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Mike Steel

Publications and source records attributed to Mike Steel.

At least 55 records · Page 3Linked to original sources
arXiv2017

Minimum triplet covers of binary phylogenetic $X$-trees

Trees with labelled leaves and with all other vertices of degree three play an important role in systematic biology and other areas of classification. A classical combinatorial result ensures that such trees can be uniquely reconstructed from the distances between the leaves (when the edges are given any strictly positive lengths). Moreover, a linear number of these pairwise distance values suffices to determine both the tree and its edge lengths. A natural set of pairs of leaves is provided by any `triplet cover' of the tree (based on the fact that each non-leaf vertex is the median vertex of three leaves). In this paper we describe a number of new results concerning triplet covers of minimum size. In particular, we characterize such covers in terms of an associated graph being a 2-tree. Also, we show that minimum triplet covers are `shellable' and thereby provide a set of pairs for which the inter-leaf distance values will uniquely determine the underlying tree and its associated branch lengths.

math.CO↗
arXiv2017

Can we 'future-proof' consensus trees?

Consensus methods are widely used for combining phylogenetic trees into a single estimate of the evolutionary tree for a group of species. As more taxa are added, the new source trees may begin to tell a different evolutionary story when restricted to the original set of taxa. However, if the new trees, restricted to the original set of taxa, were to agree exactly with the earlier trees, then we might hope that their consensus would either agree with or resolve the original consensus tree. In this paper, we ask under what conditions consensus methods exist that are 'future proof' in this sense. While we show that some methods (e.g. Adams consensus) have this property for specific types of input, we also establish a rather surprising `no-go' theorem: there is no 'reasonable' consensus method that satisfies the future-proofing property in general. We then investigate a second notion of 'future proofing' for consensus methods, in which trees (rather than taxa) are added, and establish some positive and negative results. We end with some questions for future work.

q-bio.PE↗
arXiv2016

Phylogenetic mixtures and linear invariants for equal input models

The reconstruction of phylogenetic trees from molecular sequence data relies on modelling site substitutions by a Markov process, or a mixture of such processes. In general, allowing mixed processes can result in different tree topologies becoming indistinguishable from the data, even for infinitely long sequences. However, when the underlying Markov process supports linear phylogenetic invariants, then provided these are sufficiently informative, the identifiability of the tree topology can be restored. In this paper, we investigate a class of processes that support linear invariants once the stationary distribution is fixed, the `equal input model'. This model generalizes the `Felsenstein 1981' model (and thereby the Jukes--Cantor model) from four states to an arbitrary number of states (finite or infinite), and it can also be described by a `random cluster' process. We describe the structure and dimension of the vector spaces of phylogenetic mixtures and of linear invariants for any fixed phylogenetic tree (and for all trees -- the so called `model invariants'), on any number $n$ of leaves. We also provide a precise description of the space of mixtures and linear invariants for the special case of $n=4$ leaves. By combining techniques from discrete random processes and (multi-) linear algebra, our results build on a classic result that was first established by James Lake in 1987.

q-bio.PE↗
arXiv2016

Autocatalytic sets in polymer networks with variable catalysis distributions

All living systems -- from the origin of life to modern cells -- rely on a set of biochemical reactions that are simultaneously self-sustaining and autocatalytic. This notion of an autocatalytic set has been formalized graph-theoretically (as `RAF'), leading to mathematical results and polynomial-time algorithms that have been applied to simulated and real chemical reaction systems. In this paper, we investigate the emergence of autocatalytic sets in polymer models when the catalysis rate of each molecule type is drawn from some probability distribution. We show that although the average catalysis rate $f$ for RAFs to arise depends on this distribution, a universal linear upper and lower bound for $f$ (with increasing system size) still applies. However, the probability of the appearance (and size) of autocatalytic sets can vary widely, depending on the particular catalysis distribution. We use simulations to explore how tight the mathematical bounds are, and the reasons for the observed variations. We also investigate the impact of inhibition (where molecules can also inhibit reactions) on the emergence of autocatalytic sets, deriving new mathematical and algorithmic results.

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arXiv2015

Bounds on the Expected Size of the Maximum Agreement Subtree

We prove polynomial upper and lower bounds on the expected size of the maximum agreement subtree of two random binary phylogenetic trees under both the uniform distribution and Yule-Harding distribution. This positively answers a question posed in earlier work. Determining tight upper and lower bounds remains an open problem.

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arXiv2015

Capturing a phylogenetic tree when the number of character states varies with the number of leaves

We show that for any two values $α, β>0 $ for which $α+β>1$ then there is a value $N$ so that for all $n \geq N$ the following holds. For any binary phylogenetic tree $T$ on $n$ leaves there is a set of $\lfloor n^α\rfloor$ characters that capture $T$, and for which each character takes at most $\lfloor n^β\rfloor$ distinct states. Here `capture' means that $T$ is the unique perfect phylogeny for these characters. Our short proof of this combinatorial result is based on the probabilistic method.

