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7 Cellular Respiration and Fermentation
CAMPBELL BIOLOGY IN FOCUS Urry • Cain • Wasserman • Minorsky • Reece 7 Cellular Respiration and Fermentation Kathleen Fitzpatrick and Nicole Tunbridge, Simon Fraser University Modified by James R. Jabbur, Houston Community College © 2016 Pearson Education, Inc

Overview: Life Is Work Living cells require energy from outside sources Energy flows into an ecosystem as sunlight and leaves as heat Photosynthesis generates O2 and organic molecules, which are used in cellular respiration Cells use chemical energy stored in organic molecules to regenerate ATP, which powers work For the Discovery Video Space Plants, go to Animation and Video Files.

Animation: Carbon Cycle

Organic molecules Cellular respiration in mitochondria
Light energy ECOSYSTEM Photosynthesis in chloroplasts Organic molecules CO2 + H2O + O2 Cellular respiration in mitochondria Figure 9.2 Energy flow and chemical recycling in ecosystems ATP ATP powers most cellular work Heat energy

Cellular Respiration: A CSI Investigation
Click to add notes

Today you are an intern working with the Crime Scene Investigation division of the Houston Police Department Forensic unit. You will be assisting them with the molecular analysis. It’s your first day and you get to assist on a suspected homicide case!

A call is placed to emergency services by a desperately ill woman…
Click to add notes

Operator: 911 State your problem
Moaning, gasping voice: I am dizzy, I just threw up and I’ve got a terrible headache. Operator: Can you tell me what happened before you began to feel ill? Moaning, gasping voice: My husband gave me a calcium pill before leaving for work and I feel sick, really sick. I think there is, something, something, terribly wrong. Operator: Stay calm an ambulance is dispatched. Ma’am, are you still with me? (No Response). *DATA SHEET INFO*

Obviously, we need more information to solve this puzzle!
What could have caused the woman's death? (you are thinking to yourself, “how the heck do I know”). Obviously, we need more information to solve this puzzle! Click to add notes

What is the importance of oxygen to our cells?
Why do we need to breath? Click to add notes What is the importance of oxygen to our cells? *DATA SHEET INFO*

A few days later, histology report is in!
In the histological analysis, specific staining dyes were used to access cellular damage. Tissue samples from heart, kidney, liver and lung show massive cell death. Click to add notes Heart Kidney Liver Lung *DATA SHEET INFO*

Specific Cellular Staining Indicates Organellar Damage.
What Organelle is shown below? Does this help?!!! Click to add notes *DATA SHEET INFO*

The autopsy report showed massive cell death in several tissues
The autopsy report showed massive cell death in several tissues. What do you think keeps cells alive? RIGHT! Cells metabolize food, using oxygen, to produce energy to live. Food? Oxygen? Energy? Yes, but she had eaten and she’s still DEAD Yes, but she was breathing and she’s still DEAD

What kind of food provides you with energy?
Do Carbohydrates? Do Fats? Both? **

Since the victim was breathing and had eaten, something was obviously wrong with her ability to produce energy! Let’s look at the process that provides you with energy, called Cellular Respiration. Click to add notes

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Instructor, play “big picture”

Eventually, you will need to determine exactly which part of cellular respiration was affected (Glycolysis, Kreb’s Cycle or Electron Transport System). Click to add notes

Concept 7.1: Catabolic pathways yield energy by oxidizing organic fuels
The breakdown of organic molecules is exergonic 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 (remember FONClBrISCH?) Although carbohydrates, fats, and proteins are all consumed as fuel, it is helpful to trace cellular respiration with the sugar glucose: C6H12O6 + 6 O2  6 CO2 + 6 H2O + Energy (ATP + heat) *DATA SHEET INFO*

…and yes, you must memorize that reaction!

