Phylogenetics
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Lecture Notes: the Mathematics of Phylogenetics
Lecture Notes: The Mathematics of Phylogenetics Elizabeth S. Allman, John A. Rhodes IAS/Park City Mathematics Institute June-July, 2005 University of Alaska Fairbanks Spring 2009, 2012, 2016 c 2005, Elizabeth S. Allman and John A. Rhodes ii Contents 1 Sequences and Molecular Evolution 3 1.1 DNA structure . .4 1.2 Mutations . .5 1.3 Aligned Orthologous Sequences . .7 2 Combinatorics of Trees I 9 2.1 Graphs and Trees . .9 2.2 Counting Binary Trees . 14 2.3 Metric Trees . 15 2.4 Ultrametric Trees and Molecular Clocks . 17 2.5 Rooting Trees with Outgroups . 18 2.6 Newick Notation . 19 2.7 Exercises . 20 3 Parsimony 25 3.1 The Parsimony Criterion . 25 3.2 The Fitch-Hartigan Algorithm . 28 3.3 Informative Characters . 33 3.4 Complexity . 35 3.5 Weighted Parsimony . 36 3.6 Recovering Minimal Extensions . 38 3.7 Further Issues . 39 3.8 Exercises . 40 4 Combinatorics of Trees II 45 4.1 Splits and Clades . 45 4.2 Refinements and Consensus Trees . 49 4.3 Quartets . 52 4.4 Supertrees . 53 4.5 Final Comments . 54 4.6 Exercises . 55 iii iv CONTENTS 5 Distance Methods 57 5.1 Dissimilarity Measures . 57 5.2 An Algorithmic Construction: UPGMA . 60 5.3 Unequal Branch Lengths . 62 5.4 The Four-point Condition . 66 5.5 The Neighbor Joining Algorithm . 70 5.6 Additional Comments . 72 5.7 Exercises . 73 6 Probabilistic Models of DNA Mutation 81 6.1 A first example . 81 6.2 Markov Models on Trees . 87 6.3 Jukes-Cantor and Kimura Models . -
Species Concepts Should Not Conflict with Evolutionary History, but Often Do
ARTICLE IN PRESS Stud. Hist. Phil. Biol. & Biomed. Sci. xxx (2008) xxx–xxx Contents lists available at ScienceDirect Stud. Hist. Phil. Biol. & Biomed. Sci. journal homepage: www.elsevier.com/locate/shpsc Species concepts should not conflict with evolutionary history, but often do Joel D. Velasco Department of Philosophy, University of Wisconsin-Madison, 5185 White Hall, 600 North Park St., Madison, WI 53719, USA Department of Philosophy, Building 90, Stanford University, Stanford, CA 94305, USA article info abstract Keywords: Many phylogenetic systematists have criticized the Biological Species Concept (BSC) because it distorts Biological Species Concept evolutionary history. While defences against this particular criticism have been attempted, I argue that Phylogenetic Species Concept these responses are unsuccessful. In addition, I argue that the source of this problem leads to previously Phylogenetic Trees unappreciated, and deeper, fatal objections. These objections to the BSC also straightforwardly apply to Taxonomy other species concepts that are not defined by genealogical history. What is missing from many previous discussions is the fact that the Tree of Life, which represents phylogenetic history, is independent of our choice of species concept. Some species concepts are consistent with species having unique positions on the Tree while others, including the BSC, are not. Since representing history is of primary importance in evolutionary biology, these problems lead to the conclusion that the BSC, along with many other species concepts, are unacceptable. If species are to be taxa used in phylogenetic inferences, we need a history- based species concept. Ó 2008 Elsevier Ltd. All rights reserved. When citing this paper, please use the full journal title Studies in History and Philosophy of Biological and Biomedical Sciences 1. -
In Defence of the Three-Domains of Life Paradigm P.T.S
van der Gulik et al. BMC Evolutionary Biology (2017) 17:218 DOI 10.1186/s12862-017-1059-z REVIEW Open Access In defence of the three-domains of life paradigm P.T.S. van der Gulik1*, W.D. Hoff2 and D. Speijer3* Abstract Background: Recently, important discoveries regarding the archaeon that functioned as the “host” in the merger with a bacterium that led to the eukaryotes, its “complex” nature, and its phylogenetic relationship to eukaryotes, have been reported. Based on these new insights proposals have been put forward to get rid of the three-domain Model of life, and replace it with a two-domain model. Results: We present arguments (both regarding timing, complexity, and chemical nature of specific evolutionary processes, as well as regarding genetic structure) to resist such proposals. The three-domain