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arXiv2015

Neighbourhoods of phylogenetic trees: exact and asymptotic counts

A central theme in phylogenetics is the reconstruction and analysis of evolutionary trees from a given set of data. To determine the optimal search methods for reconstructing trees, it is crucial to understand the size and structure of the neighbourhoods of trees under tree rearrangement operations. The diameter and size of the immediate neighbourhood of a tree has been well-studied, however little is known about the number of trees at distance two, three or (more generally) $k$ from a given tree. In this paper we provide a number of exact and asymptotic results concerning these quantities, and identify some key aspects of tree shape that play a role in determining these quantities. We obtain several new results for two of the main tree rearrangement operations - Nearest Neighbour Interchange and Subtree Prune and Regraft -- as well as for the Robinson-Foulds metric on trees.

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arXiv2015

Folding and unfolding phylogenetic trees and networks

Phylogenetic networks are rooted, labelled directed acyclic graphs which are commonly used to represent reticulate evolution. There is a close relationship between phylogenetic networks and multi-labelled trees (MUL-trees). Indeed, any phylogenetic network $N$ can be 'unfolded' to obtain a MUL-tree $U(N)$ and, conversely, a MUL-tree $T$ can in certain circumstances be 'folded' to obtain a phylogenetic network $F(T)$ that exhibits $T$. In this paper, we study properties of the operations $U$ and $F$ in more detail. In particular, we introduce the class of stable networks, phylogenetic networks $N$ for which $F(U(N))$ is isomorphic to $N$, characterise such networks, and show that that they are related to the well-known class of tree-sibling networks. We also explore how the concept of displaying a tree in a network $N$ can be related to displaying the tree in the MUL-tree $U(N)$. To do this, we develop a phylogenetic analogue of graph fibrations. This allows us to view $U(N)$ as the analogue of the universal cover of a digraph, and to establish a close connection between displaying trees in $U(N)$ and reconciling phylogenetic trees with networks.

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arXiv2015

Which phylogenetic networks are merely trees with additional arcs?

A binary phylogenetic network may or may not be obtainable from a tree by the addition of directed edges (arcs) between tree arcs. Here, we establish a precise and easily tested criterion (based on `2-SAT') that efficiently determines whether or not any given network can be realized in this way. Moreover, the proof provides a polynomial-time algorithm for finding one or more trees (when they exist) on which the network can be based. A number of interesting consequences are presented as corollaries; these lead to some further relevant questions and observations, which we outline in the conclusion.

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arXiv2015

Twisted trees and inconsistency of tree estimation when gaps are treated as missing data -- the impact of model mis-specification in distance corrections

Statistically consistent estimation of phylogenetic trees or gene trees is possible if pairwise sequence dissimilarities can be converted to a set of distances that are proportional to the true evolutionary distances. Susko et al. (2004) reported some strikingly broad results about the forms of inconsistency in tree estimation that can arise if corrected distances are not proportional to the true distances. They showed that if the corrected distance is a concave function of the true distance, then inconsistency due to long branch attraction will occur. If these functions are convex, then two "long branch repulsion" trees will be preferred over the true tree -- though these two incorrect trees are expected to be tied as the preferred true. Here we extend their results, and demonstrate the existence of a tree shape (which we refer to as a "twisted Farris-zone" tree) for which a single incorrect tree topology will be guaranteed to be preferred if the corrected distance function is convex. We also report that the standard practice of treating gaps in sequence alignments as missing data is sufficient to produce non-linear corrected distance functions if the substitution process is not independent of the insertion/deletion process. Taken together, these results imply inconsistent tree inference under mild conditions. For example, if some positions in a sequence are constrained to be free of substitutions and insertion/deletion events while the remaining sites evolve with independent substitutions and insertion/deletion events, then the distances obtained by treating gaps as missing data can support an incorrect tree topology even given an unlimited amount of data.

q-bio.PE↗
arXiv2015

The existence and abundance of ghost ancestors in biparental populations

In a randomly-mating biparental population of size $N$ there are, with high probability, individuals who are genealogical ancestors of every extant individual within approximately $\log_2(N)$ generations into the past. We use this result of J. Chang to prove a curious corollary under standard models of recombination: there exist, with high probability, individuals within a constant multiple of $ \log_2(N)$ generations into the past who are simultaneously (i) genealogical ancestors of {\em each} of the individuals at the present, and (ii) genetic ancestors to {\em none} of the individuals at the present. Such ancestral individuals - ancestors of everyone today that left no genetic trace -- represent `ghost' ancestors in a strong sense. In this short note, we use simple analytical argument and simulations to estimate how many such individuals exist in finite Wright-Fisher populations.

