10/18/11 chapter 9: cellular respiration. the principle of redox chemical reactions that transfer...
TRANSCRIPT
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10/18/11
Chapter 9: Cellular Respiration
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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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Fig. 9-UN1
becomes oxidized(loses electron)
becomes reduced(gains electron)
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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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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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Fig. 9-UN3
becomes oxidized
becomes 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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• 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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The Stages of Cellular Respiration: A Preview
• Cellular respiration 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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Fig. 9-6-3
Mitochondrion
Substrate-levelphosphorylation
ATP
Cytosol
Glucose Pyruvate
Glycolysis
Electronscarried
via NADH
Substrate-levelphosphorylation
ATP
Electrons carriedvia NADH and
FADH2
Oxidativephosphorylation
ATP
Citricacidcycle
Oxidativephosphorylation:electron transport
andchemiosmosis
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• The process that generates most of the ATP is called oxidative phosphorylation because it is powered by redox reactions
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• 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
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Fig. 9-7
Enzyme
ADP
PSubstrate
Enzyme
ATP+
Product
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Glycolysis
• 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
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Fig. 9-8
Energy investment phase
Glucose
2 ADP + 2 P 2 ATP used
formed4 ATP
Energy payoff phase
4 ADP + 4 P
2 NAD+ + 4 e– + 4 H+ 2 NADH + 2 H+
2 Pyruvate + 2 H2O
2 Pyruvate + 2 H2OGlucoseNet
4 ATP formed – 2 ATP used 2 ATP
2 NAD+ + 4 e– + 4 H+ 2 NADH + 2 H+
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Fig. 9-9-1
ATP
ADP
Hexokinase1
ATP
ADP
Hexokinase1
Glucose
Glucose-6-phosphate
Glucose
Glucose-6-phosphate
Glucose enters the cell and is phosphorylated by hexokinase , which transfers a phosphate group to glucose
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Fig. 9-9-2
Hexokinase
ATP
ADP
1
Phosphoglucoisomerase2
Phosphogluco-isomerase
2
Glucose
Glucose-6-phosphate
Fructose-6-phosphate
Glucose-6-phosphate
Fructose-6-phosphate
Glucose 6 phosphate is converted to its isomer fructose 6 phosphate by phosphoglucoisomerase
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1
Fig. 9-9-3
Hexokinase
ATP
ADP
Phosphoglucoisomerase
Phosphofructokinase
ATP
ADP
2
3
ATP
ADP
Phosphofructo-kinase
Fructose-1, 6-bisphosphate
Glucose
Glucose-6-phosphate
Fructose-6-phosphate
Fructose-1, 6-bisphosphate
1
2
3
Fructose-6-phosphate
3
Phosphofructokinase transfers a phosphate group from ATP to the sugar investing another ATP, sugar is now ready to be split
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Fig. 9-9-4
Glucose
ATP
ADP
Hexokinase
Glucose-6-phosphate
Phosphoglucoisomerase
Fructose-6-phosphate
ATP
ADP
Phosphofructokinase
Fructose-1, 6-bisphosphate
Aldolase
Isomerase
Dihydroxyacetonephosphate
Glyceraldehyde-3-phosphate
1
2
3
4
5
Aldolase
Isomerase
Fructose-1, 6-
bisphosphate
Dihydroxyacetonephosphate
Glyceraldehyde-
3-phosphate
4
5
Aldolase cleaves sugar into 2 3- carbon sugars
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Fig. 9-9-6
NADH2
+ 2 H+
2 P i
2
2 ADP
2 ATP
2
6
7
2
This enzyme:1.Oxidizes the sugar by transfer of electrons and H+ to NAD+ forming NADH2.Exergonic and enzyme uses energy to transfer phosphate to substrate
Fig. 9-9-52 NAD+
NADH2
+ 2 H+
2
2 P i
Triose phosphatedehydrogenase
