arXiv subjects

Leo van Iersel

Publications and source records attributed to Leo van Iersel.

At least 55 records · Page 3Linked to original sources

arXiv2019

Reconstructing Tree-Child Networks from Reticulate-Edge-Deleted Subnetworks

Network reconstruction lies at the heart of phylogenetic research. Two well studied classes of phylogenetic networks include tree-child networks and level-$k$ networks. In a tree-child network, every non-leaf node has a child that is a tree node or a leaf. In a level-$k$ network, the maximum number of reticulations contained in a biconnected component is $k$. Here, we show that level-$k$ tree-child networks are encoded by their reticulate-edge-deleted subnetworks, which are subnetworks obtained by deleting a single reticulation edge, if $k\geq 2$. Following this, we provide a polynomial-time algorithm for uniquely reconstructing such networks from their reticulate-edge-deleted subnetworks. Moreover, we show that this can even be done when considering subnetworks obtained by deleting one reticulation edge from each biconnected component with $k$ reticulations.

math.CO↗

arXiv2019

A Practical Fixed-Parameter Algorithm for Constructing Tree-Child Networks from Multiple Binary Trees

We present the first fixed-parameter algorithm for constructing a tree-child phylogenetic network that displays an arbitrary number of binary input trees and has the minimum number of reticulations among all such networks. The algorithm uses the recently introduced framework of cherry picking sequences and runs in $O((8k)^k \mathrm{poly}(n, t))$ time, where $n$ is the number of leaves of every tree, $t$ is the number of trees, and $k$ is the reticulation number of the constructed network. Moreover, we provide an efficient parallel implementation of the algorithm and show that it can deal with up to $100$ input trees on a standard desktop computer, thereby providing a major improvement over previous phylogenetic network construction methods.

cs.DM↗

arXiv2019

Deciding the existence of a cherry-picking sequence is hard on two trees

Here we show that deciding whether two rooted binary phylogenetic trees on the same set of taxa permit a cherry-picking sequence, a special type of elimination order on the taxa, is NP-complete. This improves on an earlier result which proved hardness for eight or more trees. Via a known equivalence between cherry-picking sequences and temporal phylogenetic networks, our result proves that it is NP-complete to determine the existence of a temporal phylogenetic network that contains topological embeddings of both trees. The hardness result also greatly strengthens previous inapproximability results for the minimum temporal-hybridization number problem. This is the optimization version of the problem where we wish to construct a temporal phylogenetic network that topologically embeds two given rooted binary phylogenetic trees and that has a minimum number of indegree-2 nodes, which represent events such as hybridization and horizontal gene transfer. We end on a positive note, pointing out that fixed parameter tractability results in this area are likely to ensure the continued relevance of the temporal phylogenetic network model.

q-bio.PE↗

arXiv2019

A third strike against perfect phylogeny

Perfect phylogenies are fundamental in the study of evolutionary trees because they capture the situation when each evolutionary trait emerges only once in history; if such events are believed to be rare, then by Occam's Razor such parsimonious trees are preferable as a hypothesis of evolution. A classical result states that 2-state characters permit a perfect phylogeny precisely if each subset of 2 characters permits one. More recently, it was shown that for 3-state characters the same property holds but for size-3 subsets. A long-standing open problem asked whether such a constant exists for each number of states. More precisely, it has been conjectured that for any fixed integer $r$, there exists a constant $f(r)$ such that a set of $r$-state characters $C$ has a perfect phylogeny if and only if every subset of at most $f(r)$ characters has a perfect phylogeny. In this paper, we show that this conjecture is false. In particular, we show that for any constant $t$, there exists a set $C$ of $8$-state characters such that $C$ has no perfect phylogeny, but there exists a perfect phylogeny for every subset of $t$ characters. This negative result complements the two negative results ("strikes") of Bodlaender et al. We reflect on the consequences of this third strike, pointing out that while it does close off some routes for efficient algorithm development, many others remain open.

cs.DM↗

arXiv2018

Not all phylogenetic networks are leaf-reconstructible

Unrooted phylogenetic networks are graphs used to represent evolutionary relationships. Accurately reconstructing such networks is of great relevance for evolutionary biology. It has recently been conjectured that all phylogenetic networks with at least five leaves can be uniquely reconstructed from their subnetworks obtained by deleting a single leaf and suppressing degree-2 vertices. Here, we show that this conjecture is false, by presenting a counter example for each possible number of leaves that is at least~4. Moreover, we show that the conjecture is still false when restricted to binary networks.

