Origin and Evolution of Life on Earth Bennett et al. Chapter 5

1. Origin and Evolution of Life on EarthBennett et al. Chapter 5 HNRS 228 Astrobiology w/Prof. Geller (slides by Taylor and Geller) Lectures 9 and 10

2. Origin and Evolution of Life on Earth - Chapter 5 Overview • Searching for the origin • Functional beginnings of life • From chemistry to biology at the molecular level • Prokaryotes and oxygen • Eukaryotes and explosion of diversity • Mass extinctions, asteroids and climate change • Evolutions of humans • Conclusions

3. But FirstRemember Morowitz • Order from Chaos vs. Order from Order • Origins Questions • origin of universe • origin of solar system • origin of life • origin of mind • Stars and the nucleosynthesis of heavy elements • Pauli Exclusion Exclusion • importance in bonding

4. Morowitz II • Emergence of chemistry from physics • underlying simplicity • Amphiphile • An amphiphile is a molecule that has a polar head attached to a long hydrocarbon tail. The result is that one part of the molecule (the polar head) interacts strongly with water, while the other part of the molecule (the long hydrocarbon tail) interacts strongly with nonaqueous phases or with the hydrocarbon tails of neighboring same-species molecules via dispersion forces (van der Waals forces). • All metabolism => 500/600 molecules • 1850 catalog of organics by Beilstein • Computational techniques • in search of an algorithm for distilling the 500/600 molecules of life out of millions of organics (a type of Pauli principle)

5. Morowitz III • Citric acid cycle • reductive vs. oxidative

6. Morowitz IV • Autotrophs vs. heterotrophs • autotrophs first • “Impossible by random interactions” • Glutamic Acid • generation without enzymes • Emergence Principle • “Hell and Heaven Theory” of origins • Occam’s Razor

7. Morowitz V • Calvin-Benson Cycle • glucose biosynthesis

8. Taylor Hierarchy of Life Community Population Organism Organ Tissue Cell Organelles Macromolecules Atoms

9. Taylor Hierarchy of Ecology Earth Biosphere Biome Ecosystem Community Population Organisms

10. Primordium Large-scale structure Stars Elements Solar systems Planets Geosphere Biosphere Prokaryotes Eukaryotes Multicellularity Neurons Protostomia/Deuterostomia Chordates & Vertebrates Fish Amphibians Reptiles Mammals Arboreal mammals Primates Great apes Hominids Toolmakers Language Agriculture Technology/urbanization Philosophy Spiritual Morowitz Emergences

11. Searching for the Origin • What is the question? • The tools and methodologies • Principles of physics (e.g., 1st and 2nd Law of TD) • Principles of geology (e.g., relative/absolute dating) • Principles of chemistry (e.g., chemistry of water) • Principles of biology (e.g., key macromolecules) • Occam’s razor • Conclusions: plausible scenario of the events and processes that lead to the origin of life

12. Searching for the Origin • When did life begin? • Evidence • Widespread life forms (3.5 B years ago) • Stromatolites (3.5 B years ago) • Fossilized cells (3.5 B years ago) • Radiometric dating: carbon isotopes (3.85 B years ago) • Carbon 12 versus Carbon 13 • Range of dates: 4.1 to 3.85 B years ago • Conclusions • Life arose late in the Hadean Eon • Life colonized planet in very short time frame (< 500 M years)

13. Searching for the Origin: Comparative Genomics • Comparative morphology versus comparative genomics • “Living Fossils” of DNA and RNA • Sequence of nucleotides in DNA and genome • Pattern and process of change in sequences • Comparing sequences reveals a pattern/order • Methodology of comparison – rRNA (ribosomal RNA)

14. Searching for the Origin: Three Branches of Life Forms • Results from comparative genomics • Three major domains • Bacteria • Archaea • Eukarya • Common ancestor analysis • Comparison to organisms today • Deep sea volcanic vents • Thermophiles (hyperthermophiles) • Comparison to environment of Hadean Eon

15. Searching for the Origin Domain Domain Domain Bacteria Archaea Eukarya Common Ancestor

16. Searching for the Origin: At What Place on Earth? • Options • Continental landscapes • Shallow pools • Hot springs • Deep sea vents • Data to support one or the other • Comparative genomics • Chemical energy (hydrogen sulfide) FeS + H2S FeS2 +H2 + Free Energy • Conclusion: deep sea vents • Probability of bombardment

17. Beginnings of Life on Earth • Organic chemistry* • Transition from chemistry to biology • Panspermia • The evolution of sophisticated features of metabolism and information brokers • Conclusions _________ * Enzymes first or TCA or ?

18. Catalysis in Living Systems: Enzymes • Introduction • Most reactions are “very slow” (insufficient to sustain life) • Mechanisms to accelerate specific reactions: preferential acceleration • Evolutionary significance: positive fitness • Accelerants => Catalysts => Enzymes • Proteins (relate to information brokers) • Change rate of reactions • High degree of specificity • Regenerated (not consumed)

19. Enzymes: How They Work • Base case for reactions to occur • Reactants (A-B and C-D) • Products (A-D and B-C) • Energy analysis • Free energy (net change in energy between reactants and products; DG) • Exothermic reactions (release energy/spontaneous) • Endothermic reactions (require energy to proceed) • Energy of activation (transition state or “hill”) • EA in diagram

20. Catalysis in Living Systems: Enzymes Transition State EA Free Energy %G A-B C-D %G A-D B-C

21. Catalysis in Living Systems: Enzymes • Efficacy of enzymes • “Hill”/transition state is the key • Mechanism • Lower energy of activation (EA) needed to surmount the “hill) • Selectivity/specificity (lock and key analogy) • Active site of protein due to 3-D conformation • Regeneration • Conclusion • Absence of enzymes: minutes to hours for reaction • Presence of enzymes: 1,000 times per second

22. Catalysis in Living Systems: Enzymes Transition State EA Free Energy %G A-B C-D %G A-D B-C

23. Enzymes: Modification of the Rate of Reaction • Factors controlling efficacy of enzymes • Concentration of enzyme • Concentration of reactants • Factors affecting proteins (enzymes) • Temperature (compare humans versus thermophiles in hot springs) • pH • Mechanism (3-D conformation of proteins and lock & key analogy) • Inhibitors

24. Evolutionary Perspective of Enzymes • Evolutionary advantage of enzymes • Specific acceleration of reactions • Fitness value: positive • Information broker: coded in the DNA • Mutation • Reproduction • How did enzymes come to be?