Redox Reactions: Oxidation and Reduction
The transfer of electrons during chemical reactions releases energy stored in organic molecules, ultimately used to synthesize ATP Chemical reactions that transfer electrons between reactants are called oxidation-reduction reactions, or redox reactions In oxidation, a substance loses electrons, or is oxidized In reduction, a substance gains electrons, or is reduced (the amount of positive charge is reduced)

OILRIG Oxidation Is Loss Reduction Is Gain …of electrons

Oxidation of Organic Fuel Molecules During Cellular Respiration
During cellular respiration, the organic fuel (glucose sugar) is oxidized and oxygen is reduced The series of reactions are coupled; the general (stoichiometrically balanced) equation is shown below… becomes oxidized becomes reduced *DATA SHEET INFO*

Stepwise Energy Harvest via Nicotinamide Adenine Dinucleotide (NAD+) and the Electron Transport Chain When glucose is oxidized, the covalent bonds are broken between the carbons of the sugar and the energy is transferred as electrons to another molecule, which is classified as a “cofactor” Oxidized NAD+ forms a high energy intermediate carrier, reduced NADH (which has electrons) Dehydrogenase Energy carrier

NADH NAD+ What is FAD  FADH2? *DATA SHEET INFO*
Figure 7.4 NAD+ as an electron shuttle What is FAD  FADH2? *DATA SHEET INFO*

Finally, each NADH passes its electrons into the electron transport chain, where oxygen pulls electrons down the chain in an energy-yielding tumble, regenerating ATP Of course, the reduced form of NADH, which has lost its electrons, recycles back to the oxidized form of NAD+ The same can be said of oxidized FAD and reduced FADH2 We will cover more about this, later!

Biological Energy transfer through Chemiosmosis
+ 1/2 O2 (from food via NADH) Peter Mitchell super genius Controlled release of energy for synthesis of ATP 2 H e– ATP ATP Free energy, DG Electron transport chain ATP 2 e– Figure 9.5 An introduction to electron transport chains 1/2 O2 2 H+ H2O (b) Cellular respiration Chemical (bonding)  Electrical (e-)  Potential (H+ gradient)  Mechanical (drives ATP synthase)  Metabolic (ATP made)

Hydrogen Bomb Figure 7.5 An introduction to electron transport chains

The Stages of Cellular Respiration: A Preview
Cellular respiration has four general stages: Glycolysis breaks down glucose into two molecules of pyruvate in the cell cytosol Pyruvate is ‘shipped’ into the mitochondrial matrix; during which, Pyruvate Oxidation produces acetate The Krebs/Tri Carboxylic Acid/Citric Acid - Cycle then completes the breakdown of acetate in the mitochondrial matrix Oxidative Phosphorylation via Chemiosmosis accounts for most of the ATP synthesis in the mitochondrial membrane BioFlix: Cellular Respiration *DATA SHEET INFO*

A substrate-level phosphorylation is a simple transfer of a high-energy phosphate bond from a substrate to a product (this process is not the same as oxidative phosphorylation, as you will soon see!) Enzyme Enzyme ADP P Substrate + ATP Product

Concept 9.2: Glycolysis harvests chemical energy by oxidizing glucose to pyruvate in the cytosol
Glucose (6C) (into cell cytoplasm) Main Points of Glycolysis: Split 1 molecule of glucose into 2 molecules of pyruvate 2. Energy Investment: *lose 2ATP 3. Energy Produced: *gain 2NADH *gain 4ATP Net Energy Yield: *gain 2ATP 5. Phosphofructokinase: *Conrol Point *ATP inhibits allosterically *Committed Step 1 Glucose ATP ADP Glucose-6-P Fructose-6-P phosphofructokinase enzyme ATP ADP Fructose-1,6-bisP (6C) 2 Glyeraldehyde-3-P (3C) 2 NAD 2 NADH (5 more steps) 2 ADP 2 ATP 2 ADP 2 ATP 2 Pyruvate (3C) *DATA SHEET INFO*

The Energy Input and Output of Glycolysis
Figure 7.8 The energy input and output of glycolysis

http://highered. mcgraw-hill
Instructor, play “glycolysis”

