Download - Ch 9: Cell Respiration and Fermentation
LECTURE PRESENTATIONSFor CAMPBELL BIOLOGY, NINTH EDITION
Jane B. Reece, Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson
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Lectures byErin Barley
Kathleen Fitzpatrick
Cellular Respiration and Fermentation
Chapter 9
Overview: Life Is Work
• Living cells require energy from outside sources
• Some animals, such as the chimpanzee, obtain energy by eating plants, and some animals feed on other organisms that eat plants
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• 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
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Figure 9.2
Lightenergy
ECOSYSTEM
Photosynthesisin chloroplasts
Cellular respirationin mitochondria
CO2 + H2O + O2Organic
molecules
ATP powersmost cellular workATP
Heatenergy
Concept 9.1: Catabolic pathways yield energy by oxidizing organic fuels
• Several processes are central to cellular respiration and related pathways
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Catabolic Pathways and Production of ATP
• 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
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• Cellular respiration includes both aerobic and anaerobic respiration but is often used to refer to aerobic respiration
• 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)
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Redox Reactions: Oxidation and Reduction
• The transfer of electrons during chemical reactions releases energy stored in organic molecules
• This released energy is ultimately used to synthesize ATP
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The Principle of Redox
• 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)
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• The electron donor is called the reducing agent
• The electron receptor is called the oxidizing agent
• Some redox reactions do not transfer electrons but change the electron sharing in covalent bonds
• An example is the reaction between methane and O2
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Figure 9.3
Reactants Products
Energy
WaterCarbon dioxideMethane(reducing
agent)
Oxygen(oxidizing
agent)
becomes oxidized
becomes reduced
Oxidation of Organic Fuel Molecules During Cellular Respiration
• During cellular respiration, the fuel (such as glucose) is oxidized, and O2 is reduced
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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
• As an electron acceptor, NAD+ functions as an oxidizing agent during cellular respiration
• Each NADH (the reduced form of NAD+) represents stored energy that is tapped to synthesize ATP
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Figure 9.4
Nicotinamide(oxidized form)
NAD+
(from food)
Dehydrogenase
Reduction of NAD+
Oxidation of NADH
Nicotinamide(reduced form)
NADH
• NADH passes the electrons to the electron transport chain
• Unlike an uncontrolled reaction, the electron transport chain passes electrons in a series of steps instead of one explosive reaction
• O2 pulls electrons down the chain in an energy-yielding tumble
• The energy yielded is used to regenerate ATP
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Figure 9.5
(a) Uncontrolled reaction (b) Cellular respiration
Explosiverelease of
heat and lightenergy
Controlledrelease ofenergy for
synthesis ofATP
Fre
e en
erg
y, G
Fre
e en
erg
y, G
H2 + 1/2 O2 2 H + 1/2 O2
1/2 O2
H2O H2O
2 H+ + 2 e−
2 e−
2 H+
ATP
ATP
ATP
Electro
n tran
spo
rt
chain
(from food via NADH)
The Stages of Cellular Respiration: A Preview
• Harvesting of energy from glucose has three stages
– Glycolysis (breaks down glucose into two molecules of pyruvate)
– The citric acid cycle (completes the breakdown of glucose)
– Oxidative phosphorylation (accounts for most of the ATP synthesis)
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Figure 9.UN05
Glycolysis (color-coded teal throughout the chapter)1.
Pyruvate oxidation and the citric acid cycle(color-coded salmon)
2.
Oxidative phosphorylation: electron transport andchemiosmosis (color-coded violet)
3.