25. Ribozymes • What are ribozymes in current biochemistry? • NOT ribosomes • mRNA (small fragments) • Functions • Synthesis of RNA, membranes, amino acids, ribosomes • Properties • Catalytic behavior (enhance rates ~20 times) • Genetically programmed • Naturally occurring (60-90 bases)

26. Ribozymes (continued) • Laboratory studies of ribozymes • Creation of RNA fragments at random with existence of enzyme-like properties • Variety of enzyme-like properties • Cleavage of DNA • Cleave of DNA-RNA hybrids • Linking together fragments of DNA • Linking together fragments of RNA • Transformation of polypeptides to proteins • Self-replication (2001)

27. Summary of Ribozymes • mRNA fragments • 3-D conformation like proteins (e.g., fold) • Functional ribozymes created at random in test tube • Exhibit catalytic behavior • Self replicate • Play a prominent/key role in any scenario for understanding the evolution of life at the biochemical and molecular level

28. RNA World

29. Chemical Beginnings

30. Urey-Miller Experiment

31. Significance of and Sequel to Urey Miller Experiment • Multiple variations of the study (e.g., atmosphere) • 20+ amino acids, sugars, bases for DNA and RNA, ATP, etc. • Significance: scenario for the abiotic formation of key carbon polymers (macromolecules) • Probable environments • Deep sea vents • Tidal pools (role of repeated evaporation and concentration – “evapoconcentration”; asteroid bombardment) • Chemical events leading to an “RNA World”

32. Panspermia

33. Functional Beginnings of Life: Transition from Chemistry to Biology • Ribozymes • Enzyme activity • Self replicating • Generation of biomacromolecules (C polymers; e.g., sugars, nucleotides, ATP) • via abiotic processes on Earth (Urey-Miller) • via Panspermia • via biotic processes (e.g., ribozymes) • Role of mutations, natural selection and environment: incremental changes in biomacromolecules that are inherited via RNA and DNA)

34. Functional Beginnings of Life: Transition from Chemistry to Biology Glycolysis/Fermentation 6 Carbon Sugar/ Glucose ATP (2) 3 Carbon Sugar/ Pyruvate CO2 2 Carbon Sugar/ Acetyl Coenzyme A

35. Functional Beginnings of Life: Transition from Chemistry to Biology • Evolution of Photosynthesis CO2 + H2O + Light = CH2O + O2 • Key processes • Absorption of light (pigments) • Conversion of light energy into chemical energy (ATP) • Synthesis of simple carbon compounds for storage of energy • Purple bacteria and Cyanobacteria • Primitive forms (~3.5 BYA)

36. Prokaryotes and Oxygen % of Present 4.8 4 3 2 1 0.7 0.1 0 Billions of Years Before Present

37. Functional Beginnings of Life: Transition from Chemistry to Biology Glycolysis to Respiration 2 Carbon Glycolysis/ Fermentation 4 Carbon 6 Carbon Aerobic Metabolism (TCA) ATP 5 Carbon ATP

38. Prokaryotes and Oxygen • Evolution of Photosynthesis CO2 + H2O + Energy = CH2O + O2 • Evolution of respiration CH2O + O2 = CO2 + H2O + Energy • Possibility that respiration is simply the reverse of photosynthesis • Oxygen crisis and the oxygen stimulation to evolution

39. Eukaryotes and an Explosion of Diversity • Incremental changes in evolution: role of oxygen and diversification of organisms (explain ATP fitness) • Quantum changes in evolution • Symbiosis • Lynn Margulis theory: eukaryotes are derived from prokaryotes • Compartmentalization and organelles • Bacterial origins of chloroplast and mitochondria

40. Mass Extinctions, Asteroids and Climate Change • Mass extinctions • Dramatic declines in a variety of species, families and phyla (>25%) • Timing of decline is concurrent • Rate of decline is precipitous (geological sense) • Example of catastrophism • Best example • Cretaceous/Tertiary boundary (65 M years ago) • K-T boundary and Alvarez theory of catastrophism

41. Mass Extinctions, Asteroids and Climate Change: K-T Boundary • Observations • Iridium deposits in distinct layers: suggestion of an asteroid (10-15 Km) • Other trace elements (characteristics of asteroids) • Shocked quartz • Soot deposits • Conclusive Evidence • Impact crater 200 km off Yucatan Peninsula (Chicxulub Crater)

42. Mass Extinctions, Asteroids and Climate Change: Other examples • Other mass extinctions • Five major extinctions over last 600 M years • Evidence for gradualism • First principles: evolution • Pattern in the data • Recovery response • Overall increment in number of families over geological time • Conclusions: Catastrophism coupled with gradualism

43. Origin and Evolution of Life on Earth: Conclusions • Plausible scenarios for the early origin of life on Earth (abiotic and biotic) • Role of mutation and evolution in origin of increasingly more complex forms of metabolism • Role of major evolutionary and climatological events as “pulses” of diversification in biota