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Lecture _8 - Respiration - Glycolysis and The Citric Acid Cycle

Course: BIO 203, Spring 2010
School: San Diego State
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Glycolysis Respiration: and the Citric Acid Cycle Campbell Reece, Eighth Edition Reece Eighth Reading Reading Assignment: Chapters 9 Overall equation gives the impression that it occurs in a single step. However, aerobic respiration takes place in about 20 steps, grouped into four stages: 1) Glycolysis 2) Formation of acetyl conenzyme A 3) The citric acid cycle (Krebs cycle) 4) The electron transport chain and...

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Glycolysis Respiration: and the Citric Acid Cycle Campbell Reece, Eighth Edition Reece Eighth Reading Reading Assignment: Chapters 9 Overall equation gives the impression that it occurs in a single step. However, aerobic respiration takes place in about 20 steps, grouped into four stages: 1) Glycolysis 2) Formation of acetyl conenzyme A 3) The citric acid cycle (Krebs cycle) 4) The electron transport chain and chemiosmosis BIOL 203 Spring 2010 Ralph Feuer Overview: Life Is Work Living cells require energy from from outside sources Some animals, such as the giant panda, obtain energy by eating plants ti Some animals feed on other organisms that eat plants Energy Flows Energy flows into an ecosystem as sunlight Leaves as heat Photosynthesis generates O2 and organic molecules, Used in cellular respiration Cells use chemical energy stored in organic molecules Regenerate ATP Powers work Energy Flows Energy flows into an ecosystem as as sunlight Leaves as heat Light energy ECOSYSTEM Photosynthesis generates O2 and organic molecules, Used in cellular respiration Photosynthesis in chloroplasts CO2 + H2O Cellular respiration in mitochondria Organic +O molecules 2 Cells use chemical energy stored in in organic molecules Regenerate ATP Powers work ATP ATP ATP powers most cellular work Heat energy Catabolic Pathways Yield Energy By Oxidizing Oxidizing Organic Fuels Several processes are central to cellular respiration and related pathways Catabolic Pathways and Production Production of ATP The breakdown of organic molecules is exergonic Fermentation Fermentation is a partial degradation of sugars that occurs without O2 Aerobic respiration consumes organic molecules and O2 and yields ATP Anaerobic respiration is similar to aerobic respiration but consumes compounds other than O2 Cellular Respiration Includes both aerobic and anaerobic respiration Often used to refer to aerobic respiration CYTOSOL Electron shuttles span membrane 2 NADH or 2 FADH2 2 NADH 6 NADH MITOCHONDRION 2 NADH 2 FADH2 Although carbohydrates, fats, and proteins are all consumed as fuel Helpful to trace cellular respiration with the sugar glucose: Glycolysis Glucose 2 Pyruvate 2 Acetyl CoA Citric acid cycle Oxidative phosphorylation: electron transport and chemiosmosis + 2 ATP + 2 ATP + about 32 or 34 ATP Maximum per glucose: About 36 or 38 ATP C6H12O6 + 6 O2 6 CO2 + 6 H2O + Energy (ATP + heat) Redox Reactions: Oxidation and Reduction The transfer of electrons during chemical reactions Releases energy stored in organic molecules molecules This released energy is ultimately used to synthesize ATP The Principle of Redox Chemical reactions that transfer electrons between reactants are called oxidation-reduction reactions id Redox reactions A substance loses electrons Oxidized becomes oxidized (loses electron) becomes reduced (gains electron) In oxidation becomes oxidized becomes reduced In reduction A substance gains electrons Reduced The amount of positive charge is reduced Reducing Agents and Oxidizing Agents The electron donor The