Model represents an accurate description of the differences at the most fundamental level of living organisms, as the eukaryotic lineage that arose from this unique merging event is distinct from both Archaea and Bacteria in a myriad of crucial ways. Conclusions: We maintain that “a natural system of organisms”, as proposed when the three-domain Model of life was introduced, should not be revised when considering the recent discoveries, however exciting they may be. Keywords: Eucarya, LECA, Phylogenetics, Eukaryogenesis, Three-domain model Background (English: domain) as a higher taxonomic level than The discovery that methanogenic microbes differ funda- regnum: introducing the three domains of cellular life, Ar- mentally from Bacteria such as Escherichia coli or Bacillus chaea, Bacteria and Eucarya. This paradigm was proposed subtilis constitutes one of the most important biological by Woese, Kandler and Wheelis in PNAS in 1990 [2]. -
Phylogenetics Topic 2: Phylogenetic and Genealogical Homology
Phylogenetics Topic 2: Phylogenetic and genealogical homology Phylogenies distinguish homology from similarity Previously, we examined how rooted phylogenies provide a framework for distinguishing similarity due to common ancestry (HOMOLOGY) from non-phylogenetic similarity (ANALOGY). Here we extend the concept of phylogenetic homology by making a further distinction between a HOMOLOGOUS CHARACTER and a HOMOLOGOUS CHARACTER STATE. This distinction is important to molecular evolution, as we often deal with data comprised of homologous characters with non-homologous character states. The figure below shows three hypothetical protein-coding nucleotide sequences (for simplicity, only three codons long) that are related to each other according to a phylogenetic tree. In the figure the nucleotide sequences are aligned to each other; in so doing we are making the implicit assumption that the characters aligned vertically are homologous characters. In the specific case of nucleotide and amino acid alignments this assumption is called POSITIONAL HOMOLOGY. Under positional homology it is implicit that a given position, say the first position in the gene sequence, was the same in the gene sequence of the common ancestor. In the figure below it is clear that some positions do not have identical character states (see red characters in figure below). In such a case the involved position is considered to be a homologous character, while the state of that character will be non-homologous where there are differences. Phylogenetic perspective on homologous characters and homologous character states ACG TAC TAA SYNAPOMORPHY: a shared derived character state in C two or more lineages. ACG TAT TAA These must be homologous in state. -
Phylogenetics of Buchnera Aphidicola Munson Et Al., 1991
Türk. entomol. derg., 2019, 43 (2): 227-237 ISSN 1010-6960 DOI: http://dx.doi.org/10.16970/entoted.527118 E-ISSN 2536-491X Original article (Orijinal araştırma) Phylogenetics of Buchnera aphidicola Munson et al., 1991 (Enterobacteriales: Enterobacteriaceae) based on 16S rRNA amplified from seven aphid species1 Farklı yaprak biti türlerinden izole edilen Buchnera aphidicola Munson et al., 1991 (Enterobacteriales: Enterobacteriaceae)’nın 16S rRNA’ya göre filogenetiği Gül SATAR2* Abstract The obligate symbiont, Buchnera aphidicola Munson et al., 1991 (Enterobacteriales: Enterobacteriaceae) is important for the physiological processes of aphids. Buchnera aphidicola genes detected in seven aphid species, collected in 2017 from different plants and altitudes in Adana Province, Turkey were analyzed to reveal phylogenetic interactions between Buchnera and aphids. The 16S rRNA gene was amplified and sequenced for this purpose and a phylogenetic tree built up by the neighbor-joining method. A significant correlation between B. aphidicola genes and the aphid species was revealed by this phylogenetic tree and the haplotype network. Specimens collected in Feke from Solanum melongena L. was distinguished from the other B. aphidicola genes on Aphis gossypii Glover, 1877 (Hemiptera: Aphididae) with a high bootstrap value of 99. Buchnera aphidicola in Myzus spp. was differentiated from others, and the difference between Myzus cerasi (Fabricius, 1775) and Myzus persicae (Sulzer, 1776) was clear. Although, B. aphidicola is specific to its host aphid, certain nucleotide differences obtained within the species could enable specification to geographic region or host plant in the future. Keywords: Aphid, genetic similarity, phylogenetics, symbiotic bacterium Öz Obligat simbiyont, Buchnera aphidicola Munson et al., 1991 (Enterobacteriales: Enterobacteriaceae), yaprak bitlerinin fizyolojik olaylarının sürdürülmesinde önemli bir rol oynar. -
