Phylogenomics Jer-Ming Hu 胡哲明 Plant Systematics and Evolution Group, NTU 台大植物系統分類與演化研究室 本著作除另有註明外,採取創用CC「姓名標示- 非商業性-相同方式分享」台灣3.0版授權釋出 Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike 3.0 Taiwan (CC BY-NC-SA 3.0) 1 14.4. Tue Introduction to models (Jarno) 16.4. Thu Distance-based methods (Jarno) 17.4. Fri ML analyses (Jarno) 20.4. Mon Assessing hypotheses (Jarno) 21.4. Tue Problems with molecular data (Jarno) 23.4. Thu Problems with molecular data (Jarno) Phylogenomics 24.4. Fri Search algorithms, visualization, and other computational aspects (Jarno) J Phylogenomics The uses of phylogenetic principles to make sense of genomic data Recent reviews Eisen, J.A. 1998. Gen. Res. 8: 163-167 Delsuc, F. et al. 2005. Nat. Rev. Genet. 6: 361-375 Koonin, E.V. 2005. Annu. Rev. Genet. 39: 309-338 Philippe, H. et al. 2005. Annu. Rev. Ecol. Evol. Syst. 36: 541-562 Boussau & Daubin (2010) TREE 25: 224-232 3 Current approaches Methods based on primary sequences Supermatrix Supertree Large-scale screening (e.g. ngs data) Methods based on whole-genome features Gene content Gene order DNA strings (distribution of oligonucleotides in genomes) 4 Combining data in phylogenetic analysis Data partition Different genes Coding vs. non-coding regions Among codon positions Molecular vs. morphological data Three ways to deal with partitioned data Always combine (total evidence) Never combine (separate analysis) Conditional combine (perform a test to decide whether or not to combine them) Huelsenbeck, J.P. et al. (1996) TREE 11:152 5 Separate analysis Miyamoto and Fitch (1995) Tree should be estimated from each partition Congruence among different data partitions provides stronger evidence of accurate phylogeny Miyamoto, M.M. and W.M. Fitch (1995) Syst. Biol. 44:64-76 6 Logic of total evidence All independent characters should be included without a priori consideration of data partition Not only character data, but also taxa (e.g. fossil and living) should be included in a phylogenetic analysis. 7 Conditional data combination Bull et al. (1993), de Queiroz (1993) The partitions should be subjected to a homogeneity test before any further process. e.g. KH test, partition homogeneity test, likelihood heterogeneity test, etc. Potential problem with false positives The failure to combine data sets that should have been combined Bull, J.J. et al. (1993) Syst. Biol. 42:384-397 de Queiroz, A. (1993) Syst. Biol. 42:368-372 8 Three ways for treating diverse data Data 1 Data 2 Data 3 …. 1 Test for homogeneity Accept Reject 2 3 Combined data Separated data National Taiwan University Jer-Ming Hu 9 The challenge of constructing large phylogenetic trees Data availability matrix Based on 130304 sequences of 14667 green plant species Most GenBank sequences are from commonly used genes and the model species M. J. Sanderson and A. C. Driskell 10 Construct large phylogenies M. J. Sanderson and A. C. Driskell 11 Problems of data concatenation Including multiple genes may mix phylogenetic signals from different evolutionary histories Some sequences are usually unavailable for some species and have to code them as missing data e.g. photosynthetic genes of non-photosynthetic plants 12 Supertree approach: the idea Past: (direct method) Nested source trees were grafted together O. R. P. Bininda-Emonds Present: (indirect method) Overlapping source trees are combined to yield to supertree O. R. P. Bininda-Emonds 13 Supertree: constructing strict supertree Source trees A D E F B C Strict supertree A A B C D E Strict supertree B D E F A National Taiwan University Jer-Ming Hu B C D E F 14 Supertree by supermatrix approach 1. Combination of source data sets 2. Make a new tree M. J. Sanderson, A. Purvis, and C. Henze Sanderson. et al. (1998) TREE 13:105-109 15 Strict supertree