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Teaching about time a biologists perspective Biochemistry Physiology Ecology Evolution Origins of biodiversity and estimates of divergence times 10-6s TIME 109 years Integrative Courses Vertebrate or Invertebrate Zoolgy Comparative Anatomy Biogeography *not drawn to scale Biologist’s general modus operandus • We had no training on how to teach about deep time other than memorizing the chart • We focus on teaching biological processes, not so much the time scales during which they proceed • We make modest attempts to talk about deep time (give out handouts) • We give our geology/paleontology colleagues absolute authority to provide dates of events (with no discussion of error, methods, etc.) • We do not assess whether students grasp time concepts within evolution • We avoid confrontations with students who challenge us with biblical timelines of events QuickTime™ and a decompressor are needed to see this picture. QuickTime™ and QuickTime™ and a aa QuickTime™ and decompressor decompressor decompressor are see this picture. are needed see this picture. areneeded neededtototo see this picture. Teaching deep time through macroevolution and phylogenetics QuickTime™ and a decompressor are needed to see this picture. Stories of evolution through phylogenies of many topologies QuickTime™ and a decompressor are needed to see this picture. QuickTime™ and a decompressor are needed to see this picture. QuickTime™ and a decompressor are needed to see this picture. QuickTime™ and a decompressor are needed to see this picture. Biomedcentral.com Genetic diversity/no morphological diversity Plethodontid salamanders 3.1 mya (Pliocene) mt DNA sequence P. hubrichti RM 1 N1 RM 2 BM 1 BM 2 DG VA 1 GF 1 N2 GF 2 WT VA1 DG VA 2 WT VA 2 SI 1 SI 2 S1 CW 1 CW 2 RBB 1 RBB 2 BR 1 CM 1 CM 2 BR 2 S2 PG 1 SM. 1 SM. 2 PG 2 ML 1 ML 2 CD 1 S3 CD 2 Desmognathus wrighti N1 (pygmy salamander) N2 S3 S2 S1 Combination mtDNA and allozyme Genetic variation among Indian groups of peoples QuickTime™ and a decompressor are needed to see this picture. BMC Genetics 2004, 5:23 Phylogeography of Drosophila in Hawaii (Bromham and Penny, 2003) QuickTime™ and a decompressor are needed to see this picture. Molecular clocks Zuckerkandl and Pauling 1962 • Implicitly used when choosing a region to assay for variation given the expected evolutionary distance of interest • Explicitly used when attempting to date divergence times • Need to calibrate divergence times estimated with DNA variation with historical geological dates/events • Lots of debate and criticism about the use of molecular clocks QuickTime™ and a decompressor are needed to see this picture. Inquiry-based integrated instructional unit: Phylogenetic analysis of the bony fishes: Morphological and mtDNA sequence 3-4 week module Comparative Anatomy Evolution Genetics Molecular Biology Ichthyology, etc Introductory Biology What are the phylogenetic relationships among orders of Actinopterygii? 150 mya 200 mya 250 mya 300 mya Learning activities 1. Students assign traits to a set of sea shells and generate a “tree” based on the similarity of characters--this is done by hand. [cladograms are visual representations of calculated relationships] 2. Students create character matrix and extract DNA/ sequence the 28s rRNA gene. [scaled up repetition, base pair differences are empirically determined] 3. Students align sequences, calculate the number of differences among taxa and use a computer program to generate a phylogenetic tree [more complex iteration of initial exercise; introduces bioinformatics] 4. Students download cytb sequence and generate another tree for the same taxa [repeat tree making skills] Assessment Exam questions: “What is a molecular clock, and how is it relevant to phylogenetic analysis?” “Compare the cytb tree to the tree you constructed from the 28S rRNA gene you made earlier in the week. Which gave a more robust hypothesis (and why), and what are the reasons why the two genes resolved different hypotheses?” ….but that’s it. Goals of this workshop (for me) 1. Learn more formal pedagogy behind teaching deep time 2. Learn how to integrate cognitive science in pedagogy of phylogenetics • Analogy • Diagram reasoning (“tree thinking” Catley and Novick,2009) 3. Form collaborations with geoscientists to generate a working group interested in integrating teaching methods of deep time from