Called the reducing agent Reactants Products becomes oxidized The electron receptor Called the oxidizing agent becomes reduced Some redox reactions do not transfer electrons But change the electron sharing in covalent bonds An example is the reaction between between methane and O2 Methane (reducing agent) Oxygen (oxidizing agent) Carbon dioxide Water Reactants becomes oxidized Products becomes reduced reduced Methane (reducing agent) Oxygen (oxidizing agent) Carbon dioxide di Water Oxidation of Organic Fuel Molecules During Cellular Respiration During cellular respiration The fuel (such as glucose) is oxidized O2 is reduced becomes oxidized becomes reduced Introduction to Electron Transport Chains NADH passes the electrons to the electron transport chain Electron transport chain Passes electrons in a series of steps Unlike an uncontrolled, explosive reaction Explosive release of heat and light and l ight energy H2 + 1/2 O2 2H (from food via NADH) Controlled rel ease of 2 H+ + 2 e energy for synthesis of ATP 1/ 2 O2 O2 pulls electrons down the chain in an energy-yielding tumble tumble (a) Uncontrolled reaction (b) Cellular respiration 1/ O 22 The energy yielded is used to regenerate ATP Stepwise Energy Harvest via NAD+ and the Electron Transport Chain In cellular respiration: Glucose and other organic molecules are broken down in a series of steps Electrons from organic compounds are usually first transferred to NAD+ A coenzyme Nicotinamide adenine dinucleotide NAD+ as an electron shuttle Two nucleotides joined together at their phosphate groups (yellow) Nicotinamide (shown in white) is a nitrogenous base; but not present in DNA or RNA 2 e + 2 H+ 2 e + H+ NADH Dehydrogenase NAD+ + 2[H] Reduction of NAD+ Oxidation of NADH Nicotinamide (reduced form) Nicotinamide (oxidized form) + H+ H+ Stepwise Energy Harvest via NAD+ and the Electron Transport Chain In cellular respiration: Glucose and other organic molecules are broken down in a series of steps Electrons from organic compounds are usually first transferred to NAD+ A coenzyme NAD+ functions as an oxidizing agent during cellular respiration NAD+ is an electron acceptor Dehydrogenase Each NADH (the reduced form of NAD+) Represents stored energy Tapped to synthesize ATP The Stages of Cellular Respiration: A Preview Cellular respiration has three stages: Glycolysis Breaks down glucose into two molecules of pyruvate Electrons carried via NADH Glycolysis Glucose Pyruvate Cytosol ATP Substrate-level phosphorylation The Stages of Cellular Respiration: A Preview Cellular respiration has three stages: Glycolysis Breaks down glucose into two molecules of pyruvate Electrons carried via NADH Electrons carried via NADH and FADH2 The citric acid cycle Completes the breakdown of glucose Glycolysis Glucose Pyruvate Citric acid cycle Mitochondrion Cytosol ATP Substrate-level phosphorylation ATP Substrate-level phosphorylation The Stages of Cellular Respiration: A Preview Cellular respiration has three stages: Glycolysis Breaks down glucose into two molecules of pyruvate Electrons carried via NADH Electrons carried via NADH and FADH2 The citric acid cycle Completes the breakdown of glucose Glycolysis Glucose Pyruvate Citric acid cycle Oxidative phosphorylation: electron transport and chemiosmosis Mitochondrion Cytosol ATP ATP Substrate-level phosphorylation ATP Oxidative phosphorylation Oxidative phosphorylation Accounts for most of the ATP ATP synthesis Substrate-level phosphorylation Oxidation Phosphorylation Process that generates most of the ATP Powered by redox reactions Oxidative phosphorylation Accounts for almost 90% of the ATP generated by cellular respiration A smaller amount of ATP is formed in glycolysis and the citric acid cycle