algorithm 1. Identify common taxa 2. Construct a backbone tree 3. Examine source tree corresponding to each branch 4. Strict consensus supertree M. J. Sanderson, A. Purvis, and C. Henze 16 Supertree by MRP Matrix Representation with Parsimony (MRP) Remake a new data matrix which characters refer to the topologies of the source trees Baum, B.R. (1992) Taxon 41:3-10 Ragan, M.A. (1992) Mol. Phylo. Evol. 1:53-58 Sanderson. et al. (1998) TREE 13:105-109 17 Supertree by MRP Source trees 1: present in the clade 0: absent in the clade O. R. P. Bininda-Emonds, J. L. Gittleman, and M. A. Steel 18 BUILD algorithm Aho et al. (1981) SIAM J. Comput. 10:405-421 Cluster C = X 4 3 2 1 4 6 5 1 National Taiwan University Jer-Ming Hu C1 = {{1, 2} {1, 2, 3} {1, 2, 3, 4}} C2 = {{1, 5} {1, 5, 6} {1, 5, 6, 4}} 19 BUILD algorithm C’ = {{1,2,3,5,6}, {4}} O. R. P. Bininda-Emonds, J. L. Gittleman, and M. A. Steel Repeatedly evaluating the clusters C’’ = {{1,2,5}, {3}, {6}} C = {{1,2,3,4,5,6}, {1,2,3,5,6}, {1,2,5}, {3}, {4}, {6}} O. R. P. Bininda-Emonds, J. L. Gittleman, and M. A. Steel 20 Two source trees 4 3 2 4 6 3 5 2 1 4 1 The BUILD tree 6 5 4 6 3 5 1 2 1 The MRP tree National Taiwan University Jer-Ming Hu 21 Current supertree methods O. R. P. Bininda-Emonds Bininda-Emonds, O.R.P. (2004) TREE 19.315 22 O. R. P. Bininda-Emonds 23 Supertree: the future A divide-and-conquer approach can be applied, to decompose very large phylogenetic problem into many smaller subproblems, and the solutions to which are combined to derive the global answer Similar approaches have been used in phylogenetics, e.g. compartmentalization and quartet puzzling Two new approaches • • Disk-covering methods (DCMs) The biclique method Bininda-Emonds, O.R.P. (2004) TREE 19.315 24 Disk-covering method (DCM) O. R. P. Bininda-Emonds 25 The biclique method O. R. P. Bininda-Emonds 26 Critics of supertree approach It does not take sequence info into account The dataset is non-independent MRP favors larger source trees because they contribute more characters Some direct methods cannot incorporate support values into the source trees 27 Programs available supertree Rod Page http://darwin.zoology.gla.ac.uk/~rpage/supertree/ SuperTree Salamin et al. (2002) http://www2.unil.ch/phylo/bioinformatics/supertree.html Supertrees related Olaf R.P. Bininda-Emonds http://www.molekularesystematik.uni-oldenburg.de/en/34011.html 28 ... okay, let’s see some real data Supertree of legumes related to Astragalus Sanderson, M.J. et al. (1998) TREE 13:105-109 Supertree of prokaryotes Daubin, V. et al. (2002) Genome Res. 12:1080 Supertree of the angiosperms Davies, T.J. et al. (2004) PNAS 101:1904-1909 29 M. J. Sanderson, A. Purvis, and C. Henze Bold faces: taxa in common M. J. Sanderson, A. Purvis, and C. Henze M. J. Sanderson, A. Purvis, and C. Henze 30 Genome Research (2002) 12:1080-1090 A Phylogenomic Approach to Bacterial Phylogeny: Evidence of a Core of Genes Sharing a Common History Vincent Daubin, Manolo Gouy, and Guy Perriere Supertree using MRP • 730 source trees, include 45 taxa • Bootstrap % used as character weights • BIONJ + gamma • Identify the core of genes • Principal Coordinates analysis (PCO) for comparing tree distance (Robinson-Foulds distance) • V. Daubin, M. Gouy, and G. Perriere 31 Supertree made w/ ML trees Chlamydiales Spirochaetes Hyperthermophilic bacteria Cyanobacteria Low GC Gram+ Deinococcus High GC Gram+ Proteobacteria Crenarchaeota Euryarchaeota Eukaryota V. Daubin, M. Gouy, and G. Perriere Supertree constructed from 730 gene trees 32 Supertree approach Plot of the first two axes of the PCO (Principle Coordinate Analysis) Identifying “core genes” Choose only the “core genes & perform supertree searching again V. Daubin, M. Gouy, and G. Perriere 33 Supertree made w/ ML trees after