Apicoplast. Apicoplast - history. Treatments and New drug targets
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1 Treatments and New drug targets What is the apicoplast? Where does it come from? How are proteins targeted to the organelle? How does the organelle replicate? What is the function of the organelle? - history Discovery just waiting to happen! mysterious membrane-bound organelle viewed in 1960s. Term coined only in 1995 Four membranes Fatty acid synthesis proteins are targeted to apicoplast 1998 New drug - Human trials Fosmidomycin Isoprenoid biosynthesis Good tolerance Recrudescence 1
2 Apicomplexans have 3 genomes Band 3: Nuclear genome on chromosomes Band 2: Small 6 kb mitochondrial genome (linear) Band 1: 35 kb circular genome Discovered in 1975 Isopycnic Centrifugation (Density gradient separation) CsCl Sample EtBr Before centrifugation After centrifugation Common apicoplast ancestor ~35 kb circular genome Toxoplasma Plasmodium Eimeria Gene synteny Similarity to chloroplast genes 2 large subunit rrna 2 small subunit rrna 3 subunits of RNA polymerase Ribosomal proteins Complete set of trnas Elongation factor (EF-TS) Chaperone proteins (clp) Nothing related to photosynthetic machinery Big surprise - no apicoplast DNA in Cryptosporidium, no apicoplast Subcellular Localization of the 35 kb DNA How? Methods Localization in
3 What is a plastid? General term for membrane-bound organelles found in plants and algae. Contain DNA, various pigments and various storage products. Plastid forms are interconvertible. (ie. many fall colors results from transition of chloroplast to chromoplast in deciduous trees) Originated from an endosymbiotic cyanobacteria. Undifferentiated colorless chlorophyll develops in dark colorless Apicomplexans synthesize amylopectin! - Highly branched glucose polymer Not water soluble Starch storage Pigments red, yellow carotenoids Roots, seeds, fruit leaves, stems Petals, fruit Plastid Evolution Endosymbiotic origin s Microscopic observations - pigments Discovery of extrachromosomal DNA Phylogenetic studies Close relationship to cyanobacteria Primary endosymbiosis Eukaryote + cyanobacterium Secondary endosymbiosis Eukaryote + algae Plastid Evolution A single primary endosymbiotic event gave rise to all present day plastids Primary plastid account for only a small fraction of plastid biodiversity At least 3 major secondary endosymbiotic events Numerous cases of loss of photosynthetic machinery 3
4 Evolution Primary Endosymbiosis: ORIGIN OF THE CHLOROPLAST Secondary Endosymbiosis: ORIGIN OF THE APICOPLAS T Protein targeting via secretory pathway Bipartite signal is required for apicoplast targeting At least a two-step process Hydrophobic signal peptide - secretory signal A) Transit Domain structure peptide of - nuclear-encoded translocation into apicoplast plastid proteins signal sequence plastid-targeting domain Mature Plastid Protein B) targeting Deletion / Reconstitution Identifies a Bipartite Targeting Signal... signal seq. transit peptide Acyl Carrier Protein Green Fluorescent Protein ACP(L) ACP(L) apicoplast 3Δ3 3Δ3 5Δ4 secreted P30 (signal) 5Δ2 P30-5Δ2 P30-5 Δ2 cytoplasmic apicoplast... that is evolutionarily conserved P. falciparum ACP (signal + transit) A. thaliana ftsz (transit) Pf ACP P30-At FTSZ Pf ACP apicoplast 4
5 /Mitochondria Association Toxoplasma Mitochondrion Plasmodium Mitochondrion /Mitochondria Association Plasmodium Toxoplasma Dinoflagellate Plastid Division with Cell Cycle Toxoplasma divides by Endodyogeny 2 daughter cells inside mother 1 - Interphase is apical to nucleus 2 - S phase Centrioles duplicated DNA replicated and divided Segregation Associated with spindle poles 4 - Segregation/growth U-shaped structure 5 - Division Centriole association Movement into daughter bud Nuclear fission 1 4 DNA 2 3 5
6 Division * division 6
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