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Chemoheterotrophs Chapter 5 Metabolism: Lipids, Anabolism, Autotrophs Energy & carbon extraction from organic sources Note how biochemically inferior humans are to many bacteria in this area: Many microbial chemoheterotrophs can live on a single organic food source (glucose only; citrate only; etc.) What would happen to you if all you ate was sugar? WHY? Dr. Amy Rogers Bio 139 Fall 2006 Office Hours: Mondays & Wednesdays, ~8:30-10:00 AM Some figures taken from Krogh Biology: A Guide to the Natural World Chemoheterotrophs: Fat •Catabolism of glucose for energy: •Glycolysis •Fermentation OR aerobic respiration (Krebs, ETC) •Anaerobic respiration (ETC without oxygen as electron acceptor) •Catabolism of proteins for energy: •Deamination for aerobic respiration (products enter Krebs cycle) •Decarboxylation in absence of oxygen Fat catabolism: Triacylglycerols Carbohydrates & Proteins are not the only organic energy sources: Lipids (here, all saturated) Lipids are frequently stored as fats in the form of triacylglycerol (triglycerides) 1. Hydrolysis to glycerol & free fatty acids Glycerol (3 carbons) enters glycolysis 2. Breakdown of fatty acids into “activated” 2 carbon units β (beta)-oxidation 2 carbon units are oxidized, broken off and bound to Coenzyme A to make Acetyl-CoA Repeat β-oxidation, 2 carbons at a time, until the fatty acid is gone. (“activated” 2 carbon units: remember pyruvate going from glycolysis to Krebs?) All these oxidations, also make reduced molecules: NADH & FADH2 1 Anabolism • Bacteria are remarkable chemical factories • Consider this week’s news article (bioplastics, biofuels) • Various species of bacteria can make all kinds of weird stuff (reduced electron carriers) (NADH, FADH2) • We have dissected a single catabolic pathway (glycolysis & aerobic respiration) in detail; many others exist • What about anabolism (biosynthesis)? DO NOT memorize any names/details from this figure The point: Catabolic pathways provide both the energy (ATP) and the initial substrates for anabolic pathways in complex, interconnected webs of reactions. Anabolism • Every reaction in a biochemical pathway is catalyzed by a specific enzyme • For example, synthesis of the amino acid tryptophan requires at least 13 different enzymes Produced during glycolysis • Lack of any one enzyme in a pathway prevents synthesis of the final product The product must be found in the environment, or the organism dies Essential amino acids, vitamins, etc. Krebs cycle intermediate Auxotrophs & Prototrophs A mutation that abolishes function of a single enzyme can make a bacterium dependent on an outside source of a nutrient (can’t synthesize it, gotta eat it) Photoautotrophs (as opposed to chemoheterotrophs) Produce chemical energy from light; carbon source: CO2 • Cyanobacteria, green algae, plants: Photosynthesis We call such mutated strains of bacteria auxotrophs: bacteria with specific unusual nutritional requirements as a result of mutation (lost ability to make it for themselves) • Use electrons from H2O to reduce CO2 to carbohydrate • Produce oxygen (waste) • Probably modified earth’s atmosphere by dumping O2 • Green & Purple Sulfur bacteria: Prototroph: a wild type strain of the same species, with all its genes intact, can grow without addition of that particular nutrient to the media • Use H2S, produce sulfur or sulfuric acid • Strict anaerobes, and do not produce oxygen • Came earlier in evolution, when atmosphere was hydrogen-rich 2 Chemoautotrophs • Inorganic energy sources (chemo-) – Energy extracted by oxidizing inorganic chemicals – Hydrogen, ferrous ions, ammonia, etc. • CO2 as carbon source (-auto-) – Can synthesize all organic substances they need • Carbohydrates, fats, proteins, nucleic acids (some bacteria can oxidize gold, copper, even uranium!) Example: Nitrifying bacteria that use ammonia as an energy source Archaea in deep ocean volcanic vents are in this group Nitrifying Bacteria The Nitrogen Cycle (chemoautotrophs) • Nitrifying bacteria use ammonia (NH3) & nitrite (NO2) as energy sources • Nitrogen is a necessary element for life (amino acids, nucleic acids) • The ultimate product of such oxidation is nitrate (NO3) • Nitrogen gas (N2) is abundant in the atmosphere, but in this chemical form, it is inaccessible to most living things • These chemicals can be oxidized, therefore can be energy sources! • Nitrate is the form of nitrogen most usable by plants • Nitrifying bacterial play a crucial ecological role • Atmospheric N2 must be fixed (converted to a usable form) Fixation: conversion of a gas into a solid or liquid form by chemical reaction Nitrogen Cycle Nitrogen Cycle • Nitrification is an oxidation (NH3 NO3) • autotrophic nitrifying bacteria get energy • Nitrogen fixation is a reduction; energy is consumed (N2 NH3) (costs energy) (terminal electron acceptor) • Some nitrogen fixing bacteria are symbiotic, living inside plants which provide the energy in exchange for the fertilizer • Denitrification (returning organic nitrogen back to N2 gas), in Oxidation by nitrifying bacteria (yields energy) turn, is a reduction. Under anaerobic conditions, some bacteria use nitrate instead of oxygen as a terminal electron acceptor. • See Nitrate Reduction test (lab #20) 3 Relevant reading in Black’s Microbiology: (pages from 6th edition) • Ch. 5 • Ch. 6, page 190 (auxotrophs & prototrophs) • Ch. 25, pages 751-754 (nitrogen cycle) 4