PCO Spirochaetes Chlamydiales Hyperthermophilic bacteria Low GC Gram+ Cyanobacteria Deinococcus High GC Gram+ Proteobacteria Euryarchaeota Crenarchaeota Eukaryota V. Daubin, M. Gouy, and G. Perriere 34 PNAS (2004) 101:1904-1909 Darwin's abominable mystery: Insights from a supertree of the angiosperms T. Johathan Davies, Timothy G. Barraclough, Mark W. Chase, Pamela S. Soltis, Douglas E. Soltis, and Vincent Savolainen Supertree using MRP • 46 source trees, include 379 taxa • Bootstrap % used as character weights • Examining diversification rates • V. T. J. Davies, T. G. Barraclough, M. W. Chase, P. S. Soltis, D. E. Soltis, and V. Savolainen 35 PNAS (2004) 101:1904-1909 Darwin's abominable mystery: Insights from a supertree of the angiosperms T. Johathan Davies, Timothy G. Barraclough, Mark W. Chase, Pamela S. Soltis, Douglas E. Soltis, and Vincent Savolainen Supporting Table 2 36 1 of 10000 most parsimonious supertrees with dates obtained by NPRS of ML using rbcL Diversification rates Low Low - medium Medium - high High Time calibrated by 84 mya of Fagales/Cucurbitales split V. T. J. Davies, T. G. Barraclough, M. W. Chase, P. S. Soltis, D. E. Soltis, and V. Savolainen 37 V. T. J. Davies, T. G. Barraclough, M. W. Chase, P. S. Soltis, D. E. Soltis, and V. Savolainen No ‘key innovations’ can be identified 38 Phylogenomics using whole-genome features Distribution of sequence strings Transformation of the frequencies of small oligonucleotides (up to eight) or oligopeptides (up to six) into distances and used for phylogenetic reconstruction Gene order methods Gene content methods 39 Genome sequences Analyzed the tetranucleotide usage deviations from expectations (TUD) AAAA AAAC AAAG AAAT ... CAAA CAAC ... TTTT Compare among different genomes Use the similarity measures to construct a distance based phylogeny National Taiwan University Jer-Ming Hu 40 Potential problems using sequence strings The phylogeneic information of sequence strings saturates rapidly No models of evolution is proposed for such distribution of oligonucleotide or oligopeptide frequencies 41 16S rRNA tree TUD tree D. T. Pride, R. J. Meinersmann, T. M. Wassenaar, and M. J. Blaser Gram-negative Gram-positive Archaea and thermophilic bacteria Gram-negative 42 Gene order method Information of genome organization Distances of two genomes can be computed to minimize the numbers of inversions, transpositions, insertions, and deletions Distances can also be computed for the minimum number of pairs of genes next to each other in one of the two genomes, but not in the other (“break-point distance”) 43 Distribution of conserved gene pairs among 31 clades of prokaryotes Wolf et al. (2001) BMC Evol. Biol. 1:8 Y. I. Wolf, I. B. Rogozin, N. V. Grishin, R. L. Tatusov, E. V. Koonin Gene-order phylogeny ss rRNA phylogeny J. O. Korbel, B. Snel, M. A. Huynen, and P. Bork J. O. Korbel, B. Snel, M. A. Huynen, and P. Bork 44 Gene content method Construct distance or parsimony phylogeny based on the proportion of shared orthologous genes between genomes A data matrix is made for the presence/absence of orthologous genes • Note: universal present genes will become parsimonious uninformative! 45 Example of comparing gene contents B. Snel, P. Bork, M. A. Huynen 1. Large genomes have many genes in common 2. The number of genes two genomes have in common depends on their evolutionary distance 46 Gene-content phylogeny ss rRNA phylogeny J. O. Korbel, B. Snel, M. A. Huynen, and P. Bork J. O. Korbel, B. Snel, M. A. Huynen, and P. Bork Note that proteobacteria in gene-content phylogeny is not monophyletic 47 Problems using gene-content method Big/small genome attraction Artificial grouping of unrelated species with small genomes, e.g. Mycoplasma, Buchnera, Chlamydia, or Rickettsia. Phylogenetic