arXiv subjects

Leo van Iersel

Publications and source records attributed to Leo van Iersel.

At least 73 records · Page 4Linked to original sources

arXiv2014

Satisfying ternary permutation constraints by multiple linear orders or phylogenetic trees

A ternary permutation constraint satisfaction problem (CSP) is specified by a subset Pi of the symmetric group S_3. An instance of such a problem consists of a set of variables V and a set of constraints C, where each constraint is an ordered triple of distinct elements from V. The goal is to construct a linear order alpha on V such that, for each constraint (a,b,c) in C, the ordering of a,b,c induced by alpha is in Pi. Excluding symmetries and trivial cases there are 11 such problems, and their complexity is well known. Here we consider the variant of the problem, denoted 2-Pi, where we are allowed to construct two linear orders alpha and beta and each constraint needs to be satisfied by at least one of the two. We give a full complexity classification of all 11 2-Pi problems, observing that in the switch from one to two linear orders the complexity landscape changes quite abruptly and that hardness proofs become rather intricate. We then focus on one of the 11 problems in particular, which is closely related to the '2-Caterpillar Compatibility' problem in the phylogenetics literature. We show that this particular CSP remains hard on three linear orders, and also in the biologically relevant case when we swap three linear orders for three phylogenetic trees, yielding the '3-Tree Compatibility' problem. Due to the biological relevance of this problem we also give extremal results concerning the minimum number of trees required, in the worst case, to satisfy a set of rooted triplet constraints on n leaf labels.

cs.CC↗

arXiv2014

A practical approximation algorithm for solving massive instances of hybridization number for binary and nonbinary trees

Reticulate events play an important role in determining evolutionary relationships. The problem of computing the minimum number of such events to explain discordance between two phylogenetic trees is a hard computational problem. Even for binary trees, exact solvers struggle to solve instances with reticulation number larger than 40-50. Here we present CycleKiller and NonbinaryCycleKiller, the first methods to produce solutions verifiably close to optimality for instances with hundreds or even thousands of reticulations. Using simulations, we demonstrate that these algorithms run quickly for large and difficult instances, producing solutions that are very close to optimality. As a spin-off from our simulations we also present TerminusEst, which is the fastest exact method currently available that can handle nonbinary trees: this is used to measure the accuracy of the NonbinaryCycleKiller algorithm. All three methods are based on extensions of previous theoretical work and are publicly available. We also apply our methods to real data.

q-bio.PE↗

arXiv2014

On Computing the Maximum Parsimony Score of a Phylogenetic Network

Phylogenetic networks are used to display the relationship of different species whose evolution is not treelike, which is the case, for instance, in the presence of hybridization events or horizontal gene transfers. Tree inference methods such as Maximum Parsimony need to be modified in order to be applicable to networks. In this paper, we discuss two different definitions of Maximum Parsimony on networks, "hardwired" and "softwired", and examine the complexity of computing them given a network topology and a character. By exploiting a link with the problem Multicut, we show that computing the hardwired parsimony score for 2-state characters is polynomial-time solvable, while for characters with more states this problem becomes NP-hard but is still approximable and fixed parameter tractable in the parsimony score. On the other hand we show that, for the softwired definition, obtaining even weak approximation guarantees is already difficult for binary characters and restricted network topologies, and fixed-parameter tractable algorithms in the parsimony score are unlikely. On the positive side we show that computing the softwired parsimony score is fixed-parameter tractable in the level of the network, a natural parameter describing how tangled reticulate activity is in the network. Finally, we show that both the hardwired and softwired parsimony score can be computed efficiently using Integer Linear Programming. The software has been made freely available.

q-bio.PE↗

arXiv2013

A short note on exponential-time algorithms for hybridization number

In this short note we prove that, given two (not necessarily binary) rooted phylogenetic trees T_1, T_2 on the same set of taxa X, where |X|=n, the hybridization number of T_1 and T_2 can be computed in time O^{*}(2^n) i.e. O(2^{n} poly(n)). The result also means that a Maximum Acyclic Agreement Forest (MAAF) can be computed within the same time bound.

