Chapter 9: Cellular Respiration and Fermentation - See figure 9.2 9.1: Catabolic pathways yield energy by oxidizing organic fuels Catabolic pathways and production of ATP - Compounds that can participate in exergonic reactions can act as fuels - Fermentation – partial degradation of sugars or other organic fuel that occurs without the use of oxygen - Aerobic respiration – oxygen is consumed along with organic fuel to produce carbon dioxide, water and ATP - Cellular respiration – includes both anaerobic and aerobic processes - C6H12O6 +O2 CO2 + H2O +Energy - The breakdown of glucose is exergonic, has a free energy change of -686 kcal/mol - (Recall a negative change in energy indicates the products store less energy than the reactants and the reaction can occur spontaneously without an input of energy) Redox Reactions: Oxidation and Reduction - The principle of redox o Transfer of electrons (e-) in a reaction from one reactant to another o Oxidation – loss of eo Reduction – gain of eo Reducing agent – e- donor o Oxidizing agent – e- acceptor o See figure 9.3 - Oxidation of organic fuel molecules during cellular respiration o In cellular respiration Glucose is oxidized Oxygen is reduced o Oxidation of glucose transfers electrons to a lower energy state - Stepwise energy harvest of NAD+ and the electron transport chain (ETC) o If energy is released all at once, energy can not be harvested efficiently for constructive work o H+ are transferred via electron carriers called NAD+ and FADH (electrons travel with protons) o Most electrons flow from glucose – NADH – ETC – oxygen The stages of cellular respiration: A preview 1. Glycolysis 2. Kreb’s cycle (Citric acid cycle) 3. ETC - See figure 9.6 - Eukaryotes – step 2 and three occur in mitochondria - Prokaryotes- step 2 takes place in cytosol, step 3 takes place along the plasma membrane - Oxidative phosphorylation – energy from each step is stored in a form the mitochondrion can use to make ATP o Powered by redox reactions of ETC Substrate-level phosphorylation – enxyme transfers a phosphate group from a substrate molecule to ADP The cell makes about 32 molecules of ATP from each molecule of glucose (each ATP has 7.3 kcal/mol of free energy) 9.2: Glycolysis harvests chemical energy by oxidizing glucose to pyruvate - See figure 9.9 Glycolysis – means “sugar splitting” 6-carbon glucose is broken down into 2 3-carbon molecules called pyruvate To begin glycolysis, 2 molecules of ATP are invested Net energy yield is 2 ATP (4 are made, but two are spent in the beginning) 2 NADH are formed and transferred to ETC 9.3: After pyruvate is oxidized, the citric acid cycle completes the energyyielding oxidation of organic molecules - See figure 9.10, 9.11, 9.12 Oxidation of pyruvate to acetyl CoA - Pyruvate is converted to a compound called acetyl CoA in a transition step o Releases carbon dioxide (2 total molecules, 1 per pyruvate) The citric acid cycle - The cycle generates 1 ATP per turn (2 molecules total) and 3 carbon dioxide molecules (6 in total, 2 from the conversion of pyruvate to acetyl CoA) - Most of the chemical energy is transferred to NAD+ and FAD - 8 NADH and 2 FADH2 are produced from each glucose molecule which is then transferred to the ETC 9.4: During oxidative phosphorylation, chemiosmosis couples electron transport to ATP synthesis The Pathway of Electron Transport - ETC is a collection of molecules embedded in the inner membrane of the mitochondria in eukaryotic cells - Most components are proteins, numbered I through IV - Cytochromes – electron carriers - NADH and FADH2 each carry a pair of electrons o FADH2 provides less energy for ATP synthesis than NADH does Chemiosmosis: The energy-coupling mechanism - ATP synthase – enzyme that makes ATP from ADP and inorganic phosphate o Works like an ion pump running in reverse o Uses the energy of an existing ion gradient to power ATP synthesis o Power source is a difference in the [H+] o See figure 9.14 - Chemiosmosis – energy stored in the form of a hydrogen ion gradient across a membrane is used to drive cellular work - Establishing the H+ gradient is a major function of the ETC - Proton-motive force – H+ gradient that results from H+ being pumped into intermembrane space - Chemiosmosis is an energy-coupling mechanism that uses energy stored in the form of a H+ gradient across a membrane to drive cellular work o Eukaryotic cells, energy comes from exergonic redox reactions o ATP synthesis is the work performed - See figure 9.15 An accounting of ATP production by cellular respiration - Each NADH molecule generates enough proton-motive force to synthesize approximately 2.5 ATP molecules - See figure 9.16 - Each FADH2 generates approximately 1.5 ATP molecules - Each glucose molecule generates 30-32 molecules of ATP - Approximately 34% of the energy from glucose is transferred to ATP production 9.5: Fermentation and anaerobic respiration enable cells to produce ATP without the use of oxygen - Anaerobic respiration and fermentation do not require oxygen to generate ATP Types of fermentation - Glycolysis plus reactions that generate NAD+ by transferring electrons from NADH to pyruvate - Alcohol fermentation o Pyruvate is converted to ethanol in two steps o See figure 9.17a o First step releases carbon dioxide from pyruvate o Second step acetalaldehyde is reduced by NADH to ethanol which regenerates the supply of NAD+ - Lactic acid fermentation o Pyruvate is reduced directly by NADH to form lactate with no release of carbon dioxide o Humans do this during short bursts of activity when oxygen is scarce Comparing fermentation with anaerobic and aerobic respiration - All three pathways use glycolysis to oxidize glucose to pyruvate - Difference among the three pathways is the mechanisms for oxidizing NADH back o NAD+ - Obligate anaerobes carry out fermentation or anaerobic respiration only - Faculative anaerobes make ATP using fermentation or respiration The evolutionary significance of glycolysis - Oldest known fossils of bacteria produced oxygen as a by-product of photosynthesis Therefore early prokaryotes may have generated ATP exclusively from glycolysis Glycolysis occurred in cytosol which shows that membrane-bound organelles were not necessary 9.6: Glycolysis and the citric acid cycle connect to many other metabolic pathways The versatility of catabolism - We obtain calories from fat, protein, starch and other sugars - All of these organic molecules can be used for cellular respiration to make ATP - See figure 9.19 - Beta oxidation – breaks fatty acids down to two-carbon fragments - A gram of fat oxidized by cellular respiration produces more than twice as much ATP as a gram of carbohydrate Biosynthesis (Anabolic pathways) - Some organic monomers are used to provide carbon skeletons that are required to make other molecules Regulation of cellular respiration via feedback mechanisms - See figure 9.20 - Most common mechanism fro control is feedback inhibition – end product of anabolic pathway inhibits the enzyme that catalyzes an early step of the pathway o Prevents needless diversion of metabolic intermediates from uses that are more urgent - Catabolic controls are based mainly on regulating the activity of enzymes at strategic points in the pathway