An Introduction to Phylogenetic Analysis
This article reprinted from: Kosinski, R.J. 2006. An introduction to phylogenetic analysis. Pages 57-106, in Tested Studies for Laboratory Teaching, Volume 27 (M.A. O'Donnell, Editor). Proceedings of the 27th Workshop/Conference of the Association for Biology Laboratory Education (ABLE), 383 pages. Compilation copyright © 2006 by the Association for Biology Laboratory Education (ABLE) ISBN 1-890444-09-X All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording, or otherwise, without the prior written permission of the copyright owner. Use solely at one’s own institution with no intent for profit is excluded from the preceding copyright restriction, unless otherwise noted on the copyright notice of the individual chapter in this volume. Proper credit to this publication must be included in your laboratory outline for each use; a sample citation is given above. Upon obtaining permission or with the “sole use at one’s own institution” exclusion, ABLE strongly encourages individuals to use the exercises in this proceedings volume in their teaching program. Although the laboratory exercises in this proceedings volume have been tested and due consideration has been given to safety, individuals performing these exercises must assume all responsibilities for risk. The Association for Biology Laboratory Education (ABLE) disclaims any liability with regards to safety in connection with the use of the exercises in this volume. The focus of ABLE is to improve the undergraduate biology laboratory experience by promoting the development and dissemination of interesting, innovative, and reliable laboratory exercises. -
Introductory Activities
TEACHER’S GUIDE Introduction Dean Madden Introductory NCBE, University of Reading activities Version 1.0 CaseCase Studies introduction Introductory activities The activities in this section explain the basic principles behind the construction of phylogenetic trees, DNA structure and sequence alignment. Students are also intoduced to the Geneious software. Before carrying out the activities in the DNA to Darwin Case studies, students will need to understand: • the basic principles behind the construction of an evolutionary tree or phylogeny; • the basic structure of DNA and proteins; • the reasons for and the principle of alignment; • use of some features of the Geneious computer software (basic version). The activities in this introduction are designed to achieve this. Some of them will reinforce what students may already know; others involve new concepts. The material includes extension activities for more able students. Evolutionary trees In 1837, 12 years before the publication of On the Origin of Species, Charles Darwin famously drew an evolutionary tree in one of his notebooks. The Origin also included a diagram of an evolutionary tree — the only illustration in the book. Two years before, Darwin had written to his friend Thomas Henry Huxley, saying: ‘The time will come, I believe, though I shall not live to see it, when we shall have fairly true genealogical trees of each great kingdom of Nature.’ Today, scientists are trying to produce the ‘Tree of Life’ Darwin foresaw, using protein, DNA and RNA sequence data. Evolutionary trees are covered on pages 2–7 of the Student’s guide and in the PowerPoint and Keynote slide presentations. -
A Phylogenomic Analysis of Turtles ⇑ Nicholas G