Figure 9.6-1
Electronscarried
via NADH
Glycolysis
Glucose Pyruvate
CYTOSOL MITOCHONDRION
ATP
Substrate-levelphosphorylation
Figure 9.6-2
Electronscarried
via NADH
Electrons carriedvia NADH and
FADH2
Citricacidcycle
Pyruvateoxidation
Acetyl CoA
Glycolysis
Glucose Pyruvate
CYTOSOL MITOCHONDRION
ATP ATP
Substrate-levelphosphorylation
Substrate-levelphosphorylation
Figure 9.6-3
Electronscarried
via NADH
Electrons carriedvia NADH and
FADH2
Citricacidcycle
Pyruvateoxidation
Acetyl CoA
Glycolysis
Glucose Pyruvate
Oxidativephosphorylation:electron transport
andchemiosmosis
CYTOSOL MITOCHONDRION
ATP ATP ATP
Substrate-levelphosphorylation
Substrate-levelphosphorylation
Oxidative phosphorylation
• The process that generates most of the ATP is called oxidative phosphorylation because it is powered by redox reactions
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BioFlix: Cellular Respiration
• 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 by substrate-level phosphorylation
• For each molecule of glucose degraded to CO2
and water by respiration, the cell makes up to 32 molecules of ATP
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Concept 9.2: Glycolysis harvests chemical energy by oxidizing glucose to pyruvate
• Glycolysis (“splitting of sugar”) breaks down glucose into two molecules of pyruvate
• Glycolysis occurs in the cytoplasm and has two major phases
– Energy investment phase– Energy payoff phase
• Glycolysis occurs whether or not O2 is present
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Figure 9.8
Energy Investment Phase
Glucose
2 ADP + 2 P
4 ADP + 4 P
Energy Payoff Phase
2 NAD+ + 4 e− + 4 H+
2 Pyruvate + 2 H2O
2 ATP used
4 ATP formed
2 NADH + 2 H+
NetGlucose 2 Pyruvate + 2 H2O
2 ATP
2 NADH + 2 H+ 2 NAD+ + 4 e− + 4 H+
4 ATP formed − 2 ATP used
Figure 9.9-2
Glycolysis: Energy Investment Phase
ATPGlucose Glucose 6-phosphate Fructose 6-phosphate
ADP
Hexokinase Phosphogluco-isomerase
12
Figure 9.9-3
Glycolysis: Energy Investment Phase
ATP ATPGlucose Glucose 6-phosphate Fructose 6-phosphate Fructose 1,6-bisphosphate
ADP ADP
Hexokinase Phosphogluco-isomerase
Phospho-fructokinase
12 3
Figure 9.9-4
Glycolysis: Energy Investment Phase
ATP ATPGlucose Glucose 6-phosphate Fructose 6-phosphate Fructose 1,6-bisphosphate
Dihydroxyacetonephosphate
Glyceraldehyde3-phosphate
Tostep 6
ADP ADP
Hexokinase Phosphogluco-isomerase
Phospho-fructokinase
Aldolase
Isomerase
12 3 4
5
Figure 9.9-5
Glycolysis: Energy Payoff Phase
2 NADH
2 NAD+ + 2 H+
2 P i
1,3-Bisphospho-glycerate6
Triosephosphate
dehydrogenase
Figure 9.9-6
Glycolysis: Energy Payoff Phase
2 ATP2 NADH
2 NAD+ + 2 H+
2 P i
2 ADP
1,3-Bisphospho-glycerate
3-Phospho-glycerate
2
Phospho-glycerokinase
67
Triosephosphate
dehydrogenase
Figure 9.9-7
Glycolysis: Energy Payoff Phase
2 ATP2 NADH
2 NAD+ + 2 H+
2 P i
2 ADP
1,3-Bisphospho-glycerate
3-Phospho-glycerate
2-Phospho-glycerate
2 2
Phospho-glycerokinase
Phospho-glyceromutase
67 8
Triosephosphate
dehydrogenase
Figure 9.9-8
Glycolysis: Energy Payoff Phase
2 ATP2 NADH
2 NAD+ + 2 H+
2 P i
2 ADP
1,3-Bisphospho-glycerate
3-Phospho-glycerate
2-Phospho-glycerate
Phosphoenol-pyruvate (PEP)
2 2 2
2 H2O
Phospho-glycerokinase
Phospho-glyceromutase
Enolase
67 8
9
Triosephosphate
dehydrogenase
Figure 9.9-9
Glycolysis: Energy Payoff Phase
2 ATP 2 ATP2 NADH
2 NAD+ + 2 H+
2 P i
2 ADP
1,3-Bisphospho-glycerate
3-Phospho-glycerate
2-Phospho-glycerate
Phosphoenol-pyruvate (PEP)
Pyruvate
2 ADP2 2 2
2 H2O
Phospho-glycerokinase
Phospho-glyceromutase
Enolase Pyruvatekinase
67 8
910
Triosephosphate
dehydrogenase
Figure 9.9a
Glycolysis: Energy Investment Phase
ATPGlucose Glucose 6-phosphate
ADP
Hexokinase
1
Fructose 6-phosphate
Phosphogluco-isomerase
2
Figure 9.9b
Glycolysis: Energy Investment Phase
ATPFructose 6-phosphate
ADP
3
Fructose 1,6-bisphosphate
Phospho-fructokinase