1, 3-Bisphosphoglycerate
6
2 NAD+
Glyceraldehyde-3-phosphate
Triose phosphatedehydrogenase
NADH2
+ 2 H+
2 P i
1, 3-Bisphosphoglycerate
6
2
2
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Fig. 9-9-62 NAD+
NADH2
Triose phosphatedehydrogenase
+ 2 H+
2 P i
2
2 ADP
1, 3-Bisphosphoglycerate
Phosphoglycerokinase2 ATP
2 3-Phosphoglycerate
6
7
2
2 ADP
2 ATP
1, 3-Bisphosphoglycerate
3-Phosphoglycerate
Phosphoglycero-kinase
2
7
Glycolysis produces some ATP by substrate-level phophorylation
Total of 2 ATP because there are 2 sugar molecules
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Fig. 9-9-7
3-Phosphoglycerate
Triose phosphatedehydrogenase
2 NAD+
2 NADH+ 2 H+
2 P i
2
2 ADP
Phosphoglycerokinase
1, 3-Bisphosphoglycerate
2 ATP
3-Phosphoglycerate2
Phosphoglyceromutase
2-Phosphoglycerate2
2-Phosphoglycerate2
2
Phosphoglycero-mutase
6
7
8
8
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Fig. 9-9-82 NAD+
NADH2
2
2
2
2
+ 2 H+
Triose phosphatedehydrogenase2 P i
1, 3-Bisphosphoglycerate
Phosphoglycerokinase
2 ADP
2 ATP
3-Phosphoglycerate
Phosphoglyceromutase
Enolase
2-Phosphoglycerate
2 H2O
Phosphoenolpyruvate
9
8
7
6
2 2-Phosphoglycerate
Enolase
2
2 H2O
Phosphoenolpyruvate
9
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Fig. 9-9-9
Triose phosphatedehydrogenase
2 NAD+
NADH2
2
2
2
2
2
2 ADP
2 ATP
Pyruvate
Pyruvate kinase
Phosphoenolpyruvate
Enolase2 H2O
2-Phosphoglycerate
Phosphoglyceromutase
3-Phosphoglycerate
Phosphoglycerokinase
2 ATP
2 ADP
1, 3-Bisphosphoglycerate
+ 2 H+
6
7
8
9
10
2
2 ADP
2 ATP
Phosphoenolpyruvate
Pyruvate kinase
2 Pyruvate
10
2 P i
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If Oxygen is present…
• In the presence of O2, pyruvate enters the mitochondrion
• Before the citric acid cycle can begin, pyruvate must be converted to acetyl CoA, which links the cycle to glycolysis
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Fig. 9-10
CYTOSOL MITOCHONDRION
NAD+ NADH + H+
2
1 3
Pyruvate
Transport protein
CO2Coenzyme A
Acetyl CoA
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Fig. 9-11
Pyruvate
NAD+
NADH
+ H+Acetyl CoA
CO2
CoA
CoA
CoA
Citricacidcycle
FADH2
FAD
CO22
3
3 NAD+
+ 3 H+
ADP + P i
ATP
NADH
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Fig. 9-12-1
Acetyl CoA
Oxaloacetate
CoA—SH
1
Citrate
Citricacidcycle
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Fig. 9-12-2
Acetyl CoA
Oxaloacetate
Citrate
CoA—SH
Citricacidcycle
1
2
H2O
Isocitrate
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Fig. 9-12-3
Acetyl CoA
CoA—SH
Oxaloacetate
Citrate
H2O
Citricacidcycle
Isocitrate
1
2
3
NAD+
NADH
+ H+
-Keto-glutarate
CO2
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Fig. 9-12-4
Acetyl CoA
CoA—SH
Oxaloacetate
Citrate
H2O
IsocitrateNAD+
NADH
+ H+
Citricacidcycle
-Keto-glutarate
CoA—SH
1
2
3
4
NAD+
NADH
+ H+SuccinylCoA
CO2
CO2
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Fig. 9-12-5
Acetyl CoA
CoA—SH
Oxaloacetate
Citrate
H2O
IsocitrateNAD+
NADH
+ H+
CO2
Citricacidcycle
CoA—SH -Keto-
glutarate
CO2NAD+
NADH
+ H+SuccinylCoA
1
2
3
4
5
CoA—SH
GTP GDP
ADP
P iSuccinate
ATP
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Fig. 9-12-6
Acetyl CoA
CoA—SH
Oxaloacetate
H2O
CitrateIsocitrate
NAD+
NADH
+ H+
CO2
Citricacidcycle
CoA—SH -Keto-
glutarate
CO2NAD+
NADH
+ H+
CoA—SH
P
SuccinylCoA
i
GTP GDP
ADP
ATP
Succinate
FAD
FADH2
Fumarate
1
2
3
4
5
6
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Fig. 9-12-7
Acetyl CoA
CoA—SH
Oxaloacetate
Citrate
H2O
IsocitrateNAD+
NADH
+ H+
CO2
-Keto-glutarate
CoA—SH
NAD+
NADH
SuccinylCoA
CoA—SH
PP
GDPGTP
ADP
ATP
Succinate
FAD
FADH2
Fumarate
CitricacidcycleH2O
Malate
1
2
5
6
7
i
CO2
+ H+
3
4
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Fig. 9-12-8
Acetyl CoA
CoA—SH
Citrate
H2O
IsocitrateNAD+
NADH
+ H+
CO2
-Keto-glutarate
CoA—SH
CO2NAD+
NADH
+ H+SuccinylCoA
CoA—SH
P i
GTP GDP
ADP
ATP
Succinate
FAD
FADH2
Fumarate
CitricacidcycleH2O
Malate
Oxaloacetate
NADH
+H+
NAD+
1
2
3
4
5
6
7
8
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The Pathway of Electron Transport