Aldolase Isomerase Fructose- 1, 6-bisphosphate 4 5 Dihydroxyacetone
Glucose ATP 1 Hexokinase ADP Glucose-6-phosphate 2 Phosphoglucoisomerase Fructose- 1, 6-bisphosphate 4 Fructose-6-phosphate Aldolase ATP 3 Phosphofructokinase ADP Figure 9.9 A closer look at glycolysis 5 Isomerase Fructose- 1, 6-bisphosphate 4 Aldolase 5 Isomerase Dihydroxyacetone phosphate Glyceraldehyde- 3-phosphate Dihydroxyacetone phosphate Glyceraldehyde- 3-phosphate

Substrate-level phosphorylation is shown…
2 NAD+ 6 Triose phosphate dehydrogenase 2 NADH 2 P i + 2 H+ 2 1, 3-Bisphosphoglycerate 2 ADP 7 Phosphoglycerokinase 2 ATP 2 Phosphoenolpyruvate 2 ADP 2 3-Phosphoglycerate 8 10 Phosphoglyceromutase Pyruvate kinase 2 ATP 2 2-Phosphoglycerate 9 Enolase 2 H2O Figure 9.9 A closer look at glycolysis 2 Phosphoenolpyruvate 2 ADP 10 Pyruvate kinase 2 ATP 2 Pyruvate Substrate-level phosphorylation is shown… 2 Pyruvate

Concept 7.3: After pyruvate is oxidized, the Krebs Cycle completes the energy-yielding oxidation of organic molecules in the mitochondrial matrix Pyruvate (3C) (into mitochondrial matrix) Pyruvate Dehydrogenase Enzyme Complex (PDC): 1. multi-protein complex 2. AMP stimulates allosterically Pyruvate Dehydrogenase Enzyme Complex Pyruvate (3C) + CoA + NAD+ Acetyl (2C)–CoA + NADH + CO2 Main points of Pyruvate Dehydrogenation 1. Lose (2x) 1 CO2 = 2 CO2 2. Gain (2x) 1 NADH = 2 NADH 3. Gain (2x) Acetyl = 2 Acetyl 1 mole of Glucose yields 2 moles of pyruvate, thus, the dehydrogenation goes twice, producing 2 NADH and 2 CO2. Acetyl (2C) + OAA OAA (4C) Citrate (6C) 1 mole of Glucose yields 2 moles of pyruvate, producing 2 moles of Acetyl. Thus, the cycle goes twice, producing 6 NADH, 2 FADH2, 2 GTP and 2 CO2. 3NAD 3NADH Main points of Kreb’s Cycle 1. 7 intermediate steps 2. Lose (2x) 2 CO2 = 4 CO2 3. Gain (2x) 3 NADH = 6 NADH 4. Gain (2x) 1 FADH2 = 2 FADH2 5. Gain (2x) 1 GTP = 2 GTP 6. Recycle Oxaloacetate (OAA) 7 enzyme specific catalyzed steps in Krebs Cycle 1FADH 1FADH2 1GDP 1GTP 2CO2 *DATA SHEET INFO*

Instructor, play “krebs cycle”

Citric acid cycle Succinyl CoA
Acetyl CoA CoA—SH NADH +H+ 1 H2O NAD+ 8 Oxaloacetate 2 Malate Citrate Isocitrate NAD+ Citric acid cycle NADH 3 7 + H+ H2O CO2 Fumarate CoA—SH -Keto- glutarate Figure 9.12 A closer look at the citric acid cycle 4 6 CoA—SH FADH2 5 CO2 NAD+ FAD Succinate P NADH i GTP GDP Succinyl CoA + H+ ADP ATP

A circular six carbon molecule bound together by covalent bonds!
Review Ms. Glucose A circular six carbon molecule bound together by covalent bonds! c GLYCOLYSIS (split glucose in half) Lose 2 carbons; break 2 bonds PYRUVATE DECARBOXYLATION Lose 4 carbons; break 4 bonds KREB’S CYCLE Oops! No more carbons; no more bonds to break! 40

So if Ms. Glucose loses all of her carbons (or breaks all of her covalent bonds) by the end of the Kreb’s Cycle, why do we need the electron transport system and the process of Oxidative Phosphorylation? Think about this. Why? Oops! 41