positions of some taxa do not agree with previous studies Halobacterium never clusters with Methanosarcina. This is probably due to too many HGTs 48 Programs available SHOT A web server for the construction of genome phylogenies http://coot.embl.de/~korbel/SHOT/ 49 Summary R. Delsuc, H. Brinkmann, and H. Philippe 50 Large scale screening of orthologous genes for phylogenomic analysis G. D. Edgecombe, G. Giribet, C. W. Dunn, A. Hejnol, R. M. Kristensen, R. C. Neves, G. W. Rouse, K. Worsaae, and M. V. Sørensen This work is licensed by SpringerLink for the use of “Course Database of General Education TW” ONLY. The copyright belongs to the above mentioned entity and GET does not have the authorization right. Copyright privileges have to be negotiated with the copyright owner(s) for separately. G. D. Edgecombe, G. Giribet, C. W. Dunn, A. Hejnol, R. M. Kristensen, R. C. Neves, G. W. Rouse, K. Worsaae, and M. V. Sørensen This work is licensed by SpringerLink for the use of “Course Database of General Education TW” ONLY. The copyright belongs to the above mentioned entity and GET does not have the authorization right. Copyright privileges have to be negotiated with the copyright owner(s) for separately. 51 Flow chart for EST-based phylogenomic analysis C. W. Dunn, A. Hejnol, D. Q. Matus, K. Pang, W. E. Browne, S. A. Smith, E. Seaver, G. W. Rouse, M. Obst, G. D. Edgecombe, M. V. Sørensen6, S. H. D. Haddock, A. Schmidt-Rhaesa, A. Okusu, R. M. Kristensen, W. C. Wheeler, M. Q. Martindale, and G. Giribet 52 Phylogeny based on 77 taxa, 150 genes, of the animal tree of life C. W. Dunn, A. Hejnol, D. Q. Matus, K. Pang, W. E. Browne, S. A. Smith, E. Seaver, G. W. Rouse, M. Obst, G. D. Edgecombe, M. V. Sørensen6, S. H. D. Haddock, A. Schmidt-Rhaesa, A. Okusu, R. M. Kristensen, W. C. Wheeler, M. Q. Martindale, and G. Giribet Dunn et al. (2008) Nature 452: 745 53 The myth of orthologous gene families Bastien Boussau, and Vincent Daubin 54 “Phylogenetic awareness” on making tree Bastien Boussau, and Vincent Daubin 55 Problems of just using whole-genome data Case study of seeking the root of angiosperms Amborella or monocots? Flickr scott.zona Flickr adrien2008 56 Amborella Flickr pennstatelive Amborella as the first branch has been identified in 18S rDNA, chl. atpB, phyA/C, mt coxI, matR, and combined analysis S. Stefanović, D. W. Rice, and J. D. Palmer 57 Phylogeny of monocots being basal according to complete chloroplast genomes Eudicots Monocots V. V. Goremykin, K. I. Hirsch-Ernst, S. Wolfl, and F. H. Hellwig 58 •No rate categories •1st/2nd pos only •4 gamma rate categories •1st/2nd pos only •No rate categories •Estimated proportion of invariant sites •1st/2nd pos only •No rate categories •all pos S. Stefanović, D. W. Rice, and J. D. Palmer 59 Effects of taxon sampling Method: DNA ML HKY85 with no rate categories Sampling according to Goremykin et al. (2004) Acorus instead of grasses Acorus and grasses included S. Stefanović, D. W. Rice, and J. D. Palmer Long Branch Attraction effect 60 Copyright Declaration Work Licensing Author/Source Page P8 National Taiwan University Jer-Ming Hu M. J. Sanderson and A. C. Driskell 2003. The challenge of constructing large phylogenetic trees. Trends in Plant Science 8: p. 374. http://www.sciencedirect.com/science/article/pii/S1360138503001651 It is used subject to the fair use doctrine of: •Taiwan Copyright Act Articles 52 & 65 •Elsevier Website Terms and Conditions http://www.elsevier.com/wps/find/termsconditions.cws_home/termsconditions P9 M. J. Sanderson and A. C. Driskell 2003. The challenge of constructing large phylogenetic trees. Trends in Plant Science 8: p. 375. http://www.sciencedirect.com/science/article/pii/S1360138503001651 It is used subject to the fair use doctrine of: •Taiwan Copyright Act