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Lecture 20 The origin of life 1. It must be capable of reproducing. 2. It must possess a genotype and a phenotype. 3. It must possess a metabolism. 4. It must be capable of evolving. Some problems in addressing the origin of life on earth Some problems in addressing the origin of life on earth 1. Narrow time window Some problems in addressing the origin of life on earth 1. Narrow time window • the age of the earth is 4.55 billion years old. Some problems in addressing the origin of life on earth 1. Narrow time window • the age of the earth is 4.55 billion years old. • life first appeared about 3.8 billion years ago. Some problems in addressing the origin of life on earth 1. Narrow time window • the age of the earth is 4.55 billion years old. • life first appeared about 3.8 billion years ago. • the planet could not have sustained life for its first 500 million years! Some problems in addressing the origin of life on earth 1. Narrow time window • planet could not have sustained life for its first 500 million years! Life on Earth - Timescale Precambrian (4,600 to 543 MYA) (MYA = million years ago) - life on earth evolved about 3,800 MYA. - prokaryotes evolved about 3,500 MYA - eukaryotes evolved about 2,100 MYA - multicellular eukaryotes appeared about 1,500 MYA. Life on Earth - Timescale Theories for the evolution of life Extraterrestrial origin of life Panspermia Svante August Arrhenius (1907) Theories for the evolution of life The chemical theory (Oparin-Haldane theory) A.I. Oparin 1894 - 1980 J.B.S. Haldane 1892 - 1964 Theories for the evolution of life The chemical theory (Oparin-Haldane theory) inorganic molecules ↓ organic molecules ↓ biological polymers ↓ replicating systems ↓ protobionts ↓ true cells Theories for the evolution of life The chemical theory (Oparin-Haldane theory) inorganic molecules ↓ organic molecules ↓ biological polymers ↓ replicating systems ↓ protobionts ↓ true cells First macromolecules Extraterrestrial origins F. Crick 1916- 2004 Step 1. Inorganic molecules → organic molecules 1. Extraterrestrial evidence • the Murchison meteorite contained over 70 amino acids! Step 1. Inorganic molecules → organic molecules 1. Extraterrestrial evidence • the Murchison meteorite contained over 70 amino acids! • equal mixture of D and L isomers present. First macromolecules 1953, classic experiment by Stanley Miller and Harold Urey (Production of some organic compounds under possible primitive Earth conditions) 1. gases representing the primitive atmosphere 2. electrical discharge using tungsten electrodes. Results after a few days: aldehydes, carboxylic acid and amino acids First macromolecules 1953, classic experiment by Stanley Miller and Harold Urey (Production of some organic compounds under possible primitive Earth conditions) 1. gases representing the primitive atmosphere 2. electrical discharge using tungsten electrodes. Results after a few days: aldehydes, carboxylic acid and amino acids S. Miller 1930- 2007 Theories for the evolution of life The chemical theory (Oparin-Haldane theory) inorganic molecules ↓ organic molecules ↓ biological polymers ↓ replicating systems ↓ protobionts ↓ true cells First polynucleotides clay substrates Theories for the evolution of life The chemical theory (Oparin-Haldane theory) inorganic molecules ↓ organic molecules ↓ biological polymers ↓ replicating systems ↓ protobionts ↓ true cells First polynucleotides clay substrates Ribozymes Ribozyme from Tetrahymena thermophila Evidence for an early role for RNA Evidence for an early role for RNA RNA is involved in: Evidence for an early role for RNA RNA is involved in: 1. DNA replication. Evidence for an early role for RNA RNA is involved in: 1. DNA replication. 2. Protein synthesis. Evidence for an early role for RNA RNA is involved in: 1. DNA replication. 2. Protein synthesis. 3. Ribonucleoside triphosphates (ATP, GTP) are the energy currency of cells. Evidence for an early role for RNA RNA is involved in: 1. DNA replication. 2. Protein synthesis. 3. Ribonucleoside triphosphates (ATP, GTP) are the energy currency of cells. 