New information has just come in from the pathology report!
BIOASSAY ANALYSIS OF ATP LEVELS IN DAMAGED HEART CELLS Click to add notes ATP LEVELS IN CYTOPLASM ATP LEVELS IN MITOCHONDRIA NORMAL DRASTICALLY REDUCED *DATA SHEET INFO*

ATP ANALYSIS IN DAMAGED HEART CELLS
What cellular process or processes do you think are impaired in the woman’s damaged cells? What cellular process produces the maximum ATP in the cell? Click to add notes Don’t know? Let’s dig deeper and understand the biochemistry involved!

Click to add notes Let’s see how the majority of ATP is made by the cell and why the mitochondrion is called, “the powerhouse of the cell!”

Lets review what we have covered up to now!
1. GLYCOLYSIS PYRUVATE OXIDATION CITRIC ACID CYCLE How much ATP has been produced by Substrate-Level Phosphorylation? What molecules will produce ATP by Oxidative Phosphorylation? 45

Concept 7.4: During Oxidative Phosphorylation, Chemiosmosis couples electron transport to ATP synthesis in the mitochondrial membrane by utilizing a generated Proton Motive Force NADH (from GPS) or NADH NAD+ FADH2 O2 H+ H+ MITOCHONDIAL MATRIX – LOW [H+] H+ ADP + P FAD+ NAD+ ATP H2O e- Ubiquinone Cyt C Reductase Inner Membrane (Cristae) ATP Synthase Cyt C Oxidase e- NADH dehydrogenase e- e- e- Coenzyme Q Cyt C H+ ~3 protons ~3 protons INTERMEMBRANE SPACE – HIGH [H+] ~3 protons For the Cell Biology Video ATP Synthase 3D Structure — Side View, go to Animation and Video Files. For the Cell Biology Video ATP Synthase 3D Structure — Top View, go to Animation and Video Files. Outer membrane Main Points of Energy Harvesting: 1. NADH produces 2.5 ATP 2. FADH2 produces 1.5 ATP 3. Cytosolic NADH produces 1.5 ATP (Glycerol Phosphate Shuttle - GPS) 4. Recycle electron carrier intermediate cofactors!!! CELL CYTOPLASM *DATA SHEET INFO*

Chemical Energy (bonding between atoms)
Electrical Energy (electrons from broken bonds) Potential Energy (electrons drive a proton concentration gradient) Mechanical Energy (proton motive force drives ATP synthase) Metabolic Energy (~P is attached to ADP and ATP is produced)

ATP Synthase (optional presentation)

ATP Synthase: 3-D Side and Top View

ATP Synthase, A Molecular Mill
Figure 7.13 ATP synthase, a molecular mill

Summary: The Pathway of Electron Transport
The electron transport chain is in the mitochondrial membrane Electrons are transferred from NADH or FADH2 to the electron transport chain Most of the chain’s components are multiprotein complexes Carriers alternate reduced and oxidized states as they accept and donate electrons Electrons drop in free energy as they go down the chain and are finally passed to O2, forming H2O The electron transport chain generates no ATP; electron transfer causes proteins to pump H+ from the mitochondrial matrix to the intermembrane space The chain’s function is to break the large free-energy drop from food to O2 into smaller steps that release energy

Summary: Chemiosmosis, Energy-Coupling Mechanism
Electron transfer in the electron transport chain causes proteins to pump H+ from the mitochondrial matrix to the intermembrane space H+ then moves back across the membrane, passing through channels in ATP synthase; ATP synthase uses the exergonic flow of H+ to drive the phosphorylation of ATP from ADP This is an example of chemiosmosis, the use of energy in a H+ gradient to drive cellular work The energy stored in a H+ gradient across a membrane couples the redox reactions of the electron transport chain to ATP synthesis The H+ gradient is referred to as a proton-motive force, emphasizing its capacity to do work