Articles 52 & 65 •Elsevier Website Terms and Conditions http://www.elsevier.com/wps/find/termsconditions.cws_home/termsconditions P10 O. R. P. Bininda-Emonds 2004. The evolution of supertrees. Trends in Ecology & Evolution 19: p. 315. http://www.sciencedirect.com/science/article/pii/S0169534704000709 It is used subject to the fair use doctrine of: •Taiwan Copyright Act Articles 52 & 65 •Elsevier Website Terms and Conditions http://www.elsevier.com/wps/find/termsconditions.cws_home/termsconditions P12 O. R. P. Bininda-Emonds 2004. The evolution of supertrees. Trends in Ecology & Evolution 19: p. 315. http://www.sciencedirect.com/science/article/pii/S0169534704000709 It is used subject to the fair use doctrine of: •Taiwan Copyright Act Articles 52 & 65 •Elsevier Website Terms and Conditions http://www.elsevier.com/wps/find/termsconditions.cws_home/termsconditions P12 61 Work Licensing Author/Source Page P13 National Taiwan University Jer-Ming Hu M. J. Sanderson, A. Purvis, and C. Henze 1998. Phylogenetic supertrees: Assembling the trees of life. Trends in Ecology & Evolution 13: p. 106. http://www.cell.com/trends/ecology-evolution/abstract/S0169-5347%2897%2901242-1 It is used subject to the fair use doctrine of: •Taiwan Copyright Act Articles 52 & 65 •Cell Press Copyrights and Limitations on Use •Elsevier Website Terms and Conditions •The "Code of Best Practices in Fair Use for OpenCourseWare 2009 (http://www.centerforsocialmedia.org/sites/default/files/10-305-OCW-Oct29.pdf)" by A Committee of Practitioners of OpenCourseWare in the U.S. The contents are based on Section 107 of the 1976 U.S. Copyright Act P14 M. J. Sanderson, A. Purvis, and C. Henze 1998. Phylogenetic supertrees: Assembling the trees of life. Trends in Ecology & Evolution 13: p. 107. http://www.cell.com/trends/ecology-evolution/abstract/S0169-5347%2897%2901242-1 It is used subject to the fair use doctrine of: •Taiwan Copyright Act Articles 52 & 65 •Cell Press Copyrights and Limitations on Use •Elsevier Website Terms and Conditions •The "Code of Best Practices in Fair Use for OpenCourseWare 2009 (http://www.centerforsocialmedia.org/sites/default/files/10-305-OCW-Oct29.pdf)" by A Committee of Practitioners of OpenCourseWare in the U.S. The contents are based on Section 107 of the 1976 U.S. Copyright Act P15 O. R. P. Bininda-Emonds, J. L. Gittleman, and M. A. Steel 2002. THE (SUPER)TREE OF LIFE: Procedures, Problems, and Prospects. Annual Review of Ecology and Systematics 33: p. 268. http://www.annualreviews.org/doi/pdf/10.1146/annurev.ecolsys.33.010802.150511 It is used subject to the fair use doctrine of: •Taiwan Copyright Act Articles 52 & 65 •Annual Reviews Copyright & Permissions http://www.annualreviews.org/page/about/copyright-and-permissions P17 62 Work Licensing Author/Source Page P18 National Taiwan University Jer-Ming Hu O. R. P. Bininda-Emonds, J. L. Gittleman, and M. A. Steel 2002. THE (SUPER)TREE OF LIFE: Procedures, Problems, and Prospects. Annual Review of Ecology and Systematics 33: p. 273. http://www.annualreviews.org/doi/abs/10.1146/annurev.ecolsys.33.010802.150511?journalCo de=ecolsys.1 It is used subject to the fair use doctrine of: •Taiwan Copyright Act Articles 52 & 65 •Annual Reviews Copyright & Permissions http://www.annualreviews.org/page/about/copyright-and-permissions P19 O. R. P. Bininda-Emonds, J. L. Gittleman, and M. A. Steel 2002. THE (SUPER)TREE OF LIFE: Procedures, Problems, and Prospects. Annual Review of Ecology and Systematics 33: p. 273. http://www.annualreviews.org/doi/abs/10.1146/annurev.ecolsys.33.010802.150511?journalCo de=ecolsys.1 It is used subject to the fair use doctrine of: •Taiwan Copyright Act Articles 52 & 65 •Annual Reviews Copyright & Permissions http://www.annualreviews.org/page/about/copyright-and-permissions