q-bio.PE↗

arXiv2012

Approximation algorithms for nonbinary agreement forests

Given two rooted phylogenetic trees on the same set of taxa X, the Maximum Agreement Forest problem (MAF) asks to find a forest that is, in a certain sense, common to both trees and has a minimum number of components. The Maximum Acyclic Agreement Forest problem (MAAF) has the additional restriction that the components of the forest cannot have conflicting ancestral relations in the input trees. There has been considerable interest in the special cases of these problems in which the input trees are required to be binary. However, in practice, phylogenetic trees are rarely binary, due to uncertainty about the precise order of speciation events. Here, we show that the general, nonbinary version of MAF has a polynomial-time 4-approximation and a fixed-parameter tractable (exact) algorithm that runs in O(4^k poly(n)) time, where n = |X| and k is the number of components of the agreement forest minus one. Moreover, we show that a c-approximation algorithm for nonbinary MAF and a d-approximation algorithm for the classical problem Directed Feedback Vertex Set (DFVS) can be combined to yield a d(c+3)-approximation for nonbinary MAAF. The algorithms for MAF have been implemented and made publicly available.

math.CO↗

arXiv2012

Trinets encode tree-child and level-2 phylogenetic networks

Phylogenetic networks generalize evolutionary trees, and are commonly used to represent evolutionary histories of species that undergo reticulate evolutionary processes such as hybridization, recombination and lateral gene transfer. Recently, there has been great interest in trying to develop methods to construct rooted phylogenetic networks from triplets, that is rooted trees on three species. However, although triplets determine or encode rooted phylogenetic trees, they do not in general encode rooted phylogenetic networks, which is a potential issue for any such method. Motivated by this fact, Huber and Moulton recently introduced trinets as a natural extension of rooted triplets to networks. In particular, they showed that level-1 phylogenetic networks are encoded by their trinets, and also conjectured that all "recoverable" rooted phylogenetic networks are encoded by their trinets. Here we prove that recoverable binary level-2 networks and binary tree-child networks are also encoded by their trinets. To do this we prove two decomposition theorems based on trinets which hold for all recoverable binary rooted phylogenetic networks. Our results provide some additional evidence in support of the conjecture that trinets encode all recoverable rooted phylogenetic networks, and could also lead to new approaches to construct phylogenetic networks from trinets.

q-bio.PE↗

arXiv2012

A quadratic kernel for computing the hybridization number of multiple trees

It has recently been shown that the NP-hard problem of calculating the minimum number of hybridization events that is needed to explain a set of rooted binary phylogenetic trees by means of a hybridization network is fixed-parameter tractable if an instance of the problem consists of precisely two such trees. In this paper, we show that this problem remains fixed-parameter tractable for an arbitrarily large set of rooted binary phylogenetic trees. In particular, we present a quadratic kernel.

q-bio.PE↗

arXiv2011

Cycle killer... qu'est-ce que c'est? On the comparative approximability of hybridization number and directed feedback vertex set

We show that the problem of computing the hybridization number of two rooted binary phylogenetic trees on the same set of taxa X has a constant factor polynomial-time approximation if and only if the problem of computing a minimum-size feedback vertex set in a directed graph (DFVS) has a constant factor polynomial-time approximation. The latter problem, which asks for a minimum number of vertices to be removed from a directed graph to transform it into a directed acyclic graph, is one of the problems in Karp's seminal 1972 list of 21 NP-complete problems. However, despite considerable attention from the combinatorial optimization community it remains to this day unknown whether a constant factor polynomial-time approximation exists for DFVS. Our result thus places the (in)approximability of hybridization number in a much broader complexity context, and as a consequence we obtain that hybridization number inherits inapproximability results from the problem Vertex Cover. On the positive side, we use results from the DFVS literature to give an O(log r log log r) approximation for hybridization number, where r is the value of an optimal solution to the hybridization number problem.

math.CO↗

arXiv2011

All Ternary Permutation Constraint Satisfaction Problems Parameterized Above Average Have Kernels with Quadratic Numbers of Variables

A ternary Permutation-CSP is specified by a subset $Π$ of the symmetric group $\mathcal S_3$. An instance of such a problem consists of a set of variables $V$ and a multiset of constraints, which are ordered triples of distinct variables of $V.$ The objective is to find a linear ordering $α$ of $V$ that maximizes the number of triples whose ordering (under $α$) follows a permutation in $Π$. We prove that all ternary Permutation-CSPs parameterized above average have kernels with quadratic numbers of variables.