Substrate-level phosphorylation Enzyme ADP P Substrate Enzyme + Product ATP Glycolysis Harvests Chemical Energy By Oxidizing Oxidizing Glucose To Pyruvate Glycolysis Splitting of sugar Breaks down glucose into two molecules of pyruvate Occurs in the cytoplasm Two major phases Energy investment phase Energy payoff phase Energy investment phase Glucose 2 ADP + 2 P ADP 2 ATP ATP used Energy payoff phase 4 ADP + 4 P 4 ATP formed 2 NAD+ + 4 e + 4 H+ 2 NADH + 2 H+ 2 Pyruvate + 2 H2O Pyruvate Net Glucose 4 ATP formed 2 ATP used 2 NAD+ + 4 e + 4 H+ 2 Pyruvate + 2 H2O 2 ATP 2 NADH + 2 H+ A Closer Look At Glycolysis Energy Investment Phase Phase Glucose Glucose enters the cell cell Phosphorylated by hexokinase Sugar trapped in cell More chemically reactive Transfer of a phosphate group Pair of electrons ATP 1 Hexokinase ADP Glucose Glucose-6-phosphate ATP 1 Hexoki nase ADP Glucose-6-phosphate A Closer Look At Glycolysis Energy Investment Phase Phase Glucose-6-phosphate converted to its isomer Fructose-6phosphate Glucose ATP 1 Hexokinase ADP Glucose-6-phosphate 2 Phosphoglucoisomerase Glucose-6-phosphate 2 Phosphoglucoisomerase Fructose-6-phosphate Fructose-6-phosphate The Structure and Classification of Some Monosaccharides Monosaccharides Molecular formulas that are usually multiples of CH2O Trioses (C3H6O3) Pentoses (C5H10O5) Hexoses (C6H12O6) Glucose (C6H12O6) Most common monosaccharide Grape sugar, corn sugar, blood sugar One of the main products of ph photosynthesis Source of energy Glycolysis Dihydroxyacetone Glyceraldehyde Ribose Glucose Galactose Monosaccharides are classified by The location of the carbonyl group (as (as aldose or ketose) The number of carbons in the carbon skeleton Ribulose Fructose Linear and Ring Forms Though often drawn as linear skeletons, in aqueous solutions many sugars form rings Chemical equilibrium between the linear and ring structures greatly favors the formation of rings Monosaccharides serve as a major fuel for cells and as raw material for building molecules (a) Linear and ring forms (b) Abbreviated ring structure Examples Examples of Disaccharide Synthesis Disaccharide formation Dehydration reaction joins two monosaccharides 14 glycosidic linkage This covalent bond is called a glycosidic linkage Maltose Malt sugar Glucose Glucose Maltose 12 glycosidic linkage (a) Dehydration reaction in the synthesis of maltose th Sucrose Table sugar Glucose Fructose Lactose Galactose and Glucose Milk sugar Sucrose (b) Dehydration reaction in the synthesis of sucrose A Closer Look At Glycolysis Energy Investment Phase Transfer of a phosphate group from ATP to the sugar Fructose-1,6-bisphosphate Investment of another ATP So far, 2 ATPs used Sugar now ready to be split Phosphate groups on its opposite ends Glucose-6-phosphate 2 Phosphoglucoisomerase Glucose ATP 1 Hexokinase ADP Fructose-6-phosphate ATP 3 Fructose-6-phosphate ATP 3 Phosphofructokinase ADP Key step in regulation of glycolysis Phosphofructokinase allosterically regulated by ATP and its products Inhibited by ATP and citrate (citric acid cycle) Activated by AMP and fructose 2,6bisphosphate Phosphofructokinase ADP Fructose1, 6-bisphosphate Fructose1, 6-bisphosphate Regulation Of Enzyme Activity Helps Control Metabolism Chemical chaos Result if a cells metabolic pathways were not tightly regulated Allosteric enyzme with four subunits Active site (one of four) Regulation of a cell Switching on or off the genes that encode specific enzymes Regulating the activity of enzymes Regulatory site (one (one of four) Activator Active form Stabilized active form Allosteric regulation may either inhibit or stimulate stimulate an enzymes activity Occurs when a regulatory molecule binds to a protein at one