Molecular Phylogenetics and Evolution 83 (2015) 250–257 Contents lists available at ScienceDirect Molecular Phylogenetics and Evolution journal homepage: www.elsevier.com/locate/ympev A phylogenomic analysis of turtles ⇑ Nicholas G. Crawford a,b,1, James F. Parham c, ,1, Anna B. Sellas a, Brant C. Faircloth d, Travis C. Glenn e, Theodore J. Papenfuss f, James B. Henderson a, Madison H. Hansen a,g, W. Brian Simison a a Center for Comparative Genomics, California Academy of Sciences, 55 Music Concourse Drive, San Francisco, CA 94118, USA b Department of Genetics, University of Pennsylvania, Philadelphia, PA 19104, USA c John D. Cooper Archaeological and Paleontological Center, Department of Geological Sciences, California State University, Fullerton, CA 92834, USA d Department of Biological Sciences, Louisiana State University, Baton Rouge, LA 70803, USA e Department of Environmental Health Science, University of Georgia, Athens, GA 30602, USA f Museum of Vertebrate Zoology, University of California, Berkeley, CA 94720, USA g Mathematical and Computational Biology Department, Harvey Mudd College, 301 Platt Boulevard, Claremont, CA 9171, USA article info abstract Article history: Molecular analyses of turtle relationships have overturned prevailing morphological hypotheses and Received 11 July 2014 prompted the development of a new taxonomy. Here we provide the first genome-scale analysis of turtle Revised 16 October 2014 phylogeny. We sequenced 2381 ultraconserved element (UCE) loci representing a total of 1,718,154 bp of Accepted 28 October 2014 aligned sequence. Our sampling includes 32 turtle taxa representing all 14 recognized turtle families and Available online 4 November 2014 an additional six outgroups. Maximum likelihood, Bayesian, and species tree methods produce a single resolved phylogeny. -
Phylogenetic Comparative Methods: a User's Guide for Paleontologists
Phylogenetic Comparative Methods: A User’s Guide for Paleontologists Laura C. Soul - Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC, USA David F. Wright - Division of Paleontology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024, USA and Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC, USA Abstract. Recent advances in statistical approaches called Phylogenetic Comparative Methods (PCMs) have provided paleontologists with a powerful set of analytical tools for investigating evolutionary tempo and mode in fossil lineages. However, attempts to integrate PCMs with fossil data often present workers with practical challenges or unfamiliar literature. In this paper, we present guides to the theory behind, and application of, PCMs with fossil taxa. Based on an empirical dataset of Paleozoic crinoids, we present example analyses to illustrate common applications of PCMs to fossil data, including investigating patterns of correlated trait evolution, and macroevolutionary models of morphological change. We emphasize the importance of accounting for sources of uncertainty, and discuss how to evaluate model fit and adequacy. Finally, we discuss several promising methods for modelling heterogenous evolutionary dynamics with fossil phylogenies. Integrating phylogeny-based approaches with the fossil record provides a rigorous, quantitative perspective to understanding key patterns in the history of life. 1. Introduction A fundamental prediction of biological evolution is that a species will most commonly share many characteristics with lineages from which it has recently diverged, and fewer characteristics with lineages from which it diverged further in the past. This principle, which results from descent with modification, is one of the most basic in biology (Darwin 1859). -
Lineages, Splits and Divergence Challenge Whether the Terms Anagenesis and Cladogenesis Are Necessary
Biological Journal of the Linnean Society, 2015, , – . With 2 figures. Lineages, splits and divergence challenge whether the terms anagenesis and cladogenesis are necessary FELIX VAUX*, STEVEN A. TREWICK and MARY MORGAN-RICHARDS Ecology Group, Institute of Agriculture and Environment, Massey University, Palmerston North, New Zealand Received 3 June 2015; revised 22 July 2015; accepted for publication 22 July 2015 Using the framework of evolutionary lineages to separate the process of evolution and classification of species, we observe that ‘anagenesis’ and ‘cladogenesis’ are unnecessary terms. The terms have changed significantly in meaning over time, and current usage is inconsistent and vague across many different disciplines. The most popular definition of cladogenesis is the splitting of evolutionary lineages (cessation of gene flow), whereas anagenesis is evolutionary change between splits. Cladogenesis (and lineage-splitting) is also regularly made synonymous with speciation. This definition is misleading as lineage-splitting is prolific during evolution and because palaeontological studies provide no direct estimate of gene flow. The terms also fail to incorporate speciation without being arbitrary or relative, and the focus upon lineage-splitting ignores the importance of divergence, hybridization, extinction and informative value (i.e. what is helpful to describe as a taxon) for species classification. We conclude and demonstrate that evolution and species diversity can be considered with greater clarity using simpler, more transparent terms than anagenesis and cladogenesis. Describing evolution and taxonomic classification can be straightforward, and there is no need to ‘make words mean so many different things’. