, as it is unlikely that they are direct ancestors of any extant species. Scepticism must apply when extinct species are included in trees that are wholly or partly based on DNA sequence data, due to evidence that "[[ancient DNA]]" is not preserved intact for longer than 100,000 years.{{fact}}
+==See also==
+* [[animal evolution]]
+* [[dendrogram]]
+* [[endosymbiosis]]
+* [[evolution]]
+* [[evolution of multicellularity]]
+* [[phylogeny]]
+* [[taxonomy]]
+* [[tree structure]]
+* [[:Wikipedia:WikiProject_Tree_of_Life|WikiProject Tree of Life]]
+* [[Wikispecies]]
+==References==
+<references />
+==External links==
+=== Pictures on the web ===
+*[http://tolweb.org/tree/eukaryotes/accessory/treeoverview.html Phylogenetic Trees Based on 16s rDNA]
+*[http://www.ii.uib.no/~tim/frontPages/treeOfLifeImagePage.shtml A 3D View]
+*[http://ycc.biosci.arizona.edu/nomenclature_system/fig1.html Human Y-Chromosome 2002 Phylogenetic Tree]
+* In 2003, the [[Science (journal)|''Science'']] journal dedicated a special issue to the tree of life, including an [http://www.sciencemag.org/feature/data/tol/ online version of a tree of life].
+===General===
+*[http://www.ohiou.edu/phylocode/index.html PhyloCode]
+*[http://www.mrc-lmb.cam.ac.uk/myosin/trees/trees.html A Multiple Alignment of 139 Myosin Sequences and a Phylogenetic Tree]
+*[http://tolweb.org/tree Tree of Life Web Project]
+* http://www.aisee.com/graph_of_the_month/jura.htm  The most detailed and comprehensive family tree of dinosaurs yet available
+* http://www.omne-vivum.com tree of life with lots of pictures
+{{phylo}}
+{{origin_of_life}}
+[[Category:Phylogenetics]]
+[[Category:Tree of life]]
+[[Category:Trees (structure)]]
+[[ru:Филогенетическое дерево]]
+[[de:phylogenetischer Baum]]
+[[fr:Arbre phylogénétique]]
+[[ja:系統樹]]
+[[no:Stamtre (evolusjon)]]
+[[oc:Classificacion filogenetica]]
+[[pl:Drzewo filogenetyczne]]
+[[pt:Super reino]]
+[[vi:Cây phát sinh chủng loại]]
+[[zh:系統發生樹]]
+<!-- interwiki links from the merged former [[evolutionary tree]],
+ I was sure how to handle them.
+[[da:Systema naturae]]
+[[de:Evolutionärer Stammbaum]]
+[[fr:Systématique]]
+[[nl:Classificatie en Evolutie]]
+[[pl:Drzewo życia (biologia)]]
+[[pt:Árvore da vida (biologia)]]
+[[sv:Systema naturae]]
+[[zh:演化樹]]
+-->
+{{enWP|Phylogenetic tree}}