4
5
Aldolase
Dihydroxyacetonephosphate
Glyceraldehyde3-phosphate
Tostep 6Isomerase
Figure 9.9c
Glycolysis: Energy Payoff Phase
2 NADH2 ATP
2 ADP 2
2
2 NAD+ + 2 H+
2 P i
3-Phospho-glycerate
1,3-Bisphospho-glycerate
Triosephosphate
dehydrogenase
Phospho-glycerokinase
67
Figure 9.9d
Glycolysis: Energy Payoff Phase
2 ATP
2 ADP2222
2 H2O
PyruvatePhosphoenol-pyruvate (PEP)
2-Phospho-glycerate
3-Phospho-glycerate
89
10
Phospho-glyceromutase
Enolase Pyruvatekinase
Concept 9.3: After pyruvate is oxidized, the citric acid cycle completes the energy-yielding oxidation of organic molecules
• In the presence of O2, pyruvate enters the mitochondrion (in eukaryotic cells) where the oxidation of glucose is completed
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Oxidation of Pyruvate to Acetyl CoA
• Before the citric acid cycle can begin, pyruvate must be converted to acetyl Coenzyme A (acetyl CoA), which links glycolysis to the citric acid cycle
• This step is carried out by a multienzyme complex that catalyses three reactions
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Figure 9.10
Pyruvate
Transport protein
CYTOSOL
MITOCHONDRION
CO2 Coenzyme A
NAD+ + H+NADH Acetyl CoA
1
2
3
• The citric acid cycle, also called the Krebs cycle, completes the break down of pyruvate to CO2
• The cycle oxidizes organic fuel derived from pyruvate, generating 1 ATP, 3 NADH, and 1 FADH2 per turn
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The Citric Acid Cycle
Figure 9.11Pyruvate
NAD+
NADH
+ H+Acetyl CoA
CO2
CoA
CoA
CoA
2 CO2
ADP + P i
FADH2
FAD
ATP
3 NADH
3 NAD+
Citricacidcycle
+ 3 H+
• The citric acid cycle has eight steps, each catalyzed by a specific enzyme
• The 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
• The NADH and FADH2 produced by the cycle relay electrons extracted from food to the electron transport chain
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Figure 9.12-3
1
Acetyl CoA
CitrateIsocitrate
α-Ketoglutarate
Citricacidcycle
NADH+ H+
NAD+
H2O
3
2
CoA-SH
CO2
Oxaloacetate
Figure 9.12-4
1
Acetyl CoA
CitrateIsocitrate
α-Ketoglutarate
SuccinylCoA
Citricacidcycle
NADH
NADH
+ H+
+ H+
NAD+
NAD+
H2O
3
2
4
CoA-SH
CO2
CoA-SH
CO2
Oxaloacetate
Figure 9.12-5
1
Acetyl CoA
CitrateIsocitrate
α-Ketoglutarate
SuccinylCoA
Succinate
Citricacidcycle
NADH
NADH
ATP
+ H+
+ H+
NAD+
NAD+
H2O
ADP
GTP GDP
P i
3
2
4
5
CoA-SH
CO2
CoA-SH
CoA-SH
CO2
Oxaloacetate
Figure 9.12-6
1
Acetyl CoA
CitrateIsocitrate
α-Ketoglutarate
SuccinylCoA
Succinate
Fumarate
Citricacidcycle
NADH
NADH
FADH2
ATP
+ H+
+ H+
NAD+
NAD+
H2O
ADP
GTP GDP
P i
FAD
3
2
4
5
6
CoA-SH
CO2
CoA-SH
CoA-SH
CO2
Oxaloacetate
Figure 9.12-7
1
Acetyl CoA
CitrateIsocitrate
α-Ketoglutarate
SuccinylCoA
Succinate
Fumarate
Malate
Citricacidcycle
NADH
NADH
FADH2
ATP
+ H+
+ H+
NAD+
NAD+
H2O
H2O
ADP
GTP GDP
P i
FAD
3
2
4
5
6
7
CoA-SH
CO2
CoA-SH
CoA-SH
CO2
Oxaloacetate
Figure 9.12-8
NADH
1
Acetyl CoA
CitrateIsocitrate
α-Ketoglutarate
SuccinylCoA
Succinate
Fumarate
Malate
Citricacidcycle
NAD+
NADH
NADH
FADH2
ATP
+ H+
+ H+
+ H+
NAD+
NAD+
H2O
H2O
ADP
GTP GDP
P i
FAD
3
2
4
5
6
7
8
CoA-SH
CO2
CoA-SH
CoA-SH
CO2
Oxaloacetate
Figure 9.12b
Isocitrate
α-Ketoglutarate
SuccinylCoA
NADH
NADH
NAD+
NAD+
+ H+
CoA-SH
CO2
CO2
3
4
+ H+
Concept 9.4: During oxidative phosphorylation, chemiosmosis 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, which powers ATP synthesis via oxidative phosphorylation
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The Pathway of Electron Transport
• The electron transport chain is in the inner membrane (cristae) of the mitochondrion
• Most of the chain’s components are proteins, which exist in multiprotein complexes
• The 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
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Figure 9.13
NADH
FADH2
2 H+ + 1/2 O2
2 e−
2 e−
2 e−
H2O
NAD+
