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Bacteria and Archaea The Prokaryotic Domains Prokaryotic Complexity Figure 4.5 Eukaryotic Complexity Figure 4.7 Prokaryotes • derived from ancient lineages • more biomass than all other life combined • “simple” cellular structure – no nuclear membrane – no membrane-bound organelles – no cytoskeleton • limited morphological variation Prokaryotic Morphologies Figure 27.13 Prokaryotic Morphologies Figure 27.1 photosynthetic bacteria Figure 27.7 photosynthetic archaea Figure 27.20 Prokaryotes • diverse metabolic “strategies” – photoautotrophy – chemoheterotrophy • most bacteria and archaea – chemoautotrophy – photoheterotrophy • energy from light • carbon from organic compounds Energy/carbon Table 27.2 Prokaryotes • in nearly every habitat on Earth – terrestrial – aerobic/anaerobic – marine/freshwater – deep ocean rifts/deep in crust (>2 km) – antarctic ice pack – hot/acidic (>100˚C; pH = 2-3) – salty/alkaline (pH = 11.5) – etc. Prokaryotes • a range of growth rates – generation times • 10 min • 1-3 hours • days - weeks – suspensions between growth periods • indefinite –years, decades, >century, millions? Prokaryotes • Some defy taxonomic notions – get too big – possess internal membrane systems – exhibit “eukaryote-like” growth forms Actinomycete Figure 27.16 Morphology Figure 27.3 Diplococcus Neisseria gonorrhoeae Streptococcus pyogenes Staphylococcus aureus bacterial gas vesicles Figure 27.4 Prokaryotic Taxonomy • Historically – morphology – motility (+/-) • rolling/gliding • vertical positioning • flagella & axial filaments axial filaments Figure 27.4 f l a g e l l a Figure 27.5 Prokaryotic Flagellum Figure 4.6 Gram’s Stain: Bacillus subtilis gram positive Figure 27.6 Gram’s Stain: E. coli gram negative Figure 27.6 Prokaryotic Taxonomy • Historically – morphology – motility – reactivity • Gram’s stain - peptidoglycan cell wall • metabolism –aerobic/anaerobic –resource utilization –products –inclusion bodies Mycoplasma Figure 27.17 endospore - resting body Figure 27.14 Prokaryotic Taxonomy • Historically – distinctive features • size –very large or very small • stress response –endospore formation • life style –colonial/parasitic/pathogenic Chlamydia: obligate intracellular parasite Figure 27.13 crown gall on geranium due to Agrobacterium tumefaciens Figure 27.10 Prokaryotic Taxonomy • Pathogenic requirements – contact – entry – defense evasion – multiplication – damage – infectious transfer Prokaryotic Taxonomy • Pathogen characteristics – Invasiveness – Toxigenicity • Corynebacterium diphtheriae vs. Bacillus anthracis • endotoxin vs. exotoxin –Salmonella vs. Clostridium tetani Prokaryotic Taxonomy • Koch’s postulates – Always found in diseased individuals – Grown in pure culture from host inoculant – Cultured organisms causes disease – Newly infected host produces a pure culture identical to the infective culture Prokaryotic Taxonomy • Historically – distinctive features • size –very large or very small • stress response –endospore formation • life style –parasitic/pathogenic • ecological niche Methanogens & methane using Archaea • Methanogens release 80-90% of atmospheric methane, a greenhouse gas • Methane users intercept methane seeping from sub-oceanic deposits Prokaryotic Taxonomy • Biofilm production – on solid surfaces – mixed colonies – polysaccharide matrix – resistant to treatments Recent Prokaryotic Phylogeny • Based on rRNA – evolutionarily ancient – shared by all organisms – functionally constrained – changes slowly with time – encodes signature sequences – BUT - yields a different phylogeny than other sequences analyzed Recent Prokaryotic Phylogeny • sources of phylogenetic confusion – Lateral gene transfer • among members of bacterial species • among members of different species • across domains… – phylogenetic analysis assumes cladogenic evolution • evolution may have been highly reticulate Recent Prokaryotic Phylogeny • sources of phylogenetic