cs.DS↗

arXiv2011

On the elusiveness of clusters

Rooted phylogenetic networks are often used to represent conflicting phylogenetic signals. Given a set of clusters, a network is said to represent these clusters in the "softwired" sense if, for each cluster in the input set, at least one tree embedded in the network contains that cluster. Motivated by parsimony we might wish to construct such a network using as few reticulations as possible, or minimizing the "level" of the network, i.e. the maximum number of reticulations used in any "tangled" region of the network. Although these are NP-hard problems, here we prove that, for every fixed k >= 0, it is polynomial-time solvable to construct a phylogenetic network with level equal to k representing a cluster set, or to determine that no such network exists. However, this algorithm does not lend itself to a practical implementation. We also prove that the comparatively efficient Cass algorithm correctly solves this problem (and also minimizes the reticulation number) when input clusters are obtained from two not necessarily binary gene trees on the same set of taxa but does not always minimize level for general cluster sets. Finally, we describe a new algorithm which generates in polynomial-time all binary phylogenetic networks with exactly r reticulations representing a set of input clusters (for every fixed r >= 0).

q-bio.PE↗

arXiv2010

Locating a tree in a phylogenetic network

Phylogenetic trees and networks are leaf-labelled graphs that are used to describe evolutionary histories of species. The Tree Containment problem asks whether a given phylogenetic tree is embedded in a given phylogenetic network. Given a phylogenetic network and a cluster of species, the Cluster Containment problem asks whether the given cluster is a cluster of some phylogenetic tree embedded in the network. Both problems are known to be NP-complete in general. In this article, we consider the restriction of these problems to several well-studied classes of phylogenetic networks. We show that Tree Containment is polynomial-time solvable for normal networks, for binary tree-child networks, and for level-$k$ networks. On the other hand, we show that, even for tree-sibling, time-consistent, regular networks, both Tree Containment and Cluster Containment remain NP-complete.

q-bio.PE↗

arXiv2010

Computing Rooted and Unrooted Maximum Consistent Supertrees

A chief problem in phylogenetics and database theory is the computation of a maximum consistent tree from a set of rooted or unrooted trees. A standard input are triplets, rooted binary trees on three leaves, or quartets, unrooted binary trees on four leaves. We give exact algorithms constructing rooted and unrooted maximum consistent supertrees in time O(2^n n^5 m^2 log(m)) for a set of m triplets (quartets), each one distinctly leaf-labeled by some subset of n labels. The algorithms extend to weighted triplets (quartets). We further present fast exact algorithms for constructing rooted and unrooted maximum consistent trees in polynomial space. Finally, for a set T of m rooted or unrooted trees with maximum degree D and distinctly leaf-labeled by some subset of a set L of n labels, we compute, in O(2^{mD} n^m m^5 n^6 log(m)) time, a tree distinctly leaf-labeled by a maximum-size subset X of L that all trees in T, when restricted to X, are consistent with.

cs.DM↗

arXiv2010

When two trees go to war

Rooted phylogenetic networks are often constructed by combining trees, clusters, triplets or characters into a single network that in some well-defined sense simultaneously represents them all. We review these four models and investigate how they are related. In general, the model chosen influences the minimum number of reticulation events required. However, when one obtains the input data from two binary trees, we show that the minimum number of reticulations is independent of the model. The number of reticulations necessary to represent the trees, triplets, clusters (in the softwired sense) and characters (with unrestricted multiple crossover recombination) are all equal. Furthermore, we show that these results also hold when not the number of reticulations but the level of the constructed network is minimised. We use these unification results to settle several complexity questions that have been open in the field for some time. We also give explicit examples to show that already for data obtained from three binary trees the models begin to diverge.

q-bio.PE↗

arXiv2009

A short note on the tractability of constructing phylogenetic networks from clusters