Multiproteincomplexes
(originally from NADH or FADH2)
III
III
IV
50
40
30
20
10
0
Fre
e en
erg
y (G
) re
lati
ve t
o O
2 (k
cal/
mo
l)
FMN
Fe•S Fe•S
FAD
Q
Cyt b
Cyt c1
Cyt c
Cyt a
Cyt a3
Fe•S
• Electrons are transferred from NADH or FADH2 to the electron transport chain
• Electrons are passed through a number of proteins including cytochromes (each with an iron atom) to O2
• The electron transport chain generates no ATP directly
• It breaks the large free-energy drop from food to O2 into smaller steps that release energy in manageable amounts
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Chemiosmosis: The 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 the proton, ATP synthase
• ATP synthase uses the exergonic flow of H+ to drive phosphorylation of ATP
• This is an example of chemiosmosis, the use of energy in a H+ gradient to drive cellular work
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Figure 9.14INTERMEMBRANE SPACE
Rotor
StatorH+
Internalrod
Catalyticknob
ADP+P i ATP
MITOCHONDRIAL MATRIX
Figure 9.15
Proteincomplexof electroncarriers
(carrying electronsfrom food)
Electron transport chain
Oxidative phosphorylation
Chemiosmosis
ATPsynth-ase
I
II
III
IVQ
Cyt c
FADFADH2
NADH ADP + P i
NAD+
H+
2 H+ + 1/2O2
H+
H+H+
21
H+
H2O
ATP
• 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
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An Accounting of ATP Production by Cellular Respiration
• During cellular respiration, most energy flows in this sequence:
glucose → NADH → electron transport chain → proton-motive force → ATP
• About 34% of the energy in a glucose molecule is transferred to ATP during cellular respiration, making about 32 ATP
• There are several reasons why the number of ATP is not known exactly
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Figure 9.16
Electron shuttlesspan membrane
MITOCHONDRION2 NADH
2 NADH 2 NADH 6 NADH
2 FADH2
2 FADH2
or
+ 2 ATP+ 2 ATP + about 26 or 28 ATP
Glycolysis
Glucose 2 Pyruvate
Pyruvate oxidation
2 Acetyl CoACitricacidcycle
Oxidativephosphorylation:electron transport
andchemiosmosis
CYTOSOL
Maximum per glucose:About
30 or 32 ATP
Concept 9.5: Fermentation and anaerobic respiration enable cells to produce ATP without the use of oxygen
• Most cellular respiration requires O2 to produce ATP
• Without O2, the electron transport chain will cease to operate
• In that case, glycolysis couples with fermentation or anaerobic respiration to produce ATP
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• Anaerobic respiration uses an electron transport chain with a final electron acceptor other than O2, for example sulfate
• Fermentation uses substrate-level phosphorylation instead of an electron transport chain to generate ATP
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Types of Fermentation
• Fermentation consists of glycolysis plus reactions that regenerate NAD+, which can be reused by glycolysis
• Two common types are alcohol fermentation and lactic acid fermentation
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• In alcohol fermentation, pyruvate is converted to ethanol in two steps, with the first releasing CO2
• Alcohol fermentation by yeast is used in brewing, winemaking, and baking
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Animation: Fermentation Overview
Figure 9.17
2 ADP 2 ATP
Glucose Glycolysis
2 Pyruvate
2 CO22
+2 NADH
2 Ethanol 2 Acetaldehyde
(a) Alcohol fermentation (b) Lactic acid fermentation
2 Lactate
2 Pyruvate
2 NADH
Glucose Glycolysis
2 ATP2 ADP+ 2 Pi
NAD
2 H+
+ 2 Pi
2 NAD ++
+ 2 H+
2 ADP + 2 P i 2 ATP
Glucose Glycolysis
2 Pyruvate
2 CO22 NAD
+2 NADH
2 Ethanol 2 Acetaldehyde
(a) Alcohol fermentation
+
2 H+
Figure 9.17a
• In lactic acid fermentation, pyruvate is reduced to NADH, forming lactate as an end product, with no release of CO2