math.CO↗

arXiv2017

Exploring the tiers of rooted phylogenetic network space using tail moves

Popular methods for exploring the space of rooted phylogenetic trees use rearrangement moves such as rNNI (rooted Nearest Neighbour Interchange) and rSPR (rooted Subtree Prune and Regraft). Recently, these moves were generalized to rooted phylogenetic networks, which are a more suitable representation of reticulate evolutionary histories, and it was shown that any two rooted phylogenetic networks of the same complexity are connected by a sequence of either rSPR or rNNI moves. Here, we show that this is possible using only tail moves, which are a restricted version of rSPR moves on networks that are more closely related to rSPR moves on trees. The connectedness still holds even when we restrict to distance-1 tail moves (a localized version of tail-moves). Moreover, we give bounds on the number of (distance-1) tail moves necessary to turn one network into another, which in turn yield new bounds for rSPR, rNNI and SPR (i.e. the equivalent of rSPR on unrooted networks). The upper bounds are constructive, meaning that we can actually find a sequence with at most this length for any pair of networks. Finally, we show that finding a shortest sequence of tail or rSPR moves is NP-hard.

math.CO↗

arXiv2017

Finding the most parsimonious or likely tree in a network with respect to an alignment

Phylogenetic networks are often constructed by merging multiple conflicting phylogenetic signals into a directed acyclic graph. It is interesting to explore whether a network constructed in this way induces biologically-relevant phylogenetic signals that were not present in the input. Here we show that, given a multiple alignment A for a set of taxa X and a rooted phylogenetic network N whose leaves are labelled by X, it is NP-hard to locate the most parsimonious phylogenetic tree displayed by N (with respect to A) even when the level of N - the maximum number of reticulation nodes within a biconnected component - is 1 and A contains only 2 distinct states. (If, additionally, gaps are allowed the problem becomes APX-hard.) We also show that under the same conditions, and assuming a simple binary symmetric model of character evolution, finding the most likely tree displayed by the network is NP-hard. These negative results contrast with earlier work on parsimony in which it is shown that if A consists of a single column the problem is fixed parameter tractable in the level. We conclude with a discussion of why, despite the NP-hardness, both the parsimony and likelihood problem can likely be well-solved in practice.

q-bio.PE↗

arXiv2017

Graph realizations constrained by skeleton graphs

In 2008 Amanatidis, Green and Mihail introduced the Joint Degree Matrix (JDM) model to capture the fundamental difference in assortativity of networks in nature studied by the physical and life sciences and social networks studied in the social sciences. In 2014 Czabarka proposed a direct generalization of the JDM model, the Partition Adjacency Matrix (PAM) model. In the PAM model the vertices have specified degrees, and the vertex set itself is partitioned into classes. For each pair of vertex classes the number of edges between the classes in a graph realization is prescribed. In this paper we apply the new {\em skeleton graph} model to describe the same information as the PAM model. Our model is more convenient for handling problems with low number of partition classes or with special topological restrictions among the classes. We investigate two particular cases in detail: (i) when there are only two vertex classes and (ii) when the skeleton graph contains at most one cycle.

math.CO↗

arXiv2017

Leaf-reconstructibility of phylogenetic networks

An important problem in evolutionary biology is to reconstruct the evolutionary history of a set $X$ of species. This history is often represented as a phylogenetic network, that is, a connected graph with leaves labelled by elements in $X$ (for example, an evolutionary tree), which is usually also binary, i.e. all vertices have degree 1 or 3. A common approach used in phylogenetics to build a phylogenetic network on $X$ involves constructing it from networks on subsets of $X$. Here we consider the question of which (unrooted) phylogenetic networks are leaf-reconstructible, i.e. which networks can be uniquely reconstructed from the set of networks obtained from it by deleting a single leaf (its $X$-deck). This problem is closely related to the (in)famous reconstruction conjecture in graph theory but, as we shall show, presents distinct challenges. We show that some large classes of phylogenetic networks are reconstructible from their $X$-deck. This includes phylogenetic trees, binary networks containing at least one non-trivial cut-edge, and binary level-4 networks (the level of a network measures how far it is from being a tree). We also show that for fixed $k$, almost all binary level-$k$ phylogenetic networks are leaf-reconstructible. As an application of our results, we show that a level-3 network $N$ can be reconstructed from its quarnets, that is, 4-leaved networks that are induced by $N$ in a certain recursive fashion. Our results lead to several interesting open problems which we discuss, including the conjecture that all phylogenetic networks with at least five leaves are leaf-reconstructible.