site and affects the proteins function at another site Most allosterically regulated enzymes are made from polypeptide subunits Each enzyme has active and inactive forms The binding of an activator stabilizes the active active form of the enzyme The binding of an inhibitor stabilizes the inactive form of the enzyme Oscillation NonInhibitor form functional Inactive form active site Stabilized inactive inactive form (a) Allosteric activators and inhibitors A Closer Look At Glycolysis Energy Investment Phase Cleavage reaction from which glycolysis gets its name Cleaves sugar molecule Two different three-carbon sugars Dihydroxyacetone phosophate Glyceraldehyde-3phosphate Isomers of each other Glucose ATP 1 Hexokinase ADP Glucose-6-phosphate 2 Phosphoglucoisomerase Fructose1, 4 Fructose-6-phosphate Aldolase ATP 3 6-bisphosphate Phosphofructokinase ADP 5 Isomerase Fructose1, 6-bisphosphate 4 Aldolase 5 Isomerase Dihydroxyacetone phosphate Glyceraldehyde3-phosphate Glyceraldehyde3-phosphate Dihydroxyacetone phosphate A Closer Look At Glycolysis Energy Investment Phase Isomerase catalyzes the reversible conversion between: Dihydroxyacetone and Glyceraldehyde-3-phosphate Reaction never reaches equilibrium Next enzyme uses only glyceraldehyde-3-phosphate as its substrate Glucose ATP 1 Hexokinase ADP Glucose-6-phosphate 2 Phosphoglucoisomerase Fructose1, 6-bisphosphate 4 Fructose-6-phosphate Aldolase Equilibrium is pulled in the direction of glyceraldehyde3-phosphate Removed as fast as it is formed ATP 3 Phosphofructokinase ADP 5 Isomerase Fructose1, 6-bisphosphate 4 Aldolase Net reaction: Cleavage of six-carbon sugar into two molecules of glyceraldehyde-3-phosphate Dihydroxyacetone phosphate 5 Isomerase Dihydroxyacetone phosphate Glyceraldehyde3-phosphate Glyceraldehyde3-phosphate A Closer Look At Glycolysis Energy Payoff Phase Enzyme catalyzes two sequential reactions reactions Holds glyceraldehyde-3phosphate in its active site Transfer of electrons and H+ to NAD+ Forming NADH Redox reaction 2 NAD+ 2 NADH + 2 H+ 6 Triose phosphate dehydrogenase 2 Pi 2 1, 3-Bisphosphoglycerate First, sugar is oxidized Glyceraldehyde3-phosphate 2 NAD+ 6 Triose phosphate dehydrogenase 2 Pi Very exergonic reaction Enzyme utilizes the released energy to attach a phosphate group to the oxidized substrate Product of very high energy Inorganic phosphate ions always present in the cytosol 2 NADH NADH + 2 H+ 2 1, 3-Bisphosphoglycerate Notice that the coefficient of 2 Precedes all molecules in the energy payoff phase A Closer Look At Glycolysis Energy Payoff Phase Glycolysis produces some ATP molecules molecules By substrate-level phosphorylation Exergonic reaction 2 NAD+ 6 Triose phosphate dehydrogenase 2P i 2 NADH + 2 H+ 2 1, 3-Bisphosphoglycerate 2 ADP 7 Phosphoglycerokinase 2 ATP For each glucose molecule that began glycolysis Sugar was oxidized in Step #6 Step #7 produces 2 ATP Recall that 2 ATP were invested to get sugar ready for splitting ATP debt repaid 2 1, 3-Bisphosphoglycerate 2 ADP 2 3-Phosphoglycerate 2 ATP 7 Phosphoglycerokinase Two molecules of 3phosphoglycerate molecules Not a sugar Carbonyl group that characterizes a sugar has been oxidized to a carboxyl group Hallmark of an organic acid 2 3-Phosphoglycerate A Closer Look At Glycolysis Energy Payoff Phase Enzyme relocates the remaining phosphate phosphate group Preparing the substrate for the next reaction 2 NAD+ 6 Triose phosphate dehydrogenase 2 Pi 2 NADH + 2 H+ 2 1, 3-Bisphosphoglycerate 2 ADP 7 Phosphoglycerokinase 2 ATP 2 3-Phosphoglycerate 8 Phosphoglyceromutase 2 3-Phosphoglycerate 8 2 2-Phosphoglycerate Phosphoglyceromutase 2 2-Phosphoglycerate A Closer Look At Glycolysis Energy Payoff Phase Enzyme causes a double bond to form form