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PHYOGENY & THE Tree of life
Campbell and Reece, Chapter 26
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Phylogeny Systematics
definitions Phylogeny Systematics evolutionary history of a species or group of species discipline focused on classifying organisms & determining their evolutionary relationships
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Taxonomy how organisms are classified and named
each step called a taxon (plural: taxa)
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BINOMIAL NOMENCLATURE
Man’s Genus species: Homo sapiens used to avoid ambiguity the Latin scientific name for each individual species is the Genus species portion of taxonomy
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3 DOMAINS DOMAIN ARCHAEA Prokaryotes
many live in Earth’s extreme environments as molecularly close to eukaryotes as Domain Bacteria includes multiple kingdoms
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(notice position of domain Archaea)
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Domain Archaea methanogen thermophile
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Domain Bacteria Prokaryotic very diverse group
use every major mode of nutrition & metabolism beneficial: photoautotrophs, alcoholic fermentation, Vit K production pathologic: strep throat, flesh-eating disease, ulcers, Rheumatic fever
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Legionella pneumophilia
Domain Bacteria Gram Positive Bacteria Streptococcus (cocci) Gram Negative Bacteria Legionella pneumophilia (rods)
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Domain Bacteria Spirochetes (spirillia)
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Domain Eukarya Eukaryotic cells more complex, become specialized
able to form multicellular organisms greatest diversity
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Domain Eukarya Plants Fungi
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Domain Eukarya Animal Protozoa
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Domain Eukarya Algae Cells Algal “bloom”
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Linnean Classification
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PHYLOGENETIC TREES show the evolutionary history of a group of organisms represented by a branching diagram each branch point represents the divergence of 2 evolutionary lineages from a common ancestor
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Phylogenetic Trees Branch Point
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sister taxa basal taxa
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Phylogenetic Trees patterns of descent common ancestors
What you can learn What you cannot learn patterns of descent common ancestors does not show phenotypic similarity cannot tell ages of species based on where branches are in the “tree” sister taxa did not evolve from each other; they have a common ancestor (that could be extinct)
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Uses of Phylogenetic Tree
If “close” relatives found they could be source of beneficial alleles that could be transferred to hardier taxa via genetic engineering 2. Using DNA samples are now able to differentiate legal species from illegal species of whale, tuna
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Phylogenies are inferred from morphological & molecular data
Homology: similarity in characteristics resulting from a shared ancestry
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Homologous Chromosomes in same species
When chromosomes duplicate in S Phase of Cell Cycle see genes in same loci of each sister chromatid
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Homologous Chromosomes across Species with Common Ancestor
Genes or certain DNA sequences also homologous if they descended from sequences carried by a common ancestor
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Organisms that share very similar morphologies or DNA sequences are likely to be more closely related than organisms with vastly different structures There are examples of organisms that look very different but have very similar DNA sequences because species underwent adaptive radiation.
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Homology vs. Analogy Analogy is similarity due to convergent evolution: occurs when similar environmental pressures & natural selection produce similar (analogous) adaptations even though organisms have different ancestors.
