• • • • • Molly Hunter 641C Marley 621-9350 [email protected] Will have office hours Wed. Feb 1, Fri. Feb 3 (next week) • Or make an appt. by email or phone • Lecture style: questions, questions • Key concepts/outline posted before lecture , ppt posted after lecture Prokaryotes Outline of lecture today • Why you should care about the little stuff • Today - two of the three domains of life Just like organization of life (cells, tissues….populations, communities), the organization of groups of organisms is hierarchical How can we reconstruct the evolution of living things? (Chapter 25…) s rd er Fa m ily G e Sp n u ec s ie s O as Cl yl um om • Systematists study evolutionary Ph K in gd n ai om D I. Prokaryotes - Bacteria and Archaea A. The three domains of life, phylogeny B. Morphology C. Physiology/metabolism C. Ecology and key adaptations D. A few prokaryotic groups II. The evolution of the eukaryotic cell relationships • Look for shared derived (=different from ancestor) traits to group organisms • Evidence used: morphology, development, and molecular data (especially DNA sequences) 1 Why can’t we figure it out perfectly? • More distant history is obscured by more changes • Among oldest lineages of Bacteria and Archaea in particular, lots of “lateral gene transfer.” Makes it difficult to infer relationships from phylogeny of single genes. A. Phylogeny The Archaea and Bacteria are both prokaryotic,but differ more from each other than does the Archaea from the Eukarya (=plants, animals, fungi, protists). A. Phylogeny • The three domains: Bacteria, Archaea, and Eukarya. ~3 billion years ~2 billion years How did we learn the existence of the Archaea? • Difference between prokaryotes and eukaryotes long been known. • Bacteria and Archaea are superficially similar • But they are genetically different: u I. Prokaryotes - Bacteria and Archaea • The prokaryotes are the most numerous organisms on Earth •With tremendous diversity in metabolism, habitats The first Archaean genome was sequenced in 1996 most genes were very different from bacterial genes - birth of the “three domain” concept B. Morphology of prokaryotes What are they missing? Nuclei Membrane-enclosed organelles Cytoskeletons (actin and microtubules) What have they got that’s different from protists and animals? Cell wall 2 B. Morphology of the prokaryotes Prokaryotic cells are usually much smaller than eukaryotic cells Closer to the size of a mitochondrion or chloroplast Is that a coincidence? B. Morphology of the prokaryotes - Movement Simple flagella Eukaryote Cell Prokaryote Cell Gas vesicles to adjust buoyancy cyanobacteria Or gliding mechanisms spirochaete C. Ecology of the prokaryotes B. Morphology of the prokaryotes - Cell walls • Prokaryotic cell walls differ from those of eukaryotes. • Cell walls of Bacteria contain peptidoglycan (a polymer of amino sugars). Cell walls of Archaea contain proteins. • How do prokaryotes reproduce? • Asexually by fission Bacterial cell wall C. Physiology/metabolism of the prokaryotes • How do prokaryotes reproduce? • Asexually by fission • Exchange genetic information (e.g. by conjugation). In conjugation, DNA travels from donor to recipient via a cytoplasmic bridge C. Physiology/metabolism of prokaryotes • Prokaryotes - fairly narrow range of shapes and sizes, not very exciting movement: What have prokaryotes been doing for more than 3 billion years? • Learning chemical tricks 3 C. Physiology/metabolism of prokaryotes • All ancestral prokaryotes were anaerobic, and some still are: • Anaerobic: metabolism in the absence of oxygen • Aerobic: metabolism that requires oxygen • Some can shift back and forth C. Physiology/metabolism of prokaryotes • What do plants do for energy and carbon? • They use light for energy and C02 for carbon. Some prokaryotes do as well - e.g. Cyanobacteria. These are autotrophs • Why were Cyanobacteria so important in the history of life on Earth? C. Physiology/metabolism of the prokaryotes • Some chemolithotrophs live near deep sea hydrothermal vents at up to 2,500 m deep where there is no light. • Prokaryotes (mostly Archaea) that use hydrogen sulfide from deep sea volcanic vents for energy provide food for an entire bizarre community. C. Physiology/metabolism of prokaryotes • All living organisms need a source of energy and a source of carbon in order to survive and grow • What is the source of our energy and carbon? • We ingest molecules for both energy and carbon, so we’re heterotrophs • Most Bacteria and Archaea are heterotrophs as well C. Physiology/metabolism of Prokaryotes • Others have