in the substrate Extracting a water molecule Yielding phosphenolpyruvate (PEP) Electrons of the substrate are rearranged Resulting phosphorylated compound has a very high potential energy Allows Step #10 to St #10 occur 2 2 NAD+ 6 Triose phosphate dehydrogenase 2 Pi 2 NADH + 2 H+ 2 1, 3-Bisphosphoglycerate 2 ADP 7 Phosphoglycerokinase 2 ATP 2 3-Phosphoglycerate 2 2-Phosphoglycerate 9 8 Phosphoglyceromutase 2-Phosphoglycerate 2 H2O Enolase 9 2 H 2O Enolase 2 Phosphoenolpyruvate 2 Phosphoenolpyruvate A Closer Look At Glycolysis Energy Payoff Phase Last reaction of glycolysis Produces more ATP Transferring the phosphate group from PEP to ADP Second instance of substrate-level phosphorylation Occurs twice for each glucose molecule molecule 2 ATP are produced 2 3-Phosphoglycerate 2 NAD+ 6 Triose phosphate dehydrogenase 2 Pi 2 NADH + 2 H+ 2 1, 3-Bisphosphoglycerate 2 ADP 7 Phosphoglycerokinase 2 ATP ATP 2 Phosphoenolpyruvate 2 ADP Overall, glycolysis has used 2 ATP and produced 4 ATP Net gain of 2 ATP 8 Phosphoglyceromutase Glycolysis has repaid the ATP investment with 100% interest Additional energy was stored by Step #6 NADH can be used to make ATP by oxidative phosphorylation O2 must be present 2 ATP 2 2-Phosphoglycerate 10 Pyruvate kinase 9 2 H 2O Enolase 2 Phosphoenolpyruvate 2 ADP 10 Pyruvate kinase Chemical energy in pyruvate can be extracted Citric acid cycle O2 must be present O2 not present 2 ATP 2 Pyruvate 2 Pyruvate Fermentation A Closer Look At Glycolysis Energy Payoff Phase Last reaction of glycolysis Produces more ATP Transferring the phosphate group from PEP to ADP Second instance of substrate-level phosphorylation Occurs twice for each glucose molecule molecule 2 ATP are produced Glucose Glycolysis Pyruvate No O2 present: Fermentation O2 present: Aerobic cellular respiration CYTOSOL Overall, glycolysis has used 2 ATP and produced 4 ATP Net gain of 2 ATP Glycolysis has repaid the ATP investment with 100% interest Additional energy was stored by Step #6 NADH can be used to make ATP by oxidative phosphorylation O2 must be present MITOCHONDRION Ethanol or lactate Acetyl CoA Citric acid cycle Chemical energy in pyruvate can be extracted Citric acid cycle O2 must be present O2 not present Fermentation A Closer Look At Glycolysis Energy Payoff Phase Last reaction of glycolysis Produces more ATP Transferring the phosphate group from PEP to ADP Second instance of substrate-level phosphorylation Occurs twice for each glucose molecule molecule 2 ATP are produced Overall, glycolysis has used 2 ATP and produced 4 ATP Net gain of 2 ATP Glycolysis has repaid the ATP investment with 100% interest Additional energy was stored by Step #6 NADH can be used to make ATP by oxidative phosphorylation O2 must be present Chemical energy in pyruvate can be extracted Citric acid cycle O2 must be present O2 not present Fermentation The Citric Acid Cycle Completes The Energy Yielding Oxidation Of Organic Molecules Presence of O2 Pyruvate enters the mitochondrion Help of a transport protein Before the citric acid cycle can begin Pyruvate must be converted to acetyl CoA Links the cycle to glycolysis CYTOSOL NAD+ 2 MITOCHONDRION NADH + H+ Complex of several enzymes Catalyze three steps Pyruvates carboxyl group is removed Fully oxidized (Little chemical energy) Given off as C02 Acetate Enzyme transfers extracted electrons to NAD+ Unstable bond (wavy line) Very reactive Has a high potential energy Molecule ready to feed its acetyl group into the citric acid cycle Acetyl CoA 1 Pyruvate Transport protein CO2 3 Coenzyme A Acetyl CoA Remaining 2 carbon fragment is oxidized Coenzyme A is attached to the acetate The Citric Acid Cycle Also called the Krebs cycle Takes place within the mitochondrial