Everything about Phylogenetic Tree totally explained
A
phylogenetic tree, also called an
evolutionary tree, is a
tree showing the
evolutionary relationships among various biological
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, and the edge lengths in some trees correspond to
time estimates. Each node is called a taxonomic unit. Internal nodes are generally called hypothetical taxonomic units (HTUs) as they can't be directly observed.
History
Although the idea of a "
tree of life" arose from ancient notions of a ladder-like progression from lower to higher forms of
life (such as in the
Great Chain of Being),
Charles Darwin (1859)
first illustrated and popularized the notion of an
evolutionary "tree" in his seminal book
The Origin of Species. Over a century later,
evolutionary biologists still use
tree diagrams to depict
evolution because the
floral
analogy effectively conveys the concept that
speciation occurs through the
adaptive and
random splitting of lineages. Over time, species classification has become less static and more dynamic.
Adolf Engler (1844 - 1930) and
Karl A. E. Prantl (1849 - 1893) published a system of plant classification in their monograph
Die Natürlichen Pflanzenfamilien. In it, they arranged the families and orders of flowering plants on the basis of complexity of floral morphology. Characters like a
perianth with one
whorl, unisexual flowers and
pollination by wind were considered primitive as compared to perianth with two whorls, bisexual flowers and pollination by
insects.
The
plant kingdom is further divided into
divisions, sub-divisions,
classes,
orders and
families. According to this system,
monocotyledons are considered more primitive than
dicotyledons. It also considers evolution of
angiosperms from a single source and the sequence of orders and families show
parallel evolution.
Types
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. 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 common ancestry. While unrooted trees can always be generated from rooted ones by simply omitting the root, a root can't 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 1 depicts an unrooted phylogenetic tree 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
»
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.
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, for example, its branch lengths don't represent time.
A
phylogram is a phylogenetic tree that explicitly represents number of character changes through its branch lengths.
An
ultrametric tree or
chronogram is a phylogenetic tree that explicitly represents evolutionary time through its branch lengths.
Construction
Phylogenetic trees among a nontrivial number of input sequences are constructed using
computational phylogenetics methods. Distance-matrix methods such as
neighbor-joining or
UPGMA, which calculate
genetic distance from
multiple sequence alignments, are simplest to implement, but don't invoke an evolutionary model. Many sequence alignment methods such as
ClustalW also create trees by using the simpler algorithms (for example those based on distance) of tree construction.
Maximum parsimony is another simple method of estimating phylogenetic trees, but implies an implicit model of evolution (for example parsimony). More advanced methods use the
optimality criterion of
maximum likelihood, often within a
Bayesian Framework, and apply an explicit model of evolution to phylogenetic tree estimation.
- 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 isn't good to use, for example when assumptions are violated?)
Tree-building techniques have also gained the attention of mathematicians. Trees can also be built using
T-theory.
Limitations
Although phylogenetic trees produced on the basis of sequenced
genes or
genomic data in different species can provide evolutionary insight, they've important limitations. They don't necessarily accurately represent the species evolutionary history. The data on which they're based is
noisy; the analysis can be confounded by
horizontal gene transfer,
hybridisation between species that were not nearest neighbors on the tree before hybridisation takes place,
convergent evolution, and
conserved sequences. To avoid these limitations, one method of analysis, implemented in the program
PhyloCode, doesn't assume a tree structure.
Also, there are problems in basing the analysis on a single type of character, such as a single
gene or
protein or only on morphological analysis, because such trees constructed from another unrelated data source often differ from the first, and therefore great care is needed in inferring phylogenetic relationships among species. This is most true of genetic material that's subject to lateral gene transfer and
recombination, where different
haplotype blocks can have different histories. In general, the output tree of a phylogenetic analysis is an estimate of the
character's phylogeny (for example a gene tree) and not the phylogeny of the
taxa (for example species tree) from which these characters were sampled, though ideally, both should be very close.
When extinct species are included in a tree, they're
terminal nodes, as it's unlikely that they're 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 the fact that little useful "
ancient DNA" is preserved for longer than 100,000 years, and except in the most unusual circumstances no DNA sequences long enough for use in phylogenetic analyses have yet been recovered from material over 1 million years old.
In some organisms,
endosymbionts have an independent genetic history from the host.
Phylogenetic networks are used when bifurcating trees are not suitable, due to these complications which suggest a more
reticulate evolutionary history of the organisms sampled.
Further Information
Get more info on 'Phylogenetic Tree'.
|
External Link Exchanges
Do you know how hard it is to get a link from a large encyclopaedia? Well we're different and will prove it. To get a link from us just add the following HTML to your site on a relevant page:
<a href="http://phylogenetic_tree.totallyexplained.com">Phylogenetic tree Totally Explained</a>
Then simply click through this link from your web page. Our crawlers will verify your link, extract the title of your web page and instantly add a link back to it. If you like you can remove the words Totally Explained and embed the link in article text.
As long as your link remains in place, we'll keep our link to you right here. Please play fair - our crawlers are watching. Your site must be closely related to this one's topic. Any kind of spamming, dubious practises or removing the link will result in your link from us being dropped and, potentially, your whole site being banned. |