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arXiv2015

Self-sustaining autocatalytic networks within open-ended reaction systems

Given any finite and closed chemical reaction system, it is possible to efficiently determine whether or not it contains a `self-sustaining and collectively autocatalytic' subset of reactions, and to find such subsets when they exist. However, for systems that are potentially open-ended (for example, when no prescribed upper bound is placed on the complexity or size/length of molecules types), the theory developed for the finite case breaks down. We investigate a number of subtleties that arise in such systems that are absent in the finite setting, and present several new results.

q-bio.MN↗
arXiv2015

Circumstances in which parsimony but not compatibility will be provably misleading

Phylogenetic methods typically rely on an appropriate model of how data evolved in order to infer an accurate phylogenetic tree. For molecular data, standard statistical methods have provided an effective strategy for extracting phylogenetic information from aligned sequence data when each site (character) is subject to a common process. However, for other types of data (e.g. morphological data), characters can be too ambiguous, homoplastic or saturated to develop models that are effective at capturing the underlying process of change. To address this, we examine the properties of a classic but neglected method for inferring splits in an underlying tree, namely, maximum compatibility. By adopting a simple and extreme model in which each character either fits perfectly on some tree, or is entirely random (but it is not known which class any character belongs to) we are able to derive exact and explicit formulae regarding the performance of maximum compatibility. We show that this method is able to identify a set of non-trivial homoplasy-free characters, when the number $n$ of taxa is large, even when the number of random characters is large. By contrast, we show that a method that makes more uniform use of all the data --- maximum parsimony --- can provably estimate trees in which {\em none} of the original homoplasy-free characters support splits.

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arXiv2015

Similarities as Evidence for Common Ancestry -- A Likelihood Epistemology

Darwin claims in the {\em Origin} that similarity is evidence for common ancestry, but that adaptive similarities are "almost valueless" as evidence. This claim seems reasonable for some adaptive similarities but not for others. Here we clarify and evaluate these and related matters by using the law of likelihood as an analytic tool and by considering mathematical models of three evolutionary processes -- directional selection, stabilizing selection, and drift. Our results apply both to Darwin's theory of evolution and to modern evolutionary biology.

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arXiv2014

A 'stochastic safety radius' for distance-based tree reconstruction

A variety of algorithms have been proposed for reconstructing trees that show the evolutionary relationships between species by comparing differences in genetic data across present-day taxa. If the leaf-to-leaf distances in a tree can be accurately estimated, then it is possible to reconstruct this tree from these estimated distances, using polynomial-time methods such as the popular `Neighbor-Joining' algorithm. There is a precise combinatorial condition under which distance-based methods are guaranteed to return a correct tree (in full or in part) based on the requirement that the input distances all lie within some `safety radius' of the true distances. Here, we explore a stochastic analogue of this condition, and mathematically establish upper and lower bounds on this `stochastic safety radius' for distance-based tree reconstruction methods. Using simulations, we show how this notion provides a new way to compare the performance of distance-based tree reconstruction methods. This may help explain why Neighbor-Joining performs so well, as its stochastic safety radius appears close to optimal (while its more classical safety radius is the same as many other less accurate methods).

q-bio.PE↗
arXiv2014

Impacts of terraces on phylogenetic inference

Terraces are potentially large sets of trees with precisely the same likelihood or parsimony score, which can be induced by missing sequences in partitioned multi-locus phylogenetic data matrices. The set of trees on a terrace can be characterized by enumeration algorithms or consensus methods that exploit the pattern of partial taxon coverage in the data, independent of the sequence data themselves. Terraces add ambiguity and complexity to phylogenetic inference particularly in settings where inference is already challenging: data sets with many taxa and relatively few loci. In this paper we present five new findings about terraces and their impacts on phylogenetic inference. First we clarify assumptions about model parameters that are necessary for the existence of terraces. Second, we explore the dependence of terrace size on partitioning scheme and indicate how to find the partitioning scheme associated with the largest terrace containing a given tree. Third, we highlight the impact of terraces on bootstrap estimates of confidence limits in clades, and characterize the surprising result that the bootstrap proportion for a clade can be entirely determined by the frequency of bipartitions on a terrace, with some bipartitions receiving high support even when incorrect. Fourth, we dissect some effects of prior distributions of edge lengths on the computed posterior probabilities of clades on terraces, to understand an example in which long edges "attract" each other in Bayesian inference. Fifth, we show that even if data are not partitioned, patterns of missing data studied in the terrace problem can lead to instances of apparent statistical inconsistency when even a small element of heterotachy is introduced to the model generating the sequence data. Finally, we discuss strategies for remediation of some of these problems.

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arXiv2014

Tree-like Reticulation Networks - When Do Tree-like Distances Also Support Reticulate Evolution?

Hybrid evolution and horizontal gene transfer (HGT) are processes where evolutionary relationships may more accurately be described by a reticulated network than by a tree. In such a network, there will often be several paths between any two extant species, reflecting the possible pathways that genetic material may have been passed down from a common ancestor to these species. These paths will typically have different lengths but an `average distance' can still be calculated between any two taxa. In this article, we ask whether this average distance is able to distinguish reticulate evolution from pure tree-like evolution. We consider two types of reticulation networks: hybridization networks and HGT networks. For the former, we establish a general result which shows that average distances between extant taxa can appear tree-like, but only under a single hybridization event near the root; in all other cases, the two forms of evolution can be distinguished by average distances. For HGT networks, we demonstrate some analogous but more intricate results.

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