• The electron transport chain is in the cristae of the mitochondrion
• Most of the chain’s components are proteins and multiprotein complexes
• Electrons release energy as they go down the chain and are finally passed to O2, forming H2O
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
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Fig. 9-13
NADH
NAD+2FADH2
2 FADMultiproteincomplexesFAD
Fe•S
FMN
Fe•S
Q
Fe•S
Cyt b
Cyt c1
Cyt c
Cyt a
Cyt a3
IV
Fre
e en
erg
y (G
) r e
lat i
ve t
o O
2 (
kcal
/mo
l)
50
40
30
20
10 2
(from NADHor FADH2)
0 2 H+ + 1/2 O2
H2O
e–
e–
e–
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Fig. 9-16
Protein complexof electroncarriers
H+
H+H+
Cyt c
Q
V
FADH2 FAD
NAD+NADH
(carrying electronsfrom food)
Electron transport chain
2 H+ + 1/2O2H2O
ADP + P i
Chemiosmosis
Oxidative phosphorylation
H+
H+
ATP synthase
ATP
21
http://www.youtube.com/watch?v=3y1dO4nNaKY
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• Electrons are transferred from NADH or FADH2 to the electron transport chain
• Electrons are passed through a number of proteins including cytochromes to O2
• The electron transport chain generates no ATP
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
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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 channels in ATP synthase
• ATP synthase uses the 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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
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Fig. 9-14
INTERMEMBRANE SPACE
Rotor
H+
Stator
Internalrod
Cata-lyticknob
ADP+P ATP
iMITOCHONDRIAL MATRIX
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• 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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
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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 40% of the energy in a glucose molecule is transferred to ATP during cellular respiration, making about 38 ATP
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
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Fig. 9-17
Maximum per glucose: About36 or 38 ATP
+ 2 ATP+ 2 ATP + about 32 or 34 ATP
Oxidativephosphorylation:electron transport
andchemiosmosis
Citricacidcycle
2AcetylCoA
Glycolysis
Glucose2
Pyruvate
2 NADH 2 NADH 6 NADH 2 FADH2
2 FADH2
2 NADHCYTOSOL Electron shuttles
span membrane
or
MITOCHONDRION
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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
• Glycolysis can produce ATP with or without O2 (in aerobic or anaerobic conditions)
• In the absence of O2, glycolysis couples with fermentation or anaerobic respiration to produce ATP
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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
Animation: Fermentation OverviewAnimation: Fermentation Overview
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Fig. 9-18a
2 ADP + 2 P i 2 ATP
Glucose Glycolysis
2 Pyruvate
2 NADH2 NAD+
+ 2 H+CO2
2 Acetaldehyde2 Ethanol
(a) Alcohol fermentation
2
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Fig. 9-18
2 ADP + 2 Pi 2 ATP
Glucose Glycolysis
2 NAD+ 2 NADH
2 Pyruvate
+ 2 H+
2 Acetaldehyde2 Ethanol
(a) Alcohol fermentation
2 ADP + 2 Pi2 ATP
Glucose Glycolysis
2 NAD+ 2 NADH+ 2 H+
2 Pyruvate
2 Lactate
(b) Lactic acid fermentation
2 CO2
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• 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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Fig. 9-18b
Glucose
2 ADP + 2 P i 2 ATP
Glycolysis
2 NAD+ 2 NADH+ 2 H+
2 Pyruvate
2 Lactate
(b) Lactic acid fermentation
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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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Fig. 9-19Glucose
Glycolysis
Pyruvate
CYTOSOL
No O2 present:Fermentation
O2 present:
Aerobic cellular respiration
MITOCHONDRION
Acetyl CoAEthanolor
lactateCitricacidcycle