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homoplasies: analogous structures that arose independently (Greek: to mold in same way)
Examples: bird & bat wing: their common ancestor did not fly
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The more complex the structure found in 2 species the more likely it is that they have a shared ancestor
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Molecular Evidence of Evolutionary Relationships
DNA sequence similarities have been documented among prokaryotes & eukaryotes: (comparative genomics) High degree of sequence similarity noted in some eukaryotic nuclear genes to Archaea & mitochondrial genes are similar to Bacteria
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Using DNA to map an organism’s evolutionary history
The more recently 2 species have branched from a common ancestor, the more similar their DNA sequences should be The longer ago 2 species have been on separate evolutionary paths, the more their DNA should have diverged
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Different genes evolve at different rates
Nuclear DNA Mitochondrial DNA changes slowly useful for investigating relationships between taxa that diverged hundreds of millions of yrs ago evolves rapidly useful to investigate more recent evolutionary events
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Eukaryotic genes consist of numerous coding regions (exons) that are separated by noncoding regions (introns) both transcribed into pre-mRNA and then intron sequences are removed
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in humans 90% of the exons are homologous to exons found in Drosophila & Caenorhabditis (nematode worms
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Puffer Fish is vertebrate with smallest known genome (1/7th human genome) & yet has all exons present in humans in chromosomes: “homologous” means sequences are so similar that they are not likely due to chance so are considered the result of common ancestry
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Duplication in human genome
both of genes & chromosome segments
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Based on these duplications & new combinations of exons it seems that
the vertebrate evolution has required very few new proteins evolutionary change involves making new genes by rearranging functional domains into novel combinations (called “exon shuffling”)
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Exon Shuffling Important source of genetic variation (in addition to mutations & crossing over) still investigating mechanism
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Homologous Genes 60% of human genes that encode proteins are homologous to genes from other organisms high degree of conservation of both genes & exons among widely diverse organisms from all 3 Domains is strong evidence for their common ancestry
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Nonfunctional Sequences
another bit of strong evidence for relatedness among diverse organisms is the similarity in DNA sequences that have no apparent function. one category of these are pseudogenes
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Pseudogenes 2 kinds: 1. arises from DNA replication mutations STOP codons in one of duplicates; other no mutation 2. Processed Pseudogenes: arise during transcription or translation: lack a promoter sequence so cannot be transcribed
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To date, 2909 pseudogenes in human genome
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Other functionless DNA
LINEs SINEs Long Interspersed Nucleotide Element Families: 1,2,3 Are retrotransposons Short Interspersed Nucleotide Element Also 3 families in humans Specific LINEs & SINEs found only in cloven-hooved mammals & whales
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Retroviruses RNA virus
Infects cell and turns its single strand double strand which inserts into host genome
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RetrovirusRetrotransposon
Retrovirus inserts self into host genome but somewhere along the way genes for capsids lost If LINEs in different species are homologous it is considered to be strong evidence that these 2 species share a common ancestor where that particular LINE first became established
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Review: Phylogeny can be inferred from –the fossil record,
–morphological homologies –molecular homologies
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Shared characters are used to construct phylogenetic trees
Phylogenetic trees are used to depict hypotheses about the evolutionary history of a species Shared characters are used to construct phylogenetic trees Shared ancestral characters group organisms into clades Shared derived characters distinguish clades & form branching points in the tree of life
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Shared Characteristics are used to Construct Phylogenetic Trees
Cladistics: an approach to systematics in which organisms are placed into groups based primarily on common descent Clades:groups organisms are placed in: 1 clade will include ancestor & all its descendants 3 types:
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1. Monophyletic Group equivalent to a clade
ancestral species & all its descendants
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2. Paraphyletic Group consists of an ancestral species & some of its descendants
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3. Polyphyletic Group Some members of this group will have different ancestors
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Name That Group:
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Shared Derived Character Shared Ancestral Character
character that originated in an ancestor of the taxon an evolutionary novelty unique to a clade.
