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MICROBIOLOGY MIMM211 (Biology of Microorganisms) Lecture 3 Dr. Benoit Cousineau Department of Microbiology & Immunology McGill University Phylogeny evolutionary relationship between organisms • Before the concept of evolution, organisms were grouped based on morphological similarities (taxonomy), no relationships between the groups • Many fossils of animals and plants were found and used to suggest the appearance of the different groups, built evolutionary family trees • Microorganisms were left out until late 1960s - No microbial fossils - Very similar shapes • Use ubiquitous gene sequence to compare microorganisms and construct a universal phylogenetic tree of life (include microorganisms) Evolution of early schemes for classifying microbes • Traditional early schemes (before 1866) - Plants and animals • Ernest Haeckel’s proposal (1866) - Plants, animals, and microorganisms 1 Haeckel’s tree of life Evolution of early schemes for classifying microbes • Traditional early schemes (before 1866) - Plants and animals • Ernest Haeckel’s proposal (1866) - Plants, animals, and microorganisms • Edouard Chatton (1937) - Eucaryotes and procaryotes • Roger Stainer & C.B. van Niel - Supported Chatton’s proposal Recent classifications of living organisms • The five-kingdom scheme of Robert Whittaker (1959) -Monera, Protista, Fungi, Plantae, and Animalia 2 Whittaker’s five kingdom classification Recent classifications of living organisms • The five-kingdom scheme of Robert Whittaker (1959) -Monera, Protista, Fungi, Plantae, and Animalia • The three-kingdom scheme of Carl Woese and colleagues (1977). They used a molecular approach (16S ribosomal RNA gene) - Bacteria, Archaea, and Eucaryotes - These three groups are equally distant Woese’s three kingdom scheme 3 Evolution of classification schemes Three kingdom universal tree of life The endosymbiotic theory of evolution • Procaryotes do not contain organelles (mitochondria, chloroplasts) • Almost all eucaryotes have organelles - Plants have both mitochondria and chloroplasts - All other eucaryotes only have mitochondria • Mitochondria and chloroplasts resemble procaryotes - Approximately the same size (1 µM) - The only organelles containing DNA (an active genome) - Mitochondrial and chloroplast genomes are circular - They contain 70S ribosomes (80S in eucaryotes) - Double membrane resemble cytoplasmic and outer membranes of gram-negative bacteria, however they do not have the cell envelope - They multiply and divide by binary fission 4 The endosymbiotic theory of evolution (2) • Lynn Margulis proposed in 1981 the endosymbiotic theory: A primitive eucaryotic cell engulfed an ancient procaryote to create the first eucaryotic cell - First procaryote appeared 3.5 billion years ago - First eucaryote appeared only 1.0 billion years ago - The two cells continued to evolve in a mutually beneficial symbiotic relationship for 1.0 billion year - The procaryotic cell benefited from having a sheltered environment rich in nutrients - The eucaryotic cell benefited from containing an organism that produced energy (ATP) by respiration (mitochondria) or by photosynthesis (chloroplast) - Mitochondria are the descendants of an alpha proteobacteria (gram-negative, Rickettsia prowazekii) - Interestingly R. prowazekii is an obligate intracellular parasite - Chloroplasts are the descendants of a cyanobacteria (Prochloron) The endosymbiotic theory of evolution 5 Woese’s three kingdom scheme The endosymbiotic theory of evolution (3) • More recent endosymbiosis (an evolutionary snap shot) - Eucaryotes constantly phagocytoze procaryotic cells - Need time to co-evolve and create a stable new organism • Aphids and related insects: a recent endosymbiotic relationship (200 million years) - The endosymbiont still has its gram-negative double membrane and cell envelope, phylogenetically closely related to E. coli - Stable endosymbiotic event - The endosymbiont can no longer grow outside its host - Aphids are also dependant of their endosymbiont, they die if the endosymbiont is killed (antibiotic) - Supports the endosymbiotic theory of evolution Building a phylogenetic tree • Get the sequence of an ubiquitous gene (phylogenetic marker, e.g., 16S ribosomal RNA gene, 1600 nucleotides) for all the organisms to be included in the phylogenetic tree • Align the gene sequences using a sequence alignment program • Feed the aligned sequences to a phylogenetic algorithm - Find which sequences share the highest level of homology and link closest neighbors - Create phylogenetic tree, branching the organisms related to the similarity of their gene sequence - The length of the branches linking the different organisms are proportional to their evolutionary distance 6 The bacterial kingdom 7