matrix Pyruvate NAD+ NADH + H+ Acetyl CoA CoA CoA CO2 CoA The cycle oxidizes organic fuel py Derived from pyruvate Generating per turn: 1 ATP 3 NADH 1 FADH2 ATP FADH2 FAD Citric acid cycle 2 CO2 3 NAD+ 3 NADH + 3 H+ ADP + P i The Citric Acid Cycle Also called the Krebs cycle Takes place within the mitochondrial matrix it The cycle oxidizes organic fuel Derived from pyruvate Generating per turn: 1 ATP 3 NADH 1 FADH2 Redox cofactor Energy-carrying molecule Flavin Adenine Dinucleotide (FADH2) The Citric Acid Cycle Citric acid cycle Eight steps Each catalyzed by a specific enzyme Pyruvate NAD NAD+ NADH + H+ Acetyl CoA CoA CoA CO2 CoA Acetyl group of acetyl CoA Joins the cycle by combining with oxaloacetate Forming citrate The next seven steps Decompose the citrate back to oxaloacetate Making the process a cycle FADH2 FAD ADP + P i ATP Citric acid cycle 2 CO2 3 NAD+ 3 NADH + 3 H+ The NADH and FADH2 produced by the cycle Relay electrons extracted from food to the electron transport chain The Citric Acid Cycle Acetyl CoA Adds its two-carbon acetyl group to oxaloacetate Producing citrate Acetyl CoA CoASH 1 Oxaloacetate Citrate Citric acid cycle The Citric Acid Cycle Citrate is convert to its isomer Isocitrate Removal of one water Addition of another water Acetyl CoA CoASH 1 H2 O Oxaloacetate Citrate 2 Isocitrate Citric acid cycle The Citric Acid Cycle Isocitrate is oxidized Reducing NAD+ to NADH NAD NADH Loss of one C02 molecule Acetyl CoA CoASH 1 H2 O Oxaloacetate Citrate 2 Isocitrate Citric acid cycle 3 NAD+ NADH + H+ CO2 -Ketoglutarate The Citric Acid Cycle Another CO2 is lost Resulting compound is oxidized Reducing NAD+ to NADH Remaining molecule is attached to coenzyme A Unstable bond (wavy line) Acetyl CoA CoASH 1 H2 O Oxaloacetate Citrate 2 Isocitrate Citric acid cycle 3 NAD+ NADH + H+ CO2 CoASH 4 -Ketoglutarate NAD+ NADH + H+ CO2 Succinyl CoA The Citric Acid Cycle Coenzyme A is displaced by a phosphate group Phosphate group transferred to GTP Molecule with functions similar to ATP Acetyl CoA CoASH GTP may be used to make an ATP 1 H2 O Oxaloacetate Citrate 2 Isocitrate Citric acid cycle 3 NAD+ NADH + H+ CO2 CoASH CoASH 4 -Ketoglutarate 5 NAD+ CO2 Succinate GTP GDP ADP ATP Pi Succinyl CoA NADH + H+ The Citric Acid Cycle Two hydrogens are transferred to FAD Forming FADH2 FADH Oxidizing succinate to fumarate 1 Acetyl CoA CoASH H2O Oxaloacetate Citrate 2 Isocitrate Citric acid cycle Fumarate 3 NAD+ NADH + H+ CO2 CoASH 6 CoASH 4 -Ketoglutarate FADH2 FAD 5 NAD+ CO2 Succinate GTP GDP ADP ATP Pi Succinyl CoA NADH + H+ The Citric Acid Cycle Addition of a water molecule Rearranges bones in the substrate Malate Acetyl CoA CoASH 1 H2O Oxaloacetate Malate Citric acid cycle Fumarate Citrate 2 Isocitrate 3 NAD+ NADH + H+ H2 O 7 CO2 CoASH 6 CoASH 4 -Ketoglutarate FADH2 FAD 5 NAD+ CO2 Succinate GTP GDP ADP ATP Pi Succinyl CoA NADH + H+ The Citric Acid Cycle Substrate is oxidized Reducing NAD+ to NADH Regeneration of oxaloacetate NADH +H+ NAD+ 8 Acetyl CoA CoASH 1 H2O Oxaloacetate Citrate 2 Malate Citric acid cycle Fumarate Isocitrate 3 NAD+ NADH + H+ H2 O 7 CO2 CoASH 6 CoASH 4 -Ketoglutarate FADH2 FAD 5 NAD+ CO2 Succinate GTP GDP ADP ATP Pi Succinyl CoA NADH + H+ During Oxidative Phosphorylation, Chemiosmosis Couples Couples Electron Transport To ATP Synthesis Following glycolysis and the citric acid cycle, NADH and FADH2 account for most of the energy extracted from food These two electron carriers donate electrons to the electron transport chain Powers ATP synthesis via oxidative phosphorylation CYTOSOL Electron shuttles span membrane 2 NADH or 2 FADH2 2 NADH NADH 6 NADH NADH MITOCHONDRION 2 NADH NADH 2 FADH2 FADH Glycolysis Glucose 2 Pyruvate 2 Acetyl CoA Citric acid cycle Oxidative