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204 C Constructing a Galled Phylogenetic Network
Theorem 5 Given two sets of rooted triplets T and F,
there exists an O(jLj
2
jT j(jTj + jFj))-time algorithm for
inferring a galled network N that guarantees jN(
T)j
jN(
F)j5/12(jT jjFj).
Applications
Phylogenetic networks are used by scientists to describe
evolutionary relationships that do not fit the traditional
models in which evolution is assumed to be treelike (see,
e. g., [12,16]). Evolutionary events such as horizontal gene
transfer or hybrid speciation (often referred to as recom-
bination events) that suggest convergence between objects
cannot be represented in a single tree [3,5,13,15,17]but
can be modeled in a phylogenetic network as internal
nodes having more than one parent. Galled networks are
an important type of phylogenetic network that have at-
tracted special attention in the literature [2,3,13,17]due
to their biological significance (see [3]) and their simple,
almost treelike, structure. When the number of recombi-
nation events is limited and most of the recombination
events have occurred recently, a galled network may suf-
fice to accurately describe the evolutionary process un-
der study [3].
An open challenge in the field of phylogenetics is to de-
velop efficient and reliable methods for constructing and
comparing phylogenetic networks. For example, to con-
struct a meaningful phylogenetic network for a large sub-
set of the human population (which may subsequently be
used to help locate regions in the genome associated with
some observable trait indicating a particular disease) in the
future, efficient algorithms are crucial because the input
can be expected to be very large.
The motivation behind the rooted triplet approach
taken in this paper is that a highly accurate tree for
each cardinality three subset of a leaf set can be obtained
through maximum-likelihood-based methods such as [1]
or Sibley–Ahlquist-style DNA–DNA hybridization exper-
iments (see [10]). Hence, the algorithms presented in [7]
and here can be used as the merging step in a divide-
and-conquer approach to constructing phylogenetic net-
works analogous to the quartet method paradigm for in-
ferring unrooted phylogenetic trees [9,11]andothersu-
pertree methods (see [6,14] and references therein). Dense
input sets in particular are considered since this case can
be solved in polynomial time.
Open Problems
For the rooted triplet problem, the current approxima-
tion ratio is not tight (0:4883 N(
T)/jT j5/12). It is
open if a tight approximation ratio can be found for this
problem. Similarly, a tight approximation ratio needs to
be found for the forbidden triplet problem.
Another direction is to work on a fixed-parameter
polynomial-time algorithm. Assume the number of hybrid
nodesisboundedbyh. Can an algorithm that is polyno-
mial in j
Tjwhile exponential in h be given?
Cross References
Directed Perfect Phylogeny (Binary Characters)
Distance-Based Phylogeny Reconstruction
(Fast-Converging)
Distance-Based Phylogeny Reconstruction (Optimal
Radius)
Perfect Phylogeny (Bounded Number of States)
Phylogenetic Tree Construction from a Distance
Matrix
Recommended Reading
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taxon ML
MC
trees with variable rates across sites. In: Proc. 1st
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vol. 2149, pp. 204–213. Springer, Berlin (2001)
2. Choy, C., Jansson, J., Sadakane, K., Sung, W.-K.: Computing the
maximum agreement of phylogenetic networks. In: Proc. Com-
puting: the 10th Australasian Theory Symposium (CATS 2004),
2004, pp. 33–45
3. Gusfield, D., Eddhu, S., Langley, C.: Efficient reconstruction
of phylogenetic networks with constrained recombination.
In: Proc. of Computational Systems Bioinformatics (CSB2003),
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phylogenetic relationships avoiding forbidden rooted triplets.
J Bioinform. Comput. Biol. 4(1), 59–74 (2006)
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combination using parsimony. Math. Biosci. 98(2), 185–200
(1990)
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homeomorphic subtrees, with applications to computational
evolutionary biology. Algorithmica 24(1), 1–13 (1999)
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ternational Computing and Combinatorics Conference (CO-
COON 2004), 2004
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ing rooted triplets into a galled phylogenetic network. SIAM J.
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scheme for inferring evolutionary trees from quartet topolo-
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(2001)
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