geoscience and evolutionary biology perspectives. Molecular clocks When is a molecule not appropriate? Questions to ask yourself Do molecular clocks tick evenly through time? Is there a geological or climatic date/event for calibration? Are geological calibrations accurate? Molecules can evolve at different rates than organisms (or other molecules)! Is the clock ticking at different rates among lineages? Deuterostome phylogeny: Morphology Morphological Analysis no 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 no MTR 1 WHI 2 BUT 3 TRO 4 Body form 1 lateral compression 0 lat assym body 0 Head shape 2 # dorsal fins 0 dorsal fin spike 0 hard fin rays (Y/n) 1 dorsal ray type 1 fleshy dorsal fin 0 adipose fin 0 # anal fins 1 Caudal fin type 2 Caudal fin shape 0 pectoral fin shape 0 pectoral fin ratio 1 pelvic fin ratio 2 pelvic fin (yes/no) 0 Muscular fin 0 loc pelvic fin 1 scale type 3 loss of scales 0 scale distribution 0 scale shape 2 # nostril pairs 1 Deviated nostrils (yes/no)0 nostril position 1 eye location in skull 2 eye placement 0 jaw type 2 Jaw protrusion 2 Jaw hinge type 1 teeth 1 teeth pointed/blunt 0 >1 teeth row 0 Fleshy lips 1 mouth position 2 elongate rostrum 0 Gular plate (yes/no) 0 peduncle:tail ratio 1 Lateral line coloration 0 0 2 0 3 0 0 1 1 1 0 1 2 2 2 1 2 0 0 1 2 0 1 3 0 1 1 2 0 1 1 0 0 0 1 0 3 0 0 1 1 2 1 0 2 1 1 0 0 0 0 2 2 0 0 1 ? 1 0 ? 2 1 0 3 0 0 1 1 0 2 2 1 1 1 1 0 1 0 1 1 0 1 1 0 2 0 1 0 1 0 1 1 2 3 0 0 1 0 0 0 2 0 0 3 0 0 1 0 0 2 2 1 0 0 1 0 2 0 0 2 0 2 1 1 2 1 1 1 0 0 0 1 2 2 2 0 2 0 0 1 2 0 0 3 0 0 1 1 1 2 2 1 0 0 1 1 2 0 0 1 1 2 1 0 2 1 1 1 1 1 0 1 2 0 0 1 1 0 0 1 3 0 0 2 0 0 1 1 0 1 2 0 0 1 0 1 2 0 0 1 1 Character 2 3 4 5 6 Character 1 FLO POR LUN 5 6 7 CIC 8 SUN 9 0 2 0 1 1 1 0 0 1 0 2 2 1 2 1 0 0 1 0 3 0 1 3 0 1 1 2 0 2 1 1 1 1 0 1 2 0 0 ? 0 1 2 0 1 1 1 1 1 0 0 1 2 1 0 0 2 0 0 1 3 0 1 2 0 1 1 2 0 2 1 1 1 0 0 0 2 0 0 2 0 0 1 0 1 1 1 1 1 0 0 1 2 1 0 1 1 0 0 1 3 0 0 2 1 0 1 1 0 2 2 1 0 0 0 1 2 0 0 1 0 0 0 0 2 0 0 1 1 1 0 1 2 0 0 0 2 0 0 1 ? 1 1 3 0 1 1 1 0 1 1 0 0 0 1 0 2 0 0 0 1 1 2 0 1 1 1 0 1 0 0 1 1 1 1 1 1 0 0 0 2 0 1 2 0 1 1 2 0 2 2 1 0 0 0 0 2 0 1 2 0 7 8 9 10 11 MAC BOW GAR 10 11 12 GOL 13 SHK 14 0 2 0 1 1 1 1 1 0 0 1 1 1 0 1 1 0 0 1 1 0 0 1 0 0 1 0 0 2 1 1 0 0 1 0 3 0 1 1 0 2 0 0 2 1 1 1 0 0 0 1 2 0 0 0 1 0 0 0 2 0 0 2 0 1 1 1 0 2 2 1 1 2 0 1 2 1 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 12 13 14 Genomic/mt DNA extraction PCR target gene sequence DNA sequence Alignment gi|38154450|gb|AY452491.1| gi|1144505|gb|U34341.1|OMU3434 gi|1144500|gb|U34340.1|ABU3434 GTGAGCTCTCGCTGGCCCTTGAAAATCCGGGGGAGAAGGTGTAAATCTCG GTGAGCTCTCGCTGGCCCTTGAAAATCCGGGGGAGAGGGTGTAAATCTCG GTGAGCTCTCGCTGGTCCTTGAAAATCCGGGGGAGAAGGTGTAAATCTCG *************** ******************** ************* gi|38154450|gb|AY452491.1| gi|1144505|gb|U34341.1|OMU3434 gi|1144500|gb|U34340.1|ABU3434 CGCCAGGCCGTACCCATATCCGCAGCAGGTCTCCAAGGTGAACAGCCTCT CGCCGGGCCGTACCCATATCCGCAGCAGGTCTCCAAGGTGAACAGCCTCT CGCCGGGCCGTACCCATATCCGCAGCAGGTCTCCAAGGTGAACAGCCTCT **** ********************************************* REV PRIMER GGCATGTTAGATCAAGGTAGATAAGGGAAGTCGGCAAATCAGATCCGTAA GGCATGTTAGAACAATGTATGTAAGGGAAGTCGGCAAGTCAGATCCGTAA GGCATGTTAGAACAATGTAGGTAAGGGAAGTCGGCAAGTCAGATCCGTAA *********** *** *** **************** ************ gi|38154450|gb|AY452491.1| gi|1144505|gb|U34341.1|OMU3434 gi|1144500|gb|U34340.1|ABU3434 Phylogenetic analysis gi|38154450|gb|AY452491.1| gi|1144505|gb|U34341.1|OMU3434 gi|1144500|gb|U34340.1|ABU3434 CTTCGGGATAAGGATTGGCTCTAAGGGCTGGGTCGGTCGGGCTGGAGTGC CTTCGGGATAAGGATTGGCTCTAAGGGCTGGGTCGGTCGGGCTGGGGTGC CTTCGGGATAAGGATTGGCTCTAAGGGCTGGGTCGGTCGGGCTGGGGTGC ********************************************* **** gi|38154450|gb|AY452491.1| gi|1144505|gb|U34341.1|OMU3434 gi|1144500|gb|U34340.1|ABU3434 GAAGCGGGGCTGGGCTCGTGCCGCGGCTGGGGGAGCAGTCGCCCCGTCGC GAAGCGGGGCTGGGCTCGAGCCGCGGCTGGGGGAGCAGTTGCTCCGCCTC GAAGCGGGGCTGGGCACGCGCCGCGGCTGGACGAG-----GCGTCGCCT*************** ** *********** *** ** ** * Variation in mutation rates among genomic regions Hartl & Clark, Principles of Population Genetics QuickTime™ and a decompressor are needed to see this picture. Biological factors that affect molecular clocks Neutral Theory Limitations: 1. Gene to gene variation in rates of mutation 2. Lineage to lineage variation in rates of mutation 3. Variation if portions of genome under selection (vs. neutral) Molecular clocks When is a molecule not appropriate? Saturation (homoplasy)