An Accounting of ATP Production by Cellular Respiration
Energetics of Glucose Catabolism Process Molecules Formed/Used ATP Produced Glycolysis -2 ATP ATP 4 ATP ATP (net 2ATP) 2 NADH ATP (eukaryotes) *5 ATP (prokaryotes) Pyruvate Oxidation 2 NADH ATP Krebs Cycle 6 NADH ATP 2 FADH ATP 2 GTP ATP TOTALS: net 30 ATP (eukaryotes) net 32 ATP (prokaryotes) *Note: Prokaryotes have no organelles and don’t have to pump glycolysis derived NADH into a mitochondria (this takes energy). Thus, prokaryotes derive more ATP from the catabolism of glucose. *DATA SHEET INFO*

Instructor, this material is redundant

*DATA SHEET INFO* Glycolysis Oxidative Pyruvate phosphorylation:
Electron shuttles span membrane 2 NADH or 2 FADH2 2 NADH 2 NADH 6 NADH 2 FADH2 Glycolysis Oxidative phosphorylation: electron transport and chemiosmosis Pyruvate Oxidation 2 Pyruvate 2 Acetyl CoA Citric acid cycle Glucose CELL CYTOSOL MITOCHONDRIAL MATRIX INTER-MITOCHONDRIAL MEMBRANE + 2 ATP + 2 ATP + about 26 or 28 ATP Figure 9.17 ATP yield per molecule of glucose at each stage of cellular respiration About 30 or 32 ATP Maximum per glucose: *DATA SHEET INFO*

ATP ANALYSIS IN DAMAGED HEART CELLS
So once again, what cellular process do you think is impaired in the woman’s damaged cells? Hint: What cellular process produces the majority of ATP? (remember last slide) *DATA SHEET INFO* **

Thankfully, the bloodwork results are in!
You realize that you are going to need a more detailed metabolic analysis of the woman's blood and heart tissue to support your hypothesis of her death. Thankfully, the bloodwork results are in! Click to add notes

Normal vs. Patient Blood Work
Blood Analysis (Cells or Metabolite) Normal Range Patient’s Levels White Blood cells(WBC) X 106/ml 7.1 X 106/ml Red Blood cells(WBC) X 106/ml 4.4 X 106/ml Lactate 0.5-1 mmol/L 7 mmol/L Cholesterol mg/dL 240 mg/dL Triglyceride mg/dL 297 mg/dL Calcium mg/dL 9.3 mg/dL Why? Let’s look deeper. Click to add notes *DATA SHEET INFO*

The answer to elevated lactate metabolite levels
is explained by the “Cori Cycle” ADP ATP Glucose Pyruvate NAD+ NADH NADH NAD+ Pyruvate Lactate Lactate (For Kreb’s Cycle) (TOXIC!) skeletal muscle liver 1. Muscle: Regeneration of NAD+ for Glycolysis (energy production). 2. Liver: Convert lactate back to pyruvate for Krebs Cycle. *DATA SHEET INFO*

Heart Tissue Metabolite Levels
What do these results affirm? Click to add notes *DATA SHEET INFO*

What does NAD+ (oxidized form) have to do with ATP synthesis?
Based on this data, what do you hypothesize caused the death of the woman? What does NAD+ (oxidized form) have to do with ATP synthesis? Would supplementation with NAD+ have helped save the woman’s life? Explain your reasoning. Click to add notes

Inhibitors (Poisons) that disable the Electron Transport System

QEP BIOLOGY MODULE DEVELOPMENT TEAM University of North Carolina
ACKNOWLEDGEMENTS QEP BIOLOGY MODULE DEVELOPMENT TEAM HCC Tineke Berends Terri Bubb Audrey Bush Nazanin Hebel Jennifer O’Neil Leena Sawant Pauline Ward University of North Carolina AT Baines Click to add notes

Concept 7.5: Fermentation and Anaerobic Respiration enables cells to produce ATP without the use of oxygen In the absence of O2, glycolysis couples with anaerobic respiration or fermentation to produce ATP Anaerobic respiration uses an electron transport chain with an electron acceptor other than O2, for example hydrogen sulfate (F-O-N-Cl-Br-I-S-C-H) Fermentation uses phosphorylation instead of an electron transport chain to generate ATP Cellular respiration produces 30 (eukaryote) or 32 (prokaryote) ATP per molecule of glucose; fermentation produces 2 ATP per glucose molecule

