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Register a new userMeasuring tree balance using symmetry nodes -- a new balance index and its extremal properties
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Effects like selection in evolution as well as fertility inheritance in the development of populations can lead to a higher degree of asymmetry in evolutionary trees than expected under a null hypothesis. To identify and quantify such influences, various balance indices were proposed in the phylogenetic literature and have been in use for decades. However, so far no balance index was based on the number of emph{symmetry nodes}, even though symmetry nodes play an important role in other areas of mathematical phylogenetics and despite the fact that symmetry nodes are a quite natural way to measure balance or symmetry of a given tree. The aim of this manuscript is thus twofold: First, we will introduce the emph{symmetry nodes index} as an index for measuring balance of phylogenetic trees and analyze its extremal properties. We also show that this index can be calculated in linear time. This new index turns out to be a generalization of a simple and well-known balance index, namely the emph{cherry index}, as well as a specialization of another, less established, balance index, namely emph{Rogers $J$ index}. Thus, it is the second objective of the present manuscript to compare the new symmetry nodes index to these two indices and to underline its advantages. In order to do so, we will derive some extremal properties of the cherry index and Rogers $J$ index along the way and thus complement existing studies on these indices. Moreover, we used the programming language textsf{R} to implement all three indices in the software package textsf{symmeTree}, which has been made publicly available.
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Measures of tree balance play an important role in various research areas, for example in phylogenetics. There they are for instance used to test whether an observed phylogenetic tree differs significantly from a tree generated by the Yule model of speciation. One of the most popular indices in this regard is the Colless index, which measures the degree of balance for rooted binary trees. While many statistical properties of the Colless index (e.g. asymptotic results for its mean and variance under different models of speciation) have already been discussed in different contexts, we focus on its extremal properties. While it is relatively straightforward to characterize trees with maximal Colless index, the analysis of the minimal value of the Colless index and the characterization of trees that achieve it, are much more involved. In this note, we therefore focus on the minimal value of the Colless index for any given number of leaves. We derive both a recursive formula for this minimal value, as well as an explicit expression, which shows a surprising connection between the Colless index and the so-called Blancmange curve, a fractal curve that is also known as the Takagi curve. Moreover, we characterize two classes of trees that have minimal Colless index, consisting of the set of so-called emph{maximally balanced trees} and a class of trees that we call emph{greedy from the bottom trees}. Furthermore, we derive an upper bound for the number of trees with minimal Colless index by relating these trees with trees with minimal Sackin index (another well-studied index of tree balance).
In phylogenetics it is of interest for rate matrix sets to satisfy closure under matrix multiplication as this makes finding the set of corresponding transition matrices possible without having to compute matrix exponentials. It is also advantageous to have a small number of free parameters as this, in applications, will result in a reduction of computation time. We explore a method of building a rate matrix set from a rooted tree structure by assigning rates to internal tree nodes and states to the leaves, then defining the rate of change between two states as the rate assigned to the most recent common ancestor of those two states. We investigate the properties of these matrix sets from both a linear algebra and a graph theory perspective and show that any rate matrix set generated this way is closed under matrix multiplication. The consequences of setting two rates assigned to internal tree nodes to be equal are then considered. This methodology could be used to develop parameterised models of amino acid substitution which have a small number of parameters but convey biological meaning.
Spontaneous symmetry breaking is central to our understanding of physics and explains many natural phenomena, from cosmic scales to subatomic particles. Its use for applications requires devices with a high level of symmetry, but engineered systems are always imperfect. Surprisingly, the impact of such imperfections has barely been studied, and restricted to a single asymmetry. Here, we experimentally study spontaneous symmetry breaking with two controllable asymmetries. We remarkably find that features typical of spontaneous symmetry breaking, while destroyed by one asymmetry, can be restored by introducing a second asymmetry. In essence, asymmetries are found to balance each other. Our study illustrates aspects of the universal unfolding of the pitchfork bifurcation, and provides new insights into a key fundamental process. It also has practical implications, showing that asymmetry can be exploited as an additional degree of freedom. In particular, it would enable sensors based on symmetry breaking or exceptional points to reach divergent sensitivity even in presence of imperfections. Our experimental implementation built around an optical fiber ring additionally constitutes the first observation of the polarization symmetry breaking of passive driven nonlinear resonators.
We study a continuous-time dynamical system that models the evolving distribution of genotypes in an infinite population where genomes may have infinitely many or even a continuum of loci, mutations accumulate along lineages without back-mutation, added mutations reduce fitness, and recombination occurs on a faster time scale than mutation and selection. Some features of the model, such as existence and uniqueness of solutions and convergence to the dynamical system of an approximating sequence of discrete time models, were presented in earlier work by Evans, Steinsaltz, and Wachter for quite general selective costs. Here we study a special case where the selective cost of a genotype with a given accumulation of ancestral mutations from a wild type ancestor is a sum of costs attributable to each individual mutation plus successive interaction contributions from each $k$-tuple of mutations for $k$ up to some finite ``degree. Using ideas from complex chemical reaction networks and a novel Lyapunov function, we establish that the phenomenon of mutation-selection balance occurs for such selection costs under mild conditions. That is, we show that the dynamical system has a unique equilibrium and that it converges to this equilibrium from all initial conditions.
A tree-based network $N$ on $X$ is called universal if every phylogenetic tree on $X$ is a base tree for $N$. Recently, binary universal tree-based networks have attracted great attention in the literature and their existence has been analyzed in various studies. In this note, we extend the analysis to non-binary networks and show that there exist both a rooted and an unrooted non-binary universal tree-based network with $n$ leaves for all positive integers $n$.
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