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Shared Ancestral Character Shared Derived Character
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How to Build a Cladogram
dograms/Open-This-File.swf
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Making a Phylogenetic Tree
SHOULD BE POSSIBLE TO DETERMINE THE CLADE ANY SHARED DERIVED CHARACTER 1ST APPEARED Construct a CHARACTER TABLE: 1 axis has list of organisms, 1 has characters
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CHARACTER TABLE FROG IGUANA DUCK-BILLED PLATYPUS KANGAROO BEAVER
AMNION HAIR, MAMMARY GLANDS GESTATION LONG GESTATION
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Important step in cladistics is the comparison of the
Ingroup: the taxa whose phylogeny is being investigated Outgroup: the taxon that diverged before the lineage leading to the members of the ingroup Use to identify the derived characters that define the branch points in the phylogeny of the ingroup
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PHYLOGENETIC TREES WHEN CONSTRUCTING A PHYLOGENETIC TREE, SCIENTISTS USE PARSIMONY, LOOKING FOR THE SIMPLEST EXPLANATION FOR OBSERVED PHENOMENA
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SYSTEMATISTS USE MANY KINDS OF EVIDENCE, BUT EVEN THE BEST TREE REPRESENTS ONLY THE MOST LIKELY HYPOTHESIS
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Shared characters are used to construct phylogenetic trees
The phylogenetic tree of reptiles shows that crocodiians are the closest living relatives of birds
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Crocodiles & Birds share:
4-chambered heart “singing” to defend territories parental care of eggs within nests
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Phylogenetic Trees with Proportional Branch Lengths
branch lengths are proportional to the amount of genetic change in each lineage So the longer the line the more genetic changes have occurred
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Principle of Maximum Parsimony
1st investigate the simplest explanation that is consistent with the facts aka “Occum’s Razor” for phylogenies based on DNA: the most parsimonious tree requires the fewest base changes
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Principle of Maximum Likelihood
given certain probability rules about how DNA sequences change over time, a tree can be made that reflects the most likely sequence of evolutionary events
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currently using computer programs to search for trees that are parsimonious & have a high probability
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Phylogenetic Trees as Hypotheses
Scientists can make & test predictions based on the assumption that a phylogeny (the hypothesis) can be supported or not. Prediction: features shared by 2 groups of closely related organisms are also present in their common ancestor & all of its descendants
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An organism’s evolutionary history is documented in its genome: 2 Types of Homologous Genes
Orthologous Genes Paralogous Genes Exact copies found in different species Origin: common ancestor Homology result of gene duplication: multiple copies of a gene in same species all came from same gene
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Genome Evolution By investigating entire genomes of different species see 2 patterns: Lineages that diverged long ago can share orthologous genes # of genes a given species has does not seem to increase thru duplication at same rate as increase in phenotype changes
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Molecular Clocks is a yardstick for measuring the absolute time of evolutionary change based on the observation that some genes & other regions of genomes appear to evolve at constant rates. is based on assumption that the # of nucleotide substitutions in orthologous genes is proportional to the time that has elapsed since the species branched from their common ancestor (known as divergence time)
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Molecular Clocks calibration of a molecular clock is done by graphing the # of genetic differences against the dates of evolutionary branch points that are known from fossil record
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finding the average rates of genetic change from such graphs can be used to estimate dates of events that cannot be discerned from fossil record
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genes that appear to follow molecular clock are really only acting in statistically average rate of change of course parts of the genome appear to have evolved in irregular bursts some genes seem to have different rates of change in different organisms some genes evolve a million times faster than others
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Neutral Theory states much of the evolutionary change in genes & proteins has no effect on fitness & therefore is not influenced by natural selection many new genes are harmful so are quickly removed differences in the clock rate for different genes are a function of how important a gene is
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Problems with Molecular Clocks
Natural selection favors some DNA changes Many scientists remain skeptical about: the “Neutral Theory”, about using the molecular clock beyond time span documented by fossil record (about 550 million years)
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Dating the Origin of HIV with a Molecular Clock
HIV is descended from viruses that infected chimps & other primates but did not cause same AIDS-like illness When did this happen?
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HIV Strains multiple strains have infected humans
multiple strains implies multiple origins most widespread strain in HIV-M using molecular clock best guess is HIV-M strain 1st spread to humans in the 1930’s
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Is the tree of Life Really a Ring of Life?
there has been substantial movements of genes between organisms in the 3 domains mechanism: horizontal gene transfer: plasmids, viral infection, maybe even fusion of organisms
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Ring of Life some scientists think horizontal gene transfer so common that early history of life should be represented as a ring with 3 domains emerging from the ring
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