Animation: Fermentation Overview
Types of Fermentation Fermentation consists of glycolysis & reactions that regenerate NAD+, which can be reused by glycolysis over and over again In alcohol fermentation, pyruvate is converted to ethanol in two steps, with the first releasing CO2 In lactic acid fermentation, pyruvate is reduced to NADH, forming lactate as an end product, with no release of CO2 Animation: Fermentation Overview

Figure 9.18 Fermentation 2 ADP + 2 Pi 2 ATP Glucose Glycolysis
2 Pyruvate 2 NAD+ 2 NADH 2 CO2 + 2 H+ 2 Ethanol 2 Acetaldehyde (a) Alcohol fermentation 2 ADP + 2 Pi 2 ATP Glucose Glycolysis Figure 9.18 Fermentation 2 NAD+ 2 NADH + 2 H+ 2 Pyruvate 2 Lactate (b) Lactic acid fermentation

Fermentation Pyruvate is Oxidized: Pyruvate is Fermented:
Aerobic Conditions Anaerobic Conditions Pyruvate is Oxidized: Pyruvate is Fermented: make Acetyl – CoA Lactate (in muscle), Ethanol (in yeast) Why? Due to a lack of Oxygen, the Electron Transport System stops working. NADH is not oxidized back to NAD+. NAD is a obligate cofactor in glycolysis; without it, glycolysis would not proceed and the cell would not produce life-sustaining ATP. So how do we recycle? Cori Cycle ADP ATP Glucose Pyruvate NAD+ NADH NADH NAD+ Pyruvate Lactate Lactate skeletal muscle (For Kreb’s Cycle) (TOXIC!) liver 1. Muscle: Regeneration of NAD+ for Glycolysis (energy production). 2. Liver: Convert lactate back to pyruvate for Krebs Cycle.

Summary Glucose Glycolysis CYTOSOL Pyruvate O2 present: No O2 present:
Aerobic cellular respiration No O2 present: Fermentation MITOCHONDRION Ethanol or lactate Acetyl CoA Figure 9.19 Pyruvate as a key juncture in catabolism Citric acid cycle

Concept 7.6: Glycolysis and the Kreb’s cycle connect to many other metabolic pathways
Glycolysis and the Kreb’s cycle are major intersections to various catabolic and anabolic pathways Catabolic pathways funnel electrons from many kinds of organic molecules into cellular respiration Anabolic pathways use small molecules directly from food, from glycolysis, or from the citric acid cycle to build other substances

Catabolic Pathways are shown here… Anabolic Pathways
Proteins Carbohydrates Fats Catabolic Pathways are shown here… Anabolic Pathways run in the opposite direction… Amino acids Sugars Glycerol Fatty acids Glycolysis Glucose Glyceraldehyde-3- P NH3 Pyruvate Acetyl CoA Figure 9.20 The catabolism of various molecules from food Citric acid cycle Oxidative phosphorylation

Regulation of Cellular Respiration via Feedback Mechanisms
Feedback inhibition is the most common mechanism for control If ATP concentration begins to drop, respiration speeds up; when there is plenty of ATP, respiration slows down Control of catabolism is based mainly on regulating the activity of enzymes at strategic points in the catabolic pathway

Figure 9.21 The control of cellular respiration
Glucose AMP Glycolysis Fructose-6-phosphate Stimulates + Phosphofructokinase – – Fructose-1,6-bisphosphate Inhibits Inhibits Pyruvate ATP Citrate Acetyl CoA Figure 9.21 The control of cellular respiration Citric acid cycle Oxidative phosphorylation

An Overview of Cellular Respiration
Figure 7.6-s3 An overview of cellular respiration (step 3)

