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Issues When Dealing with Polyploids. Inferring Freakish Phylogenies. Discussion Format. Assumptions of Phylogenetics Phylogenetic Incongruence. Assumptions of Phylogenetics. Phylogenetics assumes a strictly bifurcating topology involving sister taxa/lineages
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Issues When Dealing with Polyploids • Inferring Freakish Phylogenies
Discussion Format • Assumptions of Phylogenetics • Phylogenetic Incongruence
Assumptions of Phylogenetics • Phylogenetics assumes a strictly bifurcating topology involving sister taxa/lineages • How does the inclusion of polyploids affect such tree building analyses? Baum et al. 2004
Assumptions of Allopolyploids • McDade 1990 • on average, will share more derived features with its parent that has the most derived characters; therefore, placement nearest that parent • basal clade placement to the lineage that includes the most derived parent • inclusion will have limited effect on homoplasy
Incongruence • Data can tell very different stories • Morphology vs. Molecular • Molecular vs. Molecular Baum et al. 1998 Poe 1996
Sources of Incongruence • Technical • Organism-level Processes • Gene and Genome-level Processes
Morphological Continuum • Polyploidy has the potential for producing plants with morphological intermediates. Dupontia fisheri R. Br. s.l. Aiken et al. 1995 2n = 42, 44, 88, and 132
Genomes • cpDNA • Maternal vs. Paternal • nuclear DNA • Biparental http://www.gsst.kumamoto-u.ac.jp/en/org_res_act/dc_dept_mat_lif_sci.html
cpDNA Wendel and Albert 1992 61 accessions from 40 species of Gossypium resulted in 4 MP topologies differing only in the placement of G. longicalyx and G. anomalum
Gen(om)e Choice • When dealing with polyploids, which gen(om)e are you dealing with, does it possess a unique history and does it answer the question you are after? Doyle et al. 1990
Species Trees vs. Gene Trees • Species have histories, but genes can have histories of their own Sang and Zhang 1999
Polyploids at the Gene and Genome Level • Issues to Consider • Orthology vs. Paralogy • Intragenic or Interallelic Recombination • Interlocus Interactions • Subfunctionalization
Orthology vs. Paralogy A and B represent loci on the same chromosome A1 - A2 and B1 - B2 represent orthologues A1 - B2, B1 - A2, A1 - B1 and A2 - B2 all represent paralogues Inferring true history gets even stickier with polyploids...
Intragenic or Interallelic Recombination • Composite molecules are produced possessing characteristics of both parental alleles/genomes • Alleles do not arise via normal mutational processes and their relationships cannot be depicted in a hierarchical fashion • Maize Adh1 & Adh2 (Gaut and Clegg, 1993a; Hanson et al. 1996) • ArabidobsisAdh (Innan et al. 1996)
Interlocus Interactions • “exon shuffling” • “gene conversion” • “concerted evolution”
Subfunctionalization • “Silencing and relative expression levels of genes duplicated by polyploidy can be variable in different parts of the plant, indicating differential regulation of the two homoelogs during plant development.” -Adams and Wendel (2005) • eg. cotton, Tragopogon, Arabidopsis
Literature Cited • Adams, K.L. and J.F. Wendel. 2005. Novel Pattern of Gene Expression in Polyploid Plants. TRENDS in Genetics 21(10):539-543. • Baum, D.A., R.L. Small and J.F. Wendel. 1998. Biogeography and Floral Evolution of Baobobs (Adansonia, Bombacaceae) as Inferred from Multiple Data Sets. Systematic Biology 47(2):181-207. • Doyle, J.J., J.L. Doyle, J.P. Grace and A.H.D. Brown. 1990. Reproductively Isolated Polyploid Races of Glycine tacina (Leguminosae) had Different Chloroplast Genome Donors. Systematic Botany 15(2):173-181. • McDade, L.A. 1990. Hybrids and phylogenetic systematics I. Patterns of character expression in hybrids and their implications for cladistic analysis. Evolution 44:1685-1700. • Poe, S. 1996. Data Set Incongruence and the Phylogeny of Crocodilians. Systematic Biology 45(4):393-414. • Sang, T. and D. Zhang. 1999. Reconstructing Hybrid Speciation Using Sequences of Low Copy Nuclear Genes: Hybrid Origins of five Paeonia Species Based on Adh Gene Phylogenies. Systematic Botany 24(2) 148-163. • Soltis, P.S., J.J. Doyle and D.E. Soltis. 1992. Molecular Data and Polyploid Evolution in Plants. In: Molecular Systematics of Plants: pp. 177-201. Chapman and Hall: New York and London. • Wendel, J.F., and V.A. Albert. 1992. Phylogenetics of the cotton genus (Gossypium L.): character-state weighted parsimony analysis of chloroplast DNA restriction site data and its systematic and biogeographic implications. Systematic Botany 17:115-143.