phosphorylation: electron transport and chemiosmosis + 2 ATP + 2 ATP + about 32 or 34 ATP Maximum per glucose: About 36 or 38 ATP During Oxidative Phosphorylation, Chemiosmosis Couples Couples Electron Transport To ATP Synthesis Chemiosmosis Diffusion of ions across a selectively-permeable membrane Generation of ATP by the movement of hydrogen ions across a membrane during cellular respiration CYTOSOL Electron shuttles span membrane 2 NADH or 2 FADH2 2 NADH NADH 6 NADH NADH MITOCHONDRION 2 NADH NADH 2 FADH2 FADH Glycolysis Glucose 2 Pyruvate 2 Acetyl CoA Citric acid cycle Oxidative phosphorylation: electron transport and chemiosmosis + 2 ATP + 2 ATP + about 32 or 34 ATP Maximum per glucose: About 36 or 38 ATP
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UCLA - ECON - 41
UCLA - ECON - 41
. . . . . . . . . . 1995RADICALLY ELEMENTARY PROBABILITY THEORYBYEDWARD NELSONPRINCETON UNIVERSITY PRESSPRINCETON, NEW JERSEY 1987 519.2 . : . . : - - . 1995. 124 . ,
UCLA - ECON - 41
EC 41 UCLA Sample Problems #3b Regarding Chapter 2 sections 2.5 and 2.6, and Chapter 4 sections 4.1 & 4.2 These problems will NOT be collected or graded, but they will be useful for studying for exams. 1) The table below gives counts for a sample of 100 p
UCLA - ECON - 41
EC 41 UCLA Fall 2008 Sample Problems #4; RE material in Ch 4, and Ch 2 sections 4, 5, & 6 These problems will NOT be collected or graded, but they will be useful for studying for exams. 1) The true annual return on a $200 investment in Zipco stock is as f
UCLA - ECON - 41
EC 41 UCLA Fall 2008 Sample Problems #5; RE Section 4.5 and Ch 5 material Midterm 2 will cover material from Chapter 2, sections 2.4, 2.5, & 2.6 and Chapters 4 and 5 These problems will NOT be collected or graded, but they will be useful for studying for
UCLA - ECON - 41
UCLA - ECON - 41
UCLA - ECON - 41
EC 41 UCLA Fall 2008 Sample Problems #6; RE Chapters 5 & 6 We will skip Chapters 8 and 9 and end the class with section 10.1 1) Flip a fair coin and consider the number of flips required for the first head. If we graph probability of first head on the ver
UCLA - ECON - 41
UCLA - ECON - 41
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UCLA - ECON - 41
UCLA - ECON - 41
TI-84 Plus TI-84 Plus Silver Edition GuidebookImportant InformationTexas Instruments makes no warranty, either express or implied, including but not limited to any implied warranties of merchantability and fitness for a particular purpose, regarding any
UCLA - MGMT 120A - 260491200
SYLLABUS The Anderson Graduate School of Management, UCLACourse Title Course Number Quarter Instructor Meeting Times Meeting Dates Final Exam Date Class Location UCLA Office UCLA Office Phone Number UCLA Office Hours E-mail address Intermediate Financial
UCLA - MGMT 120A - 260491200
Chapter 5Income Measurement and Profitability AnalysisAACSB assurance of learning standards in accounting and business education require documentation of outcomes assessment. Although schools, departments, and faculty may approach assessment and its doc
UCLA - ECON - 171
Econ 171 - Industrial Organization Second Midterm Exam - 2009 InstructionsName: Section:There are 6 short questions and 2 problems. Short questions are worth a total of 40 points and the problems a total of 60 points. Answer in the space provided (no ne
UCLA - ECON - 171
Econ 171 - Industrial Organization First Midterm Exam - Feb 2010 InstructionsName: Section:There are 6 short questions and 2 problems. Short questions are worth a total of 40 points and the problems a total of 60 points. Answer in the space provided (no
UCLA - ECON - 171
Schedule and requirementsRequirements: The course will have 2 midterms and a final exam. The final grade is computed as follows: weight Midterms (best of 2) Final exam 40% 60%Exams: Midterm 1: Feb 2 Midterm 2: March 2 Final: Monday, March 15, 2010, 11:3