P19 P20 National Taiwan University Jer-Ming Hu O. R. P. Bininda-Emonds 2004. The evolution of supertrees. Trends in Ecology & Evolution 19: p. 317. http://www.sciencedirect.com/science/article/pii/S0169534704000709 It is used subject to the fair use doctrine of: •Taiwan Copyright Act Articles 52 & 65 •Elsevier Website Terms and Conditions http://www.elsevier.com/wps/find/termsconditions.cws_home/termsconditions P21 63 Work Licensing Author/Source Page O. R. P. Bininda-Emonds 2004. The evolution of supertrees. Trends in Ecology & Evolution 19: p. 318. http://www.sciencedirect.com/science/article/pii/S0169534704000709 It is used subject to the fair use doctrine of: •Taiwan Copyright Act Articles 52 & 65 •Elsevier Website Terms and Conditions http://www.elsevier.com/wps/find/termsconditions.cws_home/termsconditions P22 O. R. P. Bininda-Emonds 2004. The evolution of supertrees. Trends in Ecology & Evolution 19: p. 320. http://www.sciencedirect.com/science/article/pii/S0169534704000709 It is used subject to the fair use doctrine of: •Taiwan Copyright Act Articles 52 & 65 •Elsevier Website Terms and Conditions http://www.elsevier.com/wps/find/termsconditions.cws_home/termsconditions P24 O. R. P. Bininda-Emonds 2004. The evolution of supertrees. Trends in Ecology & Evolution 19: p. 320. http://www.sciencedirect.com/science/article/pii/S0169534704000709 It is used subject to the fair use doctrine of: •Taiwan Copyright Act Articles 52 & 65 •Elsevier Website Terms and Conditions http://www.elsevier.com/wps/find/termsconditions.cws_home/termsconditions P25 M. J. Sanderson, A. Purvis, and C. Henze 1998. Phylogenetic supertrees: Assembling the trees of life. Trends in Ecology & Evolution 13: p. 108. http://download.cell.com/trends/ecology-evolution/pdf/PIIS0169534797012421.pdf It is used subject to the fair use doctrine of: •Taiwan Copyright Act Articles 52 & 65 •Cell Press Copyrights and Limitations on Use •Elsevier Website Terms and Conditions •The "Code of Best Practices in Fair Use for OpenCourseWare 2009 (http://www.centerforsocialmedia.org/sites/default/files/10-305-OCW-Oct29.pdf)" by A Committee of Practitioners of OpenCourseWare in the U.S. The contents are based on Section 107 of the 1976 U.S. Copyright Act P29 64 Work Supertree using MRP 730 source trees, … Principal Coordinates analysis (PCO) for comparing tree distance (Robinson-Foulds distance) Licensing Author/Source Page M. J. Sanderson, A. Purvis, and C. Henze 1998. Phylogenetic supertrees: Assembling the trees of life. Trends in Ecology & Evolution 13: p. 108. http://download.cell.com/trends/ecology-evolution/pdf/PIIS0169534797012421.pdf It is used subject to the fair use doctrine of: •Taiwan Copyright Act Articles 52 & 65 •Cell Press Copyrights and Limitations on Use •Elsevier Website Terms and Conditions •The "Code of Best Practices in Fair Use for OpenCourseWare 2009 (http://www.centerforsocialmedia.org/sites/default/files/10-305-OCW-Oct29.pdf)" by A Committee of Practitioners of OpenCourseWare in the U.S. The contents are based on Section 107 of the 1976 U.S. Copyright Act P29 M. J. Sanderson, A. Purvis, and C. Henze 1998. Phylogenetic supertrees: Assembling the trees of life. Trends in Ecology & Evolution 13: p. 108. http://download.cell.com/trends/ecology-evolution/pdf/PIIS0169534797012421.pdf It is used subject to the fair use doctrine of: •Taiwan Copyright Act Articles 52 & 65 •Cell Press Copyrights and Limitations on Use •Elsevier Website Terms and Conditions •The "Code of Best Practices in Fair Use for OpenCourseWare 2009 (http://www.centerforsocialmedia.org/sites/default/files/10-305-OCW-Oct29.pdf)" by A Committee of Practitioners of OpenCourseWare in the U.S. The contents are based on Section 107 of the 1976 U.S. Copyright Act P29 Vincent Daubin, Manolo Gouy, and Guy Perriere A Phylogenomic Approach to Bacterial Phylogeny: Evidence of a Core of Genes Sharing a Common History http://genome.cshlp.org/content/12/7/1080.full 2012/06/14 visited P30 V. Daubin, M. Gouy, and G. Perriere. 