In [2] it was proven that the Cass algorithm is a polynomial-time algorithm for constructing level<=2 networks from clusters. Here we demonstrate, for each k>=0, a polynomial-time algorithm for constructing level-k phylogenetic networks from clusters. Unlike Cass the algorithm scheme given here is only of theoretical interest. It does, however, strengthen the hope that efficient polynomial-time algorithms (and perhaps fixed parameter tractable algorithms) exist for this problem.

q-bio.PE↗

arXiv2009

Quantifying the Extent of Lateral Gene Transfer Required to Avert a `Genome of Eden'

The complex pattern of presence and absence of many genes across different species provides tantalising clues as to how genes evolved through the processes of gene genesis, gene loss and lateral gene transfer (LGT). The extent of LGT, particularly in prokaryotes, and its implications for creating a `network of life' rather than a `tree of life' is controversial. In this paper, we formally model the problem of quantifying LGT, and provide exact mathematical bounds, and new computational results. In particular, we investigate the computational complexity of quantifying the extent of LGT under the simple models of gene genesis, loss and transfer on which a recent heuristic analysis of biological data relied. Our approach takes advantage of a relationship between LGT optimization and graph-theoretical concepts such as tree width and network flow.

q-bio.PE↗

arXiv2009

A Practical Algorithm for Reconstructing Level-1 Phylogenetic Networks

Recently much attention has been devoted to the construction of phylogenetic networks which generalize phylogenetic trees in order to accommodate complex evolutionary processes. Here we present an efficient, practical algorithm for reconstructing level-1 phylogenetic networks - a type of network slightly more general than a phylogenetic tree - from triplets. Our algorithm has been made publicly available as the program LEV1ATHAN. It combines ideas from several known theoretical algorithms for phylogenetic tree and network reconstruction with two novel subroutines. Namely, an exponential-time exact and a greedy algorithm both of which are of independent theoretical interest. Most importantly, LEV1ATHAN runs in polynomial time and always constructs a level-1 network. If the data is consistent with a phylogenetic tree, then the algorithm constructs such a tree. Moreover, if the input triplet set is dense and, in addition, is fully consistent with some level-1 network, it will find such a network. The potential of LEV1ATHAN is explored by means of an extensive simulation study and a biological data set. One of our conclusions is that LEV1ATHAN is able to construct networks consistent with a high percentage of input triplets, even when these input triplets are affected by a low to moderate level of noise.

q-bio.PE↗

arXiv2009

Phylogenetic Networks Do not Need to Be Complex: Using Fewer Reticulations to Represent Conflicting Clusters

Phylogenetic trees are widely used to display estimates of how groups of species evolved. Each phylogenetic tree can be seen as a collection of clusters, subgroups of the species that evolved from a common ancestor. When phylogenetic trees are obtained for several data sets (e.g. for different genes), then their clusters are often contradicting. Consequently, the set of all clusters of such a data set cannot be combined into a single phylogenetic tree. Phylogenetic networks are a generalization of phylogenetic trees that can be used to display more complex evolutionary histories, including reticulate events such as hybridizations, recombinations and horizontal gene transfers. Here we present the new CASS algorithm that can combine any set of clusters into a phylogenetic network. We show that the networks constructed by CASS are usually simpler than networks constructed by other available methods. Moreover, we show that CASS is guaranteed to produce a network with at most two reticulations per biconnected component, whenever such a network exists. We have implemented CASS and integrated it in the freely available Dendroscope software.

q-bio.PE↗

arXiv2008

Uniqueness, intractability and exact algorithms: reflections on level-k phylogenetic networks

Phylogenetic networks provide a way to describe and visualize evolutionary histories that have undergone so-called reticulate evolutionary events such as recombination, hybridization or horizontal gene transfer. The level k of a network determines how non-treelike the evolution can be, with level-0 networks being trees. We study the problem of constructing level-k phylogenetic networks from triplets, i.e. phylogenetic trees for three leaves (taxa). We give, for each k, a level-k network that is uniquely defined by its triplets. We demonstrate the applicability of this result by using it to prove that (1) for all k of at least one it is NP-hard to construct a level-k network consistent with all input triplets, and (2) for all k it is NP-hard to construct a level-k network consistent with a maximum number of input triplets, even when the input is dense. As a response to this intractability we give an exact algorithm for constructing level-1 networks consistent with a maximum number of input triplets.

q-bio.PE↗