NADH NAD+ What is FAD  FADH2? *DATA SHEET INFO* NADH H+ NAD+ + 2[H] +
2 e– + 2 H+ 2 e– + H+ NADH H+ Dehydrogenase Reduction of NAD+ NAD+ + 2[H] + H+ Oxidation of NADH Nicotinamide (reduced form) NADH Nicotinamide (oxidized form) What is FAD  FADH2? Figure 9.4 NAD+ as an electron shuttle NAD+ *DATA SHEET INFO*

Tough, huh? So let’s think about each answer choice!
When electrons flow along the mitochondrial electron transport chain, which of the following changes occur? A) NAD+ is reduced to NADH. B) ATP synthase pumps protons by active transport. C) The electrons gain free energy. D) The cytochromes phosphorylate ADP to form ATP. E) The pH of the inter membrane space decreases. Tough, huh? So let’s think about each answer choice! Click to add notes

A) NAD+ is reduced to NADH.
NADH (from GPS) or MITOCHONDIAL MATRIX NADH NAD+ FADH2 O2 H+ H+ H+ ADP + P FAD+ NAD+ ATP Low [H+] H2O e- Ubiquinone Cyt C Reductase Inner Membrane (Cristae) ATP Synthase Cyt C Oxidase e- NADH dehydrogenase e- e- e- Coenzyme Q Cyt C H+ ~3 protons ~3 protons High [H+] ~3 protons Outer membrane When electrons flow along the mitochondrial electron transport chain, which of the following changes occur? A) NAD+ is reduced to NADH.

B) ATP synthase pumps protons by active transport.
NADH (from GPS) or MITOCHONDIAL MATRIX NADH NAD+ FADH2 O2 H+ H+ H+ ADP + P FAD+ NAD+ ATP Low [H+] H2O e- Ubiquinone Cyt C Reductase Inner Membrane (Cristae) ATP Synthase Cyt C Oxidase e- NADH dehydrogenase e- e- e- Coenzyme Q Cyt C H+ ~3 protons ~3 protons High [H+] ~3 protons Outer membrane When electrons flow along the mitochondrial electron transport chain, which of the following changes occur? B) ATP synthase pumps protons by active transport.

C) The electrons gain free energy.
NADH (from GPS) or MITOCHONDIAL MATRIX NADH NAD+ FADH2 O2 H+ H+ H+ ADP + P FAD+ NAD+ ATP Low [H+] H2O e- Ubiquinone Cyt C Reductase Inner Membrane (Cristae) ATP Synthase Cyt C Oxidase e- NADH dehydrogenase e- e- e- Coenzyme Q Cyt C H+ ~3 protons ~3 protons High [H+] ~3 protons Outer membrane When electrons flow along the mitochondrial electron transport chain, which of the following changes occur? C) The electrons gain free energy.

D) The cytochromes phosphorylate ADP to form ATP.
NADH (from GPS) or MITOCHONDIAL MATRIX NADH NAD+ FADH2 O2 H+ H+ H+ ADP + P FAD+ NAD+ ATP Low [H+] H2O e- Ubiquinone Cyt C Reductase Inner Membrane (Cristae) ATP Synthase Cyt C Oxidase e- NADH dehydrogenase e- e- e- Coenzyme Q Cyt C H+ ~3 protons ~3 protons High [H+] ~3 protons Outer membrane When electrons flow along the mitochondrial electron transport chain, which of the following changes occur? D) The cytochromes phosphorylate ADP to form ATP.

E) The pH of the inter membrane space decreases.
NADH (from GPS) or MITOCHONDIAL MATRIX NADH NAD+ FADH2 O2 H+ H+ H+ ADP + P FAD+ NAD+ ATP Low [H+] H2O e- Ubiquinone Cyt C Reductase Inner Membrane (Cristae) ATP Synthase Cyt C Oxidase e- NADH dehydrogenase e- e- e- Coenzyme Q Cyt C H+ ~3 protons ~3 protons High [H+] ~3 protons Outer membrane When electrons flow along the mitochondrial electron transport chain, which of the following changes occur? E) The pH of the inter membrane space decreases.

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