completely unique solutions, not found in eukaryotes • Some use simple nitrogen or sulfur compounds for energy, but CO2 for carbon (needing neither light nor organic compounds for food!). • Called chemolithotrophs • Have enabled life in extremely inhospitable places! C. Physiology/metabolism of the prokaryotes Lastly, some use light for energy, but need food for carbon: photoheterotrophs 4 C. Physiology/metabolism of the prokaryotes • Other chemical tricks • Fix nitrogen from atmosphere • Digest cellulose • Produce amazing toxins like polyketides (many antibiotics, e.g. tetracycline, anti-tumor drugs) • Polyketide in Paederus beetle Paederus C. Physiology/metabolism of the prokaryotes • Paederus has long been known because if crushed against skin causes rashes • Produces pederin, a polyketide • Pederin also has anti-tumor activity • Guess where Paederus gets its pederin? Paederus beetle D. Ecology of the prokaryotes - extreme habitats D. Ecology of the prokaryotes • Some Archaea are heat-loving and acidloving. • Some live in hot sulfur springs and die of “cold” at 131°F (55°C) • Some anaerobic Archaea produce methane from CO2 as a key part of their energy metabolism. • They account for the methane in the atmosphere. Archaea in sulfurous volcanic vent D. Ecology of the prokaryotes D. Ecology of the prokaryotes Some of these methane producers live in the guts of herbivorous animals More about those prokaryotes in mammalian guts - the human gut flora How many cells relative to other cells in our body? About how many species of bacteria? How much of your weight is bacteria? 5 D. Ecology of the prokaryotes Role of gut flora? Long thought to be commensal - i.e. beneficial to the bacteria, neutral for us Recent research tells a very different story E.g. bacteria are important for digesting carbohydrates, and variation in efficiencies between individuals may explain variation in tendencies towards obesity Also influence immune system (and autoimmune diseases) in development E. Important prokaryotic groups - just a few • There are far more known Bacteria than Archaea. The Bacteria The Proteobacteria Mitochondria evolved from Proteobacteria by endosymbiosis. D. Ecology of the prokaryotes • Some prokaryotes play key roles in global nitrogen cycles. E.g., nitrogen fixers, nitrifiers, and denitrifiers. Fig. 37.8 in your text Nitrogen fixation – out of the air Nitrification – from one solid form to another better for plants Denitrification – back to the air The Bacteria The Proteobacteria • By far the biggest group of Bacteria • Includes Rhizobium, the nitrogen fixing bacterium found in legume root nodules. • Also Salmonella, cholera, and E. coli. The Bacteria: Cyanobacteria • Cyanobacteria are photosynthetic •They created the oxygen atmosphere •Chloroplasts evolved from cyanobacteria by endosymbiosis •Makes you think differently about “pond scum,” no? 6 The Bacteria: Spirochaetes • Spirochaetes are corkscrew shaped •Move by axial filaments •Some are parasites of humans, e.g. agent causing syphilis, Lyme disease The Bacteria: Chlamydias Chlamydia are among the tiniest living things. Almost all parasites, e.g. a sexually transmitted disease of humans Have complex life cycle with two stages: a resting stage gets taken into host cell, the other grows and divides E. Important prokaryotic groups : Summary of the Archaea II. The origin of the eukaryotic cell • Archeans differ from Bacteria: Cell wall: proteins not peptidoglycans • Step 1? (no one knows the sequence) Increase in size • A central problem with being big is that surface area doesn’t increase as fast as volume, yet surfaces are needed for gas exchange and feeding • What’s the solution? •Very different genetically •Archeans often live in extreme habitats: tolerate high temperature, salt, low pH, absence of oxygen. II. The origin of the eukaryotic cell What’s the solution? Lose the cell wall, allow infolding of the plasma membrane to increase area The origin of the eukaryotic cell Step 2? Infolded plasma membrane attached to the chromosome may have led to a nuclear envelope. Step 3? A primitive cytoskeleton of actin and microtubules evolved. This allowed cell to change shape and move things around 7 II. The origin of the eukaryotic cell • The first eukaryotes were anaerobic. • But as oxygen increased, the oxidizing atmosphere was poisonous to anaerobes. • Step 4? Engulfing an aerobic proteobacterium resulted in mitochondria. • (Step 5? For some) Some organisms engulfed cyanobacteria and become photosynthetic (chloroplasts). 8
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