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Bioinformatics Algorithms: Techniques and Applications

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RECONCILIATION OF GENE TREES AND SPECIES TREES 171<br />

In [6], it is shown that Definition 7.6 computes a tree that satisfies the properties<br />

for the reconciled tree <strong>and</strong> such a tree has also minimum size. Furthermore, given a<br />

gene tree TG <strong>and</strong> a species tree TS, there exists a unique minimum reconciled tree for<br />

TG <strong>and</strong> TS [6], which is also the tree inducing the minimum number of duplications<br />

<strong>and</strong> losses [21]. Next, we introduce the main problem for the lateral gene transfer<br />

model.<br />

PROBLEM 7.12 α-Active Lateral Gene Transfer Problem<br />

Input: gene tree TG <strong>and</strong> species tree TS.<br />

Output: find a minimum cost α-active lateral transfer scenario for TS <strong>and</strong> TG.<br />

The restriction of α-active Lateral Gene Transfer Problem where α = 1 is<br />

APX-hard [11,23], while it has a O(2 4T |S| 2 ) fixed-parameter algorithm [23],<br />

where the parameter T is the cost of the scenario. For arbitrary α there is an<br />

O(4 α (4T (α + T )) T |L| 2 ) time algorithm [23].<br />

The extension of the problem that considers both duplications <strong>and</strong> lateral gene<br />

transfers is known to be NP-hard [24] <strong>and</strong> admits a fixed-parameter tractable<br />

algorithm [24] that computes the minimum number of duplications <strong>and</strong> lateral<br />

transfers.<br />

7.5.3 Open Problems<br />

A deep underst<strong>and</strong>ing of the approximation complexity of the agreement problems<br />

is still needed. More precisely, the only known result is the 2-factor approximation<br />

algorithm for the variant of Optimal Species Tree Reconstruction with Duplication<br />

Cost (see Problem 7.8) in which the duplication cost is slightly modified to<br />

obtain a metric d [31]. In this variant, all gene trees are uniquely labeled. Moreover,<br />

given a gene tree TG <strong>and</strong> a species tree TS, the symmetric duplication cost<br />

between TG <strong>and</strong> TS is defined as d(TG,TS) = 1 2 (dup(TS,TG) + dup(TG,TS)). The<br />

new version of the problem remains NP-hard while admitting a 2-approximation<br />

algorithm [31].<br />

An interesting open problem on species trees <strong>and</strong> gene trees is the computational<br />

complexity of reconstructing a species tree from a set of gene trees over instances<br />

consisting of a constant number of gene trees or even of two gene trees only.<br />

An extension of the reconciliation approach (see Definition 7.5 <strong>and</strong><br />

Problem 7.11) is proposed in [20] by introducing a notion of extended reconciled<br />

tree allowing the identification of lateral gene transfers, in addition to duplication<br />

<strong>and</strong> losses. A notion of scenario is also introduced to identify lateral<br />

transfers. A dynamic programming algorithm to compute a scenario inducing<br />

a minimum reconciliation cost is given [20]. Also notice that no approximation<br />

algorithms are known for the event inference problems presented in this<br />

section.

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