ASTR/GEOL-2040: Search for life in the Universe: Lecture 7 • Metabolism first vs replication first • RNS world & LUCA On Thursday (Feb 9) • Lecture 3:30 – 4:10 – Giving out HW3, review, Q/A • Quiz 4:15 – 4:45 • Special accommodations – I have emailed everybody if you contacted me – Not everybody contacted me, so make sure you do 2 Today • • • • Metabolism first? RNA world LUCA Reading: – RGS pp. 30-40 – Lon pp. 193-195 – BS pp. 172-176, 206-208 3 Primitive cells • Lipid bilayers • Protein droplets (Oparin 1924) • Dehydration – rehydration (S Fox 1958) 4 Properties of protocells • Confinement of organics within cells is advantageous: – Facilitates chemical reactions. – Cooperative relationships evolve. • Membrane-like spheres easily made in lab experiments! – Cooled amino acids solutions. – Lipids in water. • First “cell” may have been RNA replicating within simple membrane. Role of minerals • Support – Amino acids polymerize on surfaces • Selection – Different crystal faces select left/right – Both possible natural selection chose one • Catalysis – N2 to N3H via metalic surfaces – Suitable in hydrothermal vents 6 Minerals as Templates? • How could first RNA “genetic” strands arise? • Lab experiments: clays + water + organics complex organics, including RNA strands. • Repeating chemical structure of minerals may facilitate assembly of complex organics! clay (microscopic image) pyrite Role of minerals • Support – Amino acids polymerize on surfaces • Selection – Different crystal faces select left/right – Both possible natural selection chose one • Catalysis – N2 to N3H via metalic surfaces 8 Metabolism • How to make a living (Longstaff 193) • Use of catalysts – Speeds up reaction – Regardless of direction (!) • Two types – Proteins – RNA catalysts (=ribozyme) 9 Three requirements • Source of carbon (CO2 or CH2O) • Source of energy – To reduce inorganic to org macromolecule – Electron donor (e.g. H2) • An oxidant – To harness chemical potential energy – Electron acceptor (e.g. O2) 10 “Food” in Greek? RGS 36, Lon 195, BS 174 11 Troph (Greek) = food • auto – hetero • photo – chemo • litho – organo • • • • Photoautotroph Chemoautotroph Photoheterotroph Chemoheterotroph RGS 36, Lon 195, BS 174 12 Range of possibilities • e.g.: Chemolithoheterotroph • Altogether 8 possibilities! 13 Thiobacillus denitrificans • Discovered 1904 – 0.5x1x3 mm3 • Soil & mud – Oxidize U(IV)U(VI) • Chemolithoautotroph or chemoautotroph – H2S+CO2CH2O+2S 14 Gray bacterium in rock spaces Always found to be growing Excreting CO2 Rocks mineral structure depleted in Fe A. B. C. D. Chemoautotroph Lithoautotroph Photoautotroph Lithoheterotroph 15 Blue-green in petri dish Cells grow when exposed to sunlight Excrete O2 Grow and produce O2 as long as in sunlight A. B. C. D. Chemoautotroph Lithoautotroph Photoautotroph Lithoheterotroph 16 Metabolism/replication first? • Organism needs 2 things – Replication (otherwise not self-sustaining) • Turn disorder to ordered chem reactions to extract energy from surroundings metabolism, needed to control flow of energy 17 DNA transcription • DNA messenger RNA (mRNA, transient) • mRNA read out by ribosome (rRNA) – Ribosomes contain their own type of RNA – Amino acids + RNA (tRNA, small) • Ribosome synthesizes proteins (incoming tRNA) – Forges peptide bonds between amino acids – tRNA liberated, captures new amino acids – 10-20 amino acids/second 18 Walter Gilbert (1986, Nature 319, 618) The RNA world • Central dogma of chemistry of life – DNA RNA protein store messenger enzyme • RNA world: only RNA – RNA acts as enzyme, messenger, and store for itself! – Ribonucleic acid enzyme = Ribozyme (mRNA, etc) 19 RNA itself acts as enzyme – in addition to proteins! 20 RNA itself as enzyme 21 RNA world before DNA/protein • • • • Nucleotides in RNA easier made RNA evolved to DNA (greater stability) No scenario for protein replication w/o RNA Natural selection outcompeted DNA+protein 22 RNA world evolution • RNA might produce small proteins (polypeptides), which aid RNA replication! • Best-replicating RNA “survive” and dominate. • Mutations promote RNA “genetic” variety. • RNA role in making proteins is consistent with an early RNA world. • More efficient DNA evolved later. Similarity with viruses • Replicates only inside cell (all domains of life) – RNA or DNA + protein (+lipid) coat • Edge of life: have genes, evol nat selection – – – – – But no metabolism 20-300 nm size Double-stranded: dsDNA, dsRNA Single-stranded: ssDNA, ssRNA retrovirus High mutation rate 24 Evidence for RNA world • • • • Ribozymes have been synthesized E.g. polymerase Catalyzed synthesis of 95 nucleotides Itself able to manufacture another ribosome 25 Advantage of RNA over DNA? A. Simpler, easier to synthesize? B. Less stable? C. Melting temperature lower? D. All of the above. E. Both A and B. 26 Phylogenetic Tree • Based on physical & genetic differences – Taxa: group of populations • Darwin (1857): the time would come “when we shall have very fairly true genealogical trees of each great kingdom of nature.” – RGS: life does not reject what evolution has created, but simply builds on what has gone before. 27 Phylogenetic Tree • RGS: life does not reject what evolution has created, but simply builds on what has gone before. – Biological record of continuous additions/modifications in genetic material • Tree construction: – Similar molecules in different creatures – Similar parts inherited from common ancestor 28 Position in classification scheme bacteria bacteria deinococcus-thermus proteobacteria deinococcales enterobacteriales deinococcaaceae enterobacteriaceae deinococcus escherichia radiodurans coli 29 Modern tree building • Uses ribosomal RNA (rRNA) – Branch length = difference in rRNA • Three clear “domains” – Bacteria, archea, eukarya • Root on tree LUCA – Last universal common ancestor • Closest to tree: thermophiles, hyperthermophiles – Evolution from hot cold – chemotrophs 30 Rooted Tree Unrooted Tree Modern tree building • Uses ribosomal RNA (rRNA) – Branch length = difference in rRNA • Three clear “domains” – Bacteria, archea, eukarya • Root on tree LUCA – Last universal common ancestor • What types of organisms closest to the root? 33 Modern tree building 34 Thermus, thermofilum, methanopyrus,... Deinococcus thermus 35 Closest to the root? • ……….. • ………………. 36 Closest to the root? • thermophiles, hyperthermophiles – Evolution from hot cold • Chemotrophs – Deep biosphere rather than surface 37 Cold organisms first? • • • • ….. ……… ……………. ………………… 38 Cold organisms first? • • • • Yes, also possible: Hydrothermal vent first Then ocean-sterilizing impact Only thermophiles survives 39 Origin of Life: Summary 1. 2. 3. 4. 5. “Organic soup” vs. dilute solution. Complex organics developed (mineral templates?). “Pre-cells” enclosed complex organics. Natural selection increased RNA complexity. DNA developed within some successful cell(s). A reasonable scenario, though many details are missing! On Thursday (Feb 9) • Lecture 3:30 – 4:10 – Giving out HW3, review, Q/A • Quiz 4:15 – 4:45 • Special accommodations – I have emailed everybody if you contacted me – Not everybody contacted me, so make sure you do 41 Next • • • • Summary Habitable zone Origin of life on Earth Chapter 2 of RGS pp 43-59
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