2002. A Phylogenomic Approach to Bacterial Phylogeny: Evidence of a Core of Genes Sharing a Common History. Genome Research 12: p. 1082. Copyright © 2002, by Cold Spring Harbor Laboratory Press http://genome.cshlp.org/content/12/7/1080.full.pdf+html 2012/06/14 visited P31 V. Daubin, M. Gouy, and G. Perriere. 2002. A Phylogenomic Approach to Bacterial Phylogeny: Evidence of a Core of Genes Sharing a Common History. Genome Research 12: p. 1083. Copyright © 2002, by Cold Spring Harbor Laboratory Press http://genome.cshlp.org/content/12/7/1080.full.pdf+html 2012/06/14 visited P32 65 Work Supertree using MRP 46 source trees, include 379 taxa Bootstrap % used as character weights Examining diversification rates Licensing Author/Source Page V. Daubin, M. Gouy, and G. Perriere. 2002. A Phylogenomic Approach to Bacterial Phylogeny: Evidence of a Core of Genes Sharing a Common History. Genome Research 12: p. 1084. Copyright © 2002, by Cold Spring Harbor Laboratory Press http://genome.cshlp.org/content/12/7/1080.full.pdf 2012/06/14 visited P33 V. T. J. Davies, T. G. Barraclough, M. W. Chase, P. S. Soltis, D. E. Soltis, and V. Savolainen 2004. Darwin's abominable mystery: Insights from a supertree of the angiosperms. PNAS 101: p. 1904-1909. http://www.pnas.org/content/101/7/1904.full.pdf+html It is used subject to the fair use doctrine of: •Taiwan Copyright Act Articles 52 & 65 •The National Academies Terms of Use •The "Code of Best Practices in Fair Use for OpenCourseWare 2009 (http://www.centerforsocialmedia.org/sites/default/files/10-305-OCW-Oct29.pdf)" by A Committee of Practitioners of OpenCourseWare in the U.S. The contents are based on Section 107 of the 1976 U.S. Copyright Act P34 V. T. J. Davies, T. G. Barraclough, M. W. Chase, P. S. Soltis, D. E. Soltis, and V. Savolainen 2004. Darwin's abominable mystery: Insights from a supertree of the angiosperms. PNAS 101: p. 1906. http://www.pnas.org/content/101/7/1904.full.pdf+html It is used subject to the fair use doctrine of: •Taiwan Copyright Act Articles 52 & 65 •The National Academies Terms of Use •The "Code of Best Practices in Fair Use for OpenCourseWare 2009 (http://www.centerforsocialmedia.org/sites/default/files/10-305-OCW-Oct29.pdf)" by A Committee of Practitioners of OpenCourseWare in the U.S. The contents are based on Section 107 of the 1976 U.S. Copyright Act P36 V. T. J. Davies, T. G. Barraclough, M. W. Chase, P. S. Soltis, D. E. Soltis, and V. Savolainen 2004. Darwin's abominable mystery: Insights from a supertree of the angiosperms. PNAS 101: p. 1908. http://www.pnas.org/content/101/7/1904.full.pdf+html It is used subject to the fair use doctrine of: •Taiwan Copyright Act Articles 52 & 65 •The National Academies Terms of Use •The "Code of Best Practices in Fair Use for OpenCourseWare 2009 (http://www.centerforsocialmedia.org/sites/default/files/10-305-OCW-Oct29.pdf)" by A Committee of Practitioners of OpenCourseWare in the U.S. The contents are based on Section 107 of the 1976 U.S. Copyright Act P37 P39 National Taiwan University Jer-Ming Hu 66 Work Licensing Author/Source Page D. T. Pride, R. J. Meinersmann, T. M. Wassenaar, and M. J. Blaser 2003. Evolutionary Implications of Microbial Genome Tetranucleotide Frequency Biases. Genome Research 13: p. 152. http://genome.cshlp.org/content/13/2/145.full.pdf+html 2012/06/14 visited P41 D. T. Pride, R. J. Meinersmann, T. M. Wassenaar, and M. J. Blaser 2001. Genome trees constructed using five different approaches suggest new major bacterial clades. 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