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 00, 000–000. -
Statistical Inference for the Linguistic and Non-Linguistic Past
Statistical inference for the linguistic and non-linguistic past Igor Yanovich DFG Center for Advanced Study “Words, Bones, Genes and Tools” Universität Tübingen July 6, 2017 Igor Yanovich 1 / 46 Overview of the course Overview of the course 1 Today: trees, as a description and as a process 2 Classes 2-3: simple inference of language-family trees 3 Classes 4-5: computational statistical inference of trees and evolutionary parameters 4 Class 6: histories of languages and of genes 5 Class 7: simple spatial statistics 6 Class 8: regression taking into account linguistic relationships; synthesis of the course Igor Yanovich 2 / 46 Overview of the course Learning outcomes By the end of the course, you should be able to: 1 read and engage with the current literature in linguistic phylogenetics and in spatial statistics for linguistics 2 run phylogenetic and basic spatial analyses on linguistic data 3 proceed further in the subject matter on your own, towards further advances in the field Igor Yanovich 3 / 46 Overview of the course Today’s class 1 Overview of the course 2 Language families and their structures 3 Trees as classifications and as process depictions 4 Linguistic phylogenetics 5 Worries about phylogenetics in linguistics vs. biology 6 Quick overview of the homework 7 Summary of Class 1 Igor Yanovich 4 / 46 Language families and their structures Language families and their structures Igor Yanovich 5 / 46 Language families and their structures Dravidian A modification of [Krishnamurti, 2003, Map 1.1], from Wikipedia Igor Yanovich 6 / 46 Language families and their structures Dravidian [Krishnamurti, 2003]’s classification: Dravidian family South Dravidian Central Dravidian North Dravidian SD I SD II Tamil Malay¯al.am Kannad.a Telugu Kolami Brahui (70M) (38M) (40M) (75M) (0.1M) (4M) (Numbers of speakers from Wikipedia) Igor Yanovich 7 / 46 Language families and their structures Dravidian: similarities and differences Proto-Dr. -
Phylogeny Codon Models • Last Lecture: Poor Man’S Way of Calculating Dn/Ds (Ka/Ks) • Tabulate Synonymous/Non-Synonymous Substitutions • Normalize by the Possibilities
Phylogeny Codon models • Last lecture: poor man’s way of calculating dN/dS (Ka/Ks) • Tabulate synonymous/non-synonymous substitutions • Normalize by the possibilities • Transform to genetic distance KJC or Kk2p • In reality we use codon model • Amino acid substitution rates meet nucleotide models • Codon(nucleotide triplet) Codon model parameterization Stop codons are not allowed, reducing the matrix from 64x64 to 61x61 The entire codon matrix can be parameterized using: κ kappa, the transition/transversionratio ω omega, the dN/dS ratio – optimizing this parameter gives the an estimate of selection force πj the equilibrium codon frequency of codon j (Goldman and Yang. MBE 1994) Empirical codon substitution matrix Observations: Instantaneous rates of double nucleotide changes seem to be non-zero There should be a mechanism for mutating 2 adjacent nucleotides at once! (Kosiol and Goldman) • • Phylogeny • • Last lecture: Inferring distance from Phylogenetic trees given an alignment How to infer trees and distance distance How do we infer trees given an alignment • • Branch length Topology d 6-p E 6'B o F P Edo 3 vvi"oH!.- !fi*+nYolF r66HiH- .) Od-:oXP m a^--'*A ]9; E F: i ts X o Q I E itl Fl xo_-+,<Po r! UoaQrj*l.AP-^PA NJ o - +p-5 H .lXei:i'tH 'i,x+<ox;+x"'o 4 + = '" I = 9o FF^' ^X i! .poxHo dF*x€;. lqEgrE x< f <QrDGYa u5l =.ID * c 3 < 6+6_ y+ltl+5<->-^Hry ni F.O+O* E 3E E-f e= FaFO;o E rH y hl o < H ! E Y P /-)^\-B 91 X-6p-a' 6J.