• Lactic acid fermentation by some fungi and bacteria is used to make cheese and yogurt
• Human muscle cells use lactic acid fermentation to generate ATP when O2 is scarce
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(b) Lactic acid fermentation
2 Lactate
2 Pyruvate
2 NADH
Glucose Glycolysis
2 ADP + 2 P i 2 ATP
2 NAD+
+
2 H+
Figure 9.17b
Comparing Fermentation with Anaerobic and Aerobic Respiration
• All use glycolysis (net ATP = 2) to oxidize glucose and harvest chemical energy of food
• In all three, NAD+ is the oxidizing agent that accepts electrons during glycolysis
• The processes have different final electron acceptors: an organic molecule (such as pyruvate or acetaldehyde) in fermentation and O2 in cellular respiration
• Cellular respiration produces 32 ATP per glucose molecule; fermentation produces 2 ATP per glucose molecule
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• Obligate anaerobes carry out fermentation or anaerobic respiration and cannot survive in the presence of O2
• Yeast and many bacteria are facultative anaerobes, meaning that they can survive using either fermentation or cellular respiration
• In a facultative anaerobe, pyruvate is a fork in the metabolic road that leads to two alternative catabolic routes
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Figure 9.18Glucose
CYTOSOLGlycolysis
Pyruvate
No O2 present:Fermentation
O2 present: Aerobic cellular respiration
Ethanol,lactate, or
other products
Acetyl CoA
MITOCHONDRION
Citricacidcycle
The Evolutionary Significance of Glycolysis
• Ancient prokaryotes are thought to have used glycolysis long before there was oxygen in the atmosphere
• Very little O2 was available in the atmosphere until about 2.7 billion years ago, so early prokaryotes likely used only glycolysis to generate ATP
• Glycolysis is a very ancient process
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Concept 9.6: Glycolysis and the citric acid cycle connect to many other metabolic pathways
• Gycolysis and the citric acid cycle are major intersections to various catabolic and anabolic pathways
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The Versatility of Catabolism
• Catabolic pathways funnel electrons from many kinds of organic molecules into cellular respiration
• Glycolysis accepts a wide range of carbohydrates
• Proteins must be digested to amino acids; amino groups can feed glycolysis or the citric acid cycle
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• Fats are digested to glycerol (used in glycolysis) and fatty acids (used in generating acetyl CoA)
• Fatty acids are broken down by beta oxidation and yield acetyl CoA
• An oxidized gram of fat produces more than twice as much ATP as an oxidized gram of carbohydrate
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Figure 9.19CarbohydratesProteins
Fattyacids
Aminoacids
Sugars
Fats
Glycerol
Glycolysis
Glucose
Glyceraldehyde 3- P
NH3 Pyruvate
Acetyl CoA
Citricacidcycle
Oxidativephosphorylation
Biosynthesis (Anabolic Pathways)
• The body uses small molecules to build other substances
• These small molecules may come directly from food, from glycolysis, or from the citric acid cycle
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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
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Figure 9.20
Phosphofructokinase
Glucose
GlycolysisAMP
Stimulates
− −
+Fructose 6-phosphate
Fructose 1,6-bisphosphate
Pyruvate
Inhibits Inhibits
ATP Citrate
Citricacidcycle
Oxidativephosphorylation
Acetyl CoA
Figure 9.UN07
Inputs Outputs
2 Pyruvate 2 Acetyl CoA
2 OxaloacetateCitricacidcycle
2
26
8ATP NADH
FADH2CO2
Figure 9.UN08
Protein complexof electroncarriers
(carrying electrons from food)
INTERMEMBRANESPACE
MITOCHONDRIAL MATRIX
H+
H+H+
2 H+ + 1/2 O2 H2O
NAD+
FADH2 FAD
Q
NADH
I
II
III
IV
Cyt c