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Origins “There is grandeur in this view of life, with its several powers, having been originally breathed into a few forms or into one; and that, whilst this planet has gone cycling on according to the fixed law of gravity, from so simple a beginning endless forms most beautiful and most wonderful have been, and are being, evolved.” Only figure in Origins of Species First depiction of a “tree of life” Concluding sentence in Darwin’s Origin of Species Is there a universal tree of life? Darwin’s appreciation of the scale of geologic time was key to his theory Geologists place the beginning of Earth as ~4.55 Bya Origin of the earth 4 3 2 Billions of years ago (Bya) 1 Now! Life arose in the ancient oceans ~ 4 billions years ago But where? Many possibilities…. Hydrothermal vents? Energy barriers had to be overcome Life requires a continual input of energy! Tidal pools? Clays may act as catalysts Miller-Urey experiment (1953) Objective: to determine if biomolecules could be generated from simple compounds in a mixture of gases designed to simulate a primitive reducing atmosphere Collect products, including amino acids, nucleotides, sugars, lipids Chemical origins of life http://www.pbs.org/wgbh/nova/evolution/origins-life.html What are the requirements for life? Life requires: We accept the cell as the fundamental unit of life “I took a good clear piece of cork, and with a Pen-knife sharpen’d keen as a Razor, I cut a piece of it off, and….then examining it with a Microscope, me thought I could perceive it to appear a little porous…much like a Honeycomb” Latin “cellulae” means “little rooms” from Robert Hooke’s Micrographia How did the first cell(s) form and how long did it take? Interior of the cell is separated from the environment, allowing complex process to evolve Proto-cell has an aqueous interior with RNA Surrounded by a lipid shell (membrane) that isolates it from the environment From “The Origins of Life on Earth” by Alonso Ricardo and Jack Szostak Scientific American September, 2009 RNA serves both catalytic and informational functions Ribozymes catalyze reactions Evolve to catalyze many different reactions Eventually able to selfreplicate Hypothetical sequence of events is based on experiments with synthetic ribozymes (RNA catalysts) Metabolism begins Cells take up nutrients and release waste products Ribozymes work in sequence and pathways develop Proteins appear Systems of ribozymes begin to translate strings of RNAlike nucleotides into strings of amino acids Proteins take on a variety of different catalytic and structural functions Proteins diversify eventually replace ribozymes for most catalytic reactions DNA takes on the primary informational role in cells Enzymes synthesize DNA DNA is more stable than RNA and becomes primary genetic material RNA serves as intermediates in information transfer Close to 1 billion years were required for the first life form to emerge 4.5 billion years ~billion years before life emerged 3.5-3.8 billion years prokaryotes diversify and some harvest energy from the sun Modern day example: Halobacteria are archaea that thrive in salty environments Red-purple pigments in membranes capture solar energy and convert it to ATP from “The Planet of Bacteria” by Stephen J. Gould “Our planet has always been in the "Age of Bacteria," ever since the first fossils— bacteria, of course—were entombed in rocks more than 3 billion years ago. On any possible, reasonable or fair criterion, bacteria are—and always have been— the dominant forms of life on Earth. Our failure to grasp this most evident of biological facts arises in part from the blindness of our arrogance but also, in large measure, as an effect of scale. We are so accustomed to viewing phenomena of our scale—sizes measured in feet and ages in decades—as typical of nature.” First cells appear and some become photosynthetic, but do not produce oxygen Prokaryotes will be the only life forms for the next 2 billion years Origin of the earth 4 3 2 Billions of years ago (Bya) 1 Now! Primitive cyanobacteria appear O2 is a product of photosynthesis Were they responsible for the great oxygenation event ~2.4 Bya? Layers of cyanobacteria and carbonates are apparent in both modern and ancient stromatolites Shark Bay, Australia Ancient cyanobacteria were important in the evolution of modern plants The rise in atmospheric oxygen sets the stage for eukaryotes Cells can get bigger! Cyanobacteria produce O2 photosynthesis Origin of the earth Great Oxidation Event!! First prokaryotes 4 3 2 Billions of years ago (Bya) 1 Now! Consult the homework assignment on cell size. A human skin cell is ~10 times larger than E.coli, a bacterium. Use the equation for volume to estimate how many bacteria would fit into a skin cell? (Hint: find the ratio of the volumes) volume = 4/3πr3 Neutrophil chasing a bacterium Classic movie filmed in the 1950’s by David Rogers at Vanderbilt University Note three kinds of cell: Human neutrophil Human red blood cell Staphylococcus aureus Energy considerations associated with growth The eukaryotic cell will need 1000 times more energy than E. coli. The surface area of the cell membrane must be considered. Cells take up nutrients through their membranes. Respiratory complexes in the bacterial cell membrane also generate energy. If a cell is 10 times larger than another one with the same geometry, how much more energy will it be able to make? surface area = 4πr2 Several adaptations occurred during the evolution of eukaryotes Endosymbiont hypothesis Uptake of prokaryotes capable of respiration and photosynthesis was critical to the evolution of multicellular eukaryotes Eukaryotes appear photosynthesis Origin of the earth Cyanobacteria produce O2 First prokaryotes 4 3 2 Billions of years ago (Bya) ~1 billion years to first cell additional ~1.5-2 billion years to first eukaryotic cell 1 Now! Consensus view is consistent with sequences of ribosomal RNA eukaryotic evolution involved the symbiosis of bacteria that gave rise to mitochondria and chloroplasts LUCA Changes in rDNA sequences used to predict evolutionary relationships What is the smallest number of sequence changes that can account for the data? Human cells have hundreds of mitochondria inherited from the egg Inner membrane of mitochondria contains proteins involved in energy production – many infoldings increase surface area for energy production Mitochondrial DNA (mtDNA) circular genome with 37 genes •small and large rDNA •22 tRNA genes •small number inner membrane proteins protein synthesis is more similar to prokaryotes Mitochondrial DNA: A closer look from DNA Learning Center What happened to disturb this nice tree? sequences of other genes did not produce the same tree structure both archaeal and bacterial sequences could be found in the same genome LUCA Genome projects suggest complex microbial communities at the root of the tree Data with individual genes varies Lateral transfer of DNA has occurred many times, particularly in prokaryotes LUCA is replaced with a community of cells