NYU - ORGO 1 AND - V.0243
Answers to Exam 3, Chemistry 0243 2008 1. a.H H H (CH3)3C H H H H H H H Hb.CH3CH3CH3 ClClClCl +CH3ClCH3 CH3 Cl2. (a) CH2N2 (diazomethane) and light (b) CF3COOOH (c) KMnO4 or (b) and either HO /H2O or H3O /H2O (d) Br2/H2O (e) 1. BH3 2. HOOH/HO
NYU - ORGO 1 AND - V.0243
You must do this First Question. It should take no more than five minutes) 1. (a). Draw a perfect structure for tert-butylcyclohexane in its energy minimum form. Include the hydrogens attached to the six-membered ring.(CH3)3C(b) Draw a perfect mechanism
NYU - ORGO 1 AND - V.0243
Some answers to the Chemistry 301-301A Final Examination, January 12, 2005 1.H3 C O H OH2 H3O + H2O H3 C O + CH3H3 CO CH3H3 C+O CH3HO+ H H2OH2OH H3 C H O CH3 OOH2 H H3 C O+ CH3 H O H2O H H +CH3 OH H CH3 OH CH3 H OH3O+++ OCH3IR: 1725, 28
NYU - ORGO 1 AND - V.0243
C hemistry 301A-301X Final Examination: January 12, 2005 Why, for example, should a group of simple, stable compounds of carbon, oxygen, and nitrogen struggle for billions of years to organize themselves into a professor of chemistry? What is the motive?
NYU - ORGO 1 AND - V.0243
Some Answers to the Final Examination, Chemistry 301X- 2006 1. (a) OH HO A HO H H OH BCDOH E H (b) Isomer A is an enol and will isomerize to the ketone form: HO A enol (c) Problem 15.46H F OHOketone(d) One Possibility - there are others. one dibrom
NYU - ORGO 1 AND - V.0243
Chemistry 301X Final Examination: January 24, 2006 If you are obliged to neglect any thing, let it be your chemistry. It is the least useful and the least amusing to a country gentleman of all the ordinary branches of science. Thomas JeffersonThis Final
NYU - ORGO 1 AND - V.0243
Final Examination, Chemistry 301X, January 24, 2007 This Final Examination is different in very few respects from the hour exams with which you are all too familiar. However, there is choice. DO ONLY EIGHT (8) QUESTIONS PLEASE do not do all the questions.
NYU - ORGO 1 AND - V.0243
Some Answers to the 301X Final Examination, January 24, 2007 1. The two diastereomers are, of course, the cis and trans diols. CH3 OH OH cis 1 trans 1 CH3 OH OHThe simplest mechanism through which both diols would give 2 and 3, is an SN1-like process goi
NYU - ORGO 1 AND - V.0243
Answers to Hour examination #1, Chemistry 301-301A, 2003 1. (a) yes (b) no - atoms moved (c) yes (d) no - valence violated (e) no - valence violated (f) yes (g) yes (h) no - atoms moved 2. CH 3 O C H O C C N(CH 3 )2 H O H CH 3 O C C C N(CH 3 )2 HCH 3 O C
NYU - ORGO 1 AND - V.0243
Some Answers to Hour Examination #2, Chemistry 301X - 2006 1a and b. Here are the drawings: The two diastereomers are shown at the top and the products at the bottom. In these SN2 reactions, the leaving group tosylate must be in the position as shown so t
NYU - ORGO 1 AND - V.0243
C hemistry 301-301A - Hour Examination #2, November 17, 2003 There aint no answer. There aint going to be any answer. There never has been an answer. Thats the answer. Gertrude Stein1 (20 points). Explain carefully why reaction (a) fails but (b) succeeds
NYU - ORGO 1 AND - V.0243
Some answers to Hour Examination #2, Chemistry 301-301A, 20031. (a) Bromide is not a strong nucleophile, and would never displace the poor leaving group ethoxide. SN1 is impossible as a primary cation would have to be formed. (b) Here, protonation of the
NYU - ORGO 1 AND - V.0243
H our Examination #2, Chemistry 301-301A, November 15, 2004 Then, when the work was done my mother and father would stand there in the middle of the big, bright room and say poems or sing. How strange it seemed to me that all these serious, hard-working p