math.CO↗

arXiv2017

Binets: fundamental building blocks for phylogenetic networks

Phylogenetic networks are a generalization of evolutionary trees that are used by biologists to represent the evolution of organisms which have undergone reticulate evolution. Essentially, a phylogenetic network is a directed acyclic graph having a unique root in which the leaves are labelled by a given set of species. Recently, some approaches have been developed to construct phylogenetic networks from collections of networks on 2- and 3-leaved networks, which are known as binets and trinets, respectively. Here we study in more depth properties of collections of binets, one of the simplest possible types of networks into which a phylogenetic network can be decomposed. More specifically, we show that if a collection of level-1 binets is compatible with some binary network, then it is also compatible with a binary level-1 network. Our proofs are based on useful structural results concerning lowest stable ancestors in networks. In addition, we show that, although the binets do not determine the topology of the network, they do determine the number of reticulations in the network, which is one of its most important parameters. We also consider algorithmic questions concerning binets. We show that deciding whether an arbitrary set of binets is compatible with some network is at least as hard as the well-known Graph Isomorphism problem. However, if we restrict to level-1 binets, it is possible to decide in polynomial time whether there exists a binary network that displays all the binets. We also show that to find a network that displays a maximum number of the binets is NP-hard, but that there exists a simple polynomial-time 1/3-approximation algorithm for this problem. It is hoped that these results will eventually assist in the development of new methods for constructing phylogenetic networks from collections of smaller networks.

q-bio.PE↗

arXiv2016

Nonbinary tree-based phylogenetic networks

Rooted phylogenetic networks are used to describe evolutionary histories that contain non-treelike evolutionary events such as hybridization and horizontal gene transfer. In some cases, such histories can be described by a phylogenetic base-tree with additional linking arcs, which can for example represent gene transfer events. Such phylogenetic networks are called tree-based. Here, we consider two possible generalizations of this concept to nonbinary networks, which we call tree-based and strictly-tree-based nonbinary phylogenetic networks. We give simple graph-theoretic characterizations of tree-based and strictly-tree-based nonbinary phylogenetic networks. Moreover, we show for each of these two classes that it can be decided in polynomial time whether a given network is contained in the class. Our approach also provides a new view on tree-based binary phylogenetic networks. Finally, we discuss two examples of nonbinary phylogenetic networks in biology and show how our results can be applied to them.

q-bio.PE↗

arXiv2016

On unrooted and root-uncertain variants of several well-known phylogenetic network problems

The hybridization number problem requires us to embed a set of binary rooted phylogenetic trees into a binary rooted phylogenetic network such that the number of nodes with indegree two is minimized. However, from a biological point of view accurately inferring the root location in a phylogenetic tree is notoriously difficult and poor root placement can artificially inflate the hybridization number. To this end we study a number of relaxed variants of this problem. We start by showing that the fundamental problem of determining whether an \emph{unrooted} phylogenetic network displays (i.e. embeds) an \emph{unrooted} phylogenetic tree, is NP-hard. On the positive side we show that this problem is FPT in reticulation number. In the rooted case the corresponding FPT result is trivial, but here we require more subtle argumentation. Next we show that the hybridization number problem for unrooted networks (when given two unrooted trees) is equivalent to the problem of computing the Tree Bisection and Reconnect (TBR) distance of the two unrooted trees. In the third part of the paper we consider the "root uncertain" variant of hybridization number. Here we are free to choose the root location in each of a set of unrooted input trees such that the hybridization number of the resulting rooted trees is minimized. On the negative side we show that this problem is APX-hard. On the positive side, we show that the problem is FPT in the hybridization number, via kernelization, for any number of input trees.

cs.DS↗

arXiv2016

Do branch lengths help to locate a tree in a phylogenetic network?

Phylogenetic networks are increasingly used in evolutionary biology to represent the history of species that have undergone reticulate events such as horizontal gene transfer, hybrid speciation and recombination. One of the most fundamental questions that arise in this context is whether the evolution of a gene with one copy in all species can be explained by a given network. In mathematical terms, this is often translated in the following way: is a given phylogenetic tree contained in a given phylogenetic network? Recently this tree containment problem has been widely investigated from a computational perspective, but most studies have only focused on the topology of the phylo- genies, ignoring a piece of information that, in the case of phylogenetic trees, is routinely inferred by evolutionary analyses: branch lengths. These measure the amount of change (e.g., nucleotide substitutions) that has occurred along each branch of the phylogeny. Here, we study a number of versions of the tree containment problem that explicitly account for branch lengths. We show that, although length information has the potential to locate more precisely a tree within a network, the problem is computationally hard in its most general form. On a positive note, for a number of special cases of biological relevance, we provide algorithms that solve this problem efficiently. This includes the case of networks of limited complexity, for which it is possible to recover, among the trees contained by the network with the same topology as the input tree, the closest one in terms of branch lengths.