4. Deoxyribonucleotides are synthesized from RNA precursors. Theories for the evolution of life The chemical theory (Oparin-Haldane theory) inorganic molecules ↓ organic molecules ↓ biological polymers ↓ replicating systems ↓ protobionts ↓ true cells Step 4. Replicating systems → protobionts Step 4. Replicating systems → protobionts Replicating systems → protobionts • heating and cooling mixtures of amino acids can form spherical proteinoids. Replicating systems → protobionts • heating and cooling mixtures of amino acids can form spherical proteinoids. • mixtures of lipids and proteins can form liposomes. Replicating systems → protobionts • heating and cooling mixtures of amino acids can form spherical proteinoids. • mixtures of lipids and proteins can form liposomes. • liposomes can “reproduce” by budding off smaller units. Replicating systems → protobionts • heating and cooling mixtures of amino acids can form spherical proteinoids. • mixtures of lipids and proteins can form liposomes. • liposomes can “reproduce” by budding off smaller units. • if an RNA-protein based metabolism evolved, natural selection can again occur. Theories for the evolution of life The chemical theory (Oparin-Haldane theory) inorganic molecules ↓ organic molecules ↓ biological polymers ↓ replicating systems ↓ protobionts ↓ true cells Protobionts → true cells Protobionts → true cells • natural selection would have acted to biochemical sophistication of protobionts. Protobionts → true cells • natural selection would have acted to biochemical sophistication of protobionts. • DNA became the “repository” of the genetic information. Protobionts → true cells • natural selection would have acted to biochemical sophistication of protobionts. • DNA became the “repository” of the genetic information. Why? Protobionts → true cells • natural selection would have acted to biochemical sophistication of protobionts. • DNA became the “repository” of the genetic information. Why? 1. DNA is more stable than RNA. Protobionts → true cells • natural selection would have acted to biochemical sophistication of protobionts. • DNA became the “repository” of the genetic information. Why? 1. DNA is more stable than RNA. 2. RNA freed to act only in one arena (catalysis). What is the evidence for prokaryotic life 3.5 to 4.0 BYA? What is the evidence for prokaryotic life 3.5 to 4.0 BYA? 1. Stromatolites Shark Bay, Western Australia Stromatolites What is the evidence for prokaryotic life 3.5 to 4.0 BYA? 1. Stromatolites What is the evidence for prokaryotic life 3.5 to 4.0 BYA? 1. Stromatolites • are bun-shaped structures made by cyanobacteria. What is the evidence for prokaryotic life 3.5 to 4.0 BYA? 1. Stromatolites • are bun-shaped structures made by cyanobacteria. • fossil stromatolites are abundant 3.5 bya. What is the evidence for prokaryotic life 3.5 to 4.0 BYA? 1. Stromatolites • are bun-shaped structures made by cyanobacteria. • fossil stromatolites are abundant 3.5 bya. • recently “re-discovered” in shallow, hypersaline environments (e.g., Hamelin pool in Shark Bay, western Australia). Stromatolites Stromatolites are the oldest known fossils, more than 3 billion years. They are colonial structures formed by photosynthesizing cyanobacteria (blue green algae) and other microbes. Cyanobacteria were likely responsible for the creation of earth's oxygen atmosphere. They were the dominant lifeform on Earth for over 2 billion years. Today they are nearly extinct, living a precarious existence in only a few localities worldwide. What is the evidence for prokaryotic life 3.5 to 4.0 BYA? 2. C isotope ratios suggest early photosynthesis What is the evidence for prokaryotic life 3.5 to 4.0 BYA? 2. C isotope ratios suggest early photosynthesis • 12CO2 is preferentially fixed in photosynthesis than the heavier 13CO2. What is the evidence for prokaryotic life 3.5 to 4.0 BYA? 2. C isotope ratios suggest early photosynthesis • 12CO2 is preferentially fixed in photosynthesis than the heavier 13CO2. • isotopic ratios of 13C to 12C indicate that autotrophic bacteria fixing carbon via the Calvin cycle by 3.5 bya. What is the evidence for prokaryotic life 3.5 to 4.0 BYA? 2. C isotope ratios suggest early photosynthesis • 12CO2 is preferentially fixed in photosynthesis than the heavier 13CO2. • isotopic ratios of 13C to 12C indicate that autotrophic bacteria fixing carbon via the Calvin cycle by 3.5 bya. • where did the O2 go? Into the oceans to form banded ironstone sediments. The origin and early evolution of the eukaryotes The origin and early evolution of the eukaryotes 1. The universal gene-exchange pool hypothesis The origin and early evolution of the eukaryotes 1. The universal gene-exchange pool hypothesis • Archaea, Bacteria, and Eucarya evolved only after extensive lateral gene transfer ceased. The universal gene-exchange pool hypothesis The origin and early evolution of the eukaryotes 1. The universal gene-exchange pool hypothesis • Archaea, Bacteria, and Eucarya evolved only after extensive lateral gene transfer ceased. 2. The ring of life hypothesis The origin and early evolution of the eukaryotes 1. The universal gene-exchange pool hypothesis • Archaea, Bacteria, and Eucarya evolved only after extensive lateral gene transfer ceased. 2. The ring of life hypothesis • Eucarya evolved from a fusion of a bacterium and an archean. The ring of life hypothesis Endosymbiotic hypothesis Lynn Margulis 1938-2011 Endosymbiotic hypothesis Respiration - First use of Oxygen Multicellular organisms