confusion – Mutation • prokaryotes are haploid –“recessive” mutations are not masked • prokaryotes have very little non-coding DNA • many prokaryotes have very short generation times Recent Prokaryotic Phylogeny • rRNA led to three domains – Archaea: more similar to Eukarya than to Bacteria – An ancient split between Bacteria and Archaea was followed by a more recent split between Archaea and Eukarya The Three Domain Phylogeny Figure 27.2 Shared Features of the Three Domains • • • • • • plasma membrane ribosome structure glycolysis encode polypeptide sequences in DNA replicate DNA semi-conservatively transcribe, translate with same genetic code Table 27.1 some major bacterial groups Figure 27.8 Bacterial Phylogeny • Molecular comparisons suggest several higher level groups – Proteobacteria are highly diversified • gram negative • bacteriochlorophyll • source of mitochondria • N2-fixers, Rhizobium, Agrobacterium, E. coli, Yersinia, Vibrio, Salmonella, etc. Proteobacteria Figure 27.9 Bacterial Phylogeny • Molecular comparisons suggest several higher level groups – Proteobacteria are highly diversified – ancient Cyanobacteria produced oxygen and chloroplasts • “blue-green algae” • fix CO2 & N2 • single or colonial - sheets, filaments, balls Cyanobacteria fix CO2 & N2 Figure 27.11 Cyanobacteria are pond scum Figure 27.11 Bacterial Phylogeny • Molecular comparisons suggest several higher level groups – Proteobacteria are highly diversified – ancient Cyanobacteria produced oxygen and chloroplasts – Spirochetes have axial filaments • human parasites & pathogens • free living in water sediments Spirochetes have axial filaments Figure 27.12 Bacterial Phylogeny • Molecular comparisons suggest several higher level groups – Proteobacteria are highly diversified – ancient Cyanobacteria produced oxygen and chloroplasts – Spirochetes have axial filaments – Chlamydias have a complex life cycle • obligate intracellular parasites Chlamydia Figure 27.13 Bacterial Phylogeny • Molecular comparisons suggest several higher level groups – Firmicutes: a diverse (mostly) Gram positive group • some produce endospores • some are native flora –Staphylococcus Figure 27.15 Gram + staphylococci Bacterial Phylogeny • Molecular comparisons suggest several higher level groups – Firmicutes: a diverse (mostly) Gram positive group • some produce endospores • some are native flora • some are filamentous (actinomycetes) –Mycobacterium tuberculosis –Streptomyces spp. filamentous Actinomycete Figure 27.16 Bacterial Phylogeny • Molecular comparisons suggest several higher level groups – Firmicutes: a diverse (mostly) Gram positive group • some produce endospores • some are native flora • some are filamentous (actinomycetes) • Mycoplasmas –small (~0.2 µm), no cell wall, low DNA Mycoplasma Figure 27.17 unique membrane structure Figure 27.18 unique membrane structure See page 539 Archaean Phylogeny • Most known archaea are extremophiles – many are not • Archaea cell walls lack peptidoglycan • Archaea possess unique cell membranes lipids • Archaea share rRNA signature sequences • >1/2 of Archaean genes are unlike genes known from Bacteria or Eukaryotes Archaean Phylogeny • Crenarchaeota – most live in hot, acidic habitats • 70-75˚C; pH 2-3 –Sulfolobus pH = 0.9 –Ferroplasma pH = 0.0 –some maintain internal pH 7.0 a hot, acidic home Figure 27.19 Archaean Phylogeny • Crenarchaeota – most live in hot, acidic habitats • Euryarchaeota – Methanogens [CO2 => CH4] • strict anaerobes in cow guts, rice paddies and hydrothermal vents • all share rRNA similarities Archaean Phylogeny • Crenarchaeota – most live in hot, acidic habitats • Euryarchaeota – Methanogens – extreme halophiles • e.g. in the Dead Sea • some use bacteriorhodopsin (retinal), not bacteriochlorophyll Archaean Phylogeny • Crenarchaeota – most live in hot, acidic habitats • Euryarchaeota – Methanogens – extreme halophiles – Thermoplasma • thermoacidophile, no cell wall • genome size = Mycoplasmas (1.1 x 106)