Bacteria and Archaea – morphologically simple, metabolically diverse Evolutionary relationships – the tree of life Earliest organisms on earth resembled modern forms Metabolic diversity ecological diversity A few cause diseases (+/- with humans) Others provide services (+/+ with humans) Critical for nutrient cycling Eukarya evolved from symbiosis among prokaryotes Earliest organisms on Earth Diversification over >3.5 billion years soon after Earth’s origin Earliest organisms on Earth • early Earth was a hostile environment • “oxygen revolution” – early Earth’s atmosphere was anoxic – cyanobacteria produce O2 as byproduct of photosynthesis Prokaryotes Eukaryotes 3.5 billion year old fossilized filamentous cyanobacterium modern stromatolites in Western Australia 1 Characteristics: Bacteria are ubiquitous Bacteria & Archaea Unicellular Prokaryotes (no membrane-bound nucleus) Smaller than Eukaryotic cells Domain Bacteria Peptidoglycan cell walls Plasma membrane similar to Eukarya Unique ribosomes Unique RNA Polymerase Domain Archaea Polysaccharide cell walls Unique plasma membrane Ribosomes similar to Eukarya RNA polymerase similar to Eukarya review Table 27.2 in textbook Bacteria are ubiquitous Lyme disease Yellowstone National Park Hydrothermal vent Metabolic diversity ecological diversity How do various organisms capture energy and carbon? BACTERIA & ARCHAEA PLANTS CHEMOSYNTHETIC AUTOTROPHS ANIMALS & FUNGI How can prokaryotes thrive in so many environments? 2 Metabolic diversity ecological diversity A few bacteria are pathogenic Tuberculosis Lyme disease Prokaryotes can use many energy sources in cellular respiration Prokaryotes can metabolize many inorganic compounds (e.g., NH3, CH4, H2S) Prokaryotes can ferment organic compounds other than sugar critical in decomposition (and stinky!) Gastric ulcer Photosynthetic prokaryotes – can use molecules other than H2O and O2 – several unique chlorophylls Niche partitioning for light? Borrelia burgdorferi Mycobacterium tuberculosis Heliobacter pylori salt ponds Many prokaryotes provide services Extremophiles bioremediation • halophiles (e.g., salt flats) dairy products • anaerobes (e.g. pluff mud) nutrient cycling hot springs antibiotics • thermophiles (e.g., hydrothermal vents) bacterium from Yellowstone hot spring Taq polymerase for PCR (polymerase chain reaction) revolutionized the study of genetics Thermus aquaticus Why do astrobiologists study extremophiles? 3 Phylogenetic diversity of Bacteria Ecological diversity in prokaryotes cholera anthrax Lyme disease leprosy botox Bern and Goldberg, 2005 BACTERIA Phylogenetic diversity of Archaea Cyanobacteria thermophiles ~80 species, but very abundant metabolize iron (Ferroplasma sp.) Misnomer “blue-green algae” Nostoc sp. Photosynthetic Some colonial halophiles thermophiles stromatolites are living fossils in symbiosis with fungi: metabolize sulfur Spirulina sp. methanogens 4 Does metabolic diversity reflect phylogenetic diversity? Does metabolic diversity reflect phylogenetic diversity? Predict how metabolism maps onto the phylogeny Independent evolution of complex metabolic pathways? Lateral gene transfer Major groups of Bacteria and Archaea Some placement not fully resolved Evolutionary relationships – the tree of life Domain Eukarya nuclear envelope nucleus and organelles complex cytoskeleton cells usually larger than prokaryotes evolved from prokaryotes 5 Eukaryote innovations Protists – paraphyletic and diverse unicellular or multicellular • complex cell structures • sexual reproduction & multicellularity Eukaryotes sexual repro & multicellularity variation in cell covering cell walls shells cell membrane only Ecological diversity of Protists Protists – paraphyletic and diverse pathogens unicellular or multicellular variation in cell covering absorption variation in feeding mode ingestion predators primary producers African sleeping sickness decomposers and parasites motility photosynthesis producers How did eukaryotes evolve from prokaryotes? cilia flagella predators & filter feeders Potato blight decomposers symbionts How did plants, fungi and animals evolve from protists? 6 1. How did eukaryotes evolve from prokaryotes? Inferred common ancestor of all modern eukaryotes: BA unicellular membrane-bound nucleus cytoskeleton no cell wall mitochondria 1. How did eukaryotes evolve from prokaryotes? B. EVOLUTIONARY INNOVATION: Mitochondria • site of cellular respiration • double membrane A. EVOLUTIONARY INNOVATION: Endomembrane system • • • Allows cells to be bigger Surface area to volume ratio Compartmentalization – Endoplasmic Reticulum – Golgi Complex – Vacuoles – Nuclear Membrane 1. How did eukaryotes evolve from prokaryotes? • α-proteobacteria-like • mutualistic, symbiotic relationship Evolution of eukaryotes: membrane infolding and endosymbiosis C. EVOLUTIONARY INNOVATION: Chloroplasts • site of photosynthesis • double membrane • cyanobacteria-like 7 2. How plausible is the Endosymbiosis Theory? 2. How plausible is Endosymbiosis Theory? Evidence: More evidence: Traits shared by prokaryotes and mitochondria/chloroplasts: Molecular (DNA) phylogeny: predict location of mitochondria and chloroplasts • Circular DNA • Divide by fission • Small size • Distinct ribosomes Double membranes Prochloron Morphological similarity to cyanobacteria Modern examples exist of protists with bacterial endosymbionts What if mitochondria did not arise via endosymbiosis? Phylogenetic diversity of protists Eight major, morphologically distinct groups Does ecological diversity reflect phylogenetic diversity? Parasites Malaria Relationships among groups not completely resolved Potato blight Amoebic dysentery African sleeping sickness Intestinal giardia 8 Does ecological diversity reflect phylogenetic diversity? Primary producers Photosynthetic protists are not closely related Dinoflagellates Diatoms Red algae PLANTAE Red Algae – most are multicellular Green Algae – unicellular, colonial, multicellular STRAMENOPILA Brown Algae – multicellular Diatoms – unicellular or chains “Algae” are polyphyletic ALVEOLATA Dinoflagellates – unicellular or colonial DISCICRISTATA Euglenids – unicellular, also ingest food Paraphyletic vs. polyphyletic? Euglena How did this arise? Green algae Brown algae 3. How did photosynthetic eukaryotes evolve? What are three hypotheses for the polyphyly of photosynthesis in protists? 1) 2) Secondary endosymbiosis – lateral transfer of chloroplasts among protists 3) clue: some protist groups have chloroplasts with >2 membranes solution: lateral transmission of chloroplasts = secondary endosymbiosis 9 4. How did large, complex organisms evolve? Tree of life or web of life? D. EVOLUTIONARY INNOVATION: Multicellularity • differentiation among cells • interdependent cells Rare examples of cell specialization within bacterial colonies (δ-proteobacteria) What two processes have contributed to this pattern? 4. How did large, complex organisms evolve? 5. How did sexual reproduction evolve? E. EVOLUTIONARY INNOVATION: Multicellularity F. EVOLUTIONARY INNOVATION: Meiosis fertilization = haploid diploid meiosis = diploid haploid haploid or diploid stage can be dominant Alternation of generations – multicellular haploid and diploid How many independent origins of multicellularity? 10