q-bio.PE↗

arXiv2016

Hybridization Number on Three Rooted Binary Trees is EPT

Phylogenetic networks are leaf-labelled directed acyclic graphs that are used to describe non-treelike evolutionary histories and are thus a generalization of phylogenetic trees. The hybridization number of a phylogenetic network is the sum of all indegrees minus the number of nodes plus one. The Hybridization Number problem takes as input a collection of phylogenetic trees and asks to construct a phylogenetic network that contains an embedding of each of the input trees and has a smallest possible hybridization number. We present an algorithm for the Hybridization Number problem on three binary trees on $n$ leaves, which runs in time $O(c^k poly(n))$, with $k$ the hybridization number of an optimal network and $c$ a constant. For two trees, an algorithm with running time $O(3.18^k n)$ was proposed before whereas an algorithm with running time $O(c^k poly(n))$ had prior to this article remained elusive for more than two trees. The algorithm for two trees uses the close connection to acyclic agreement forests to achieve a linear exponent in the running time, while previous algorithms for more than two trees (explicitly or implicitly) relied on a brute force search through all possible underlying network topologies, leading to running times that are not $O(c^k poly(n))$ for any $c$. The connection to acyclic agreement forests is much weaker for more than two trees, so even given the right agreement forest, reconstructing the network poses major challenges. We prove novel structural results that allow us to reconstruct a network without having to guess the underlying topology. Our techniques generalize to more than three input trees with the exception of one key lemma that maps nodes in the network to tree nodes and, thus, minimizes the amount of guessing involved in constructing the network. The main open problem therefore is to establish a similar mapping for more than three trees.

cs.DS↗

arXiv2016

Kernelizations for the hybridization number problem on multiple nonbinary trees

Given a finite set $X$, a collection $\mathcal{T}$ of rooted phylogenetic trees on $X$ and an integer $k$, the Hybridization Number problem asks if there exists a phylogenetic network on $X$ that displays all trees from $\mathcal{T}$ and has reticulation number at most $k$. We show two kernelization algorithms for Hybridization Number, with kernel sizes $4k(5k)^t$ and $20k^2(Δ^+-1)$ respectively, with $t$ the number of input trees and $Δ^+$ their maximum outdegree. Experiments on simulated data demonstrate the practical relevance of these kernelization algorithms. In addition, we present an $n^{f(k)}t$-time algorithm, with $n=|X|$ and $f$ some computable function of $k$.

cs.DM↗

arXiv2015

Phylogenetic incongruence through the lens of Monadic Second Order logic

Within the field of phylogenetics there is growing interest in measures for summarising the dissimilarity, or 'incongruence', of two or more phylogenetic trees. Many of these measures are NP-hard to compute and this has stimulated a considerable volume of research into fixed parameter tractable algorithms. In this article we use Monadic Second Order logic (MSOL) to give alternative, compact proofs of fixed parameter tractability for several well-known incongruency measures. In doing so we wish to demonstrate the considerable potential of MSOL - machinery still largely unknown outside the algorithmic graph theory community - within phylogenetics. A crucial component of this work is the observation that many of these measures, when bounded, imply the existence of an 'agreement forest' of bounded size, which in turn implies that an auxiliary graph structure, the display graph, has bounded treewidth. It is this bound on treewidth that makes the machinery of MSOL available for proving fixed parameter tractability. We give a variety of different MSOL formulations. Some are based on explicitly encoding agreement forests, while some only use them implicitly to generate the treewidth bound. Our formulations introduce a number of "phylogenetics MSOL primitives" which will hopefully be of use to other researchers.

cs.DS↗

arXiv2014

Reconstructing phylogenetic level-1 networks from nondense binet and trinet sets

Binets and trinets are phylogenetic networks with two and three leaves, respectively. Here we consider the problem of deciding if there exists a binary level-1 phylogenetic network displaying a given set $\mathcal{T}$ of binary binets or trinets over a set $X$ of taxa, and constructing such a network whenever it exists. We show that this is NP-hard for trinets but polynomial-time solvable for binets. Moreover, we show that the problem is still polynomial-time solvable for inputs consisting of binets and trinets as long as the cycles in the trinets have size three. Finally, we present an $O(3^{|X|} poly(|X|))$ time algorithm for general sets of binets and trinets. The latter two algorithms generalise to instances containing level-1 networks with arbitrarily many leaves, and thus provide some of the first supernetwork algorithms for computing networks from a set of rooted phylogenetic networks.

cs.DS↗

arXiv2014

Exact reconciliation of undated trees

Reconciliation methods aim at recovering macro evolutionary events and at localizing them in the species history, by observing discrepancies between gene family trees and species trees. In this article we introduce an Integer Linear Programming (ILP) approach for the NP-hard problem of computing a most parsimonious time-consistent reconciliation of a gene tree with a species tree when dating information on speciations is not available. The ILP formulation, which builds upon the DTL model, returns a most parsimonious reconciliation ranging over all possible datings of the nodes of the species tree. By studying its performance on plausible simulated data we conclude that the ILP approach is significantly faster than a brute force search through the space of all possible species tree datings. Although the ILP formulation is currently limited to small trees, we believe that it is an important proof-of-concept which opens the door to the possibility of developing an exact, parsimony based approach to dating species trees. The software (ILPEACE) is freely available for download.

q-bio.PE↗