10/18/11 Chapter 9: Cellular Respiration. The Principle of Redox Chemical reactions that transfer electrons between reactants are called oxidation- reduction.

10/18/11

Chapter 9: Cellular Respiration

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)

Fig. 9-UN1

becomes oxidized(loses electron)

becomes reduced(gains electron)

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

Oxidation of Organic Fuel Molecules During Cellular

Respiration• During cellular respiration, the fuel (such as

glucose) is oxidized, and O2 is reduced:

Fig. 9-UN3

becomes oxidized

becomes reduced

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

• 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

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)

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

• The process that generates most of the ATP is called oxidative phosphorylation because it is 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 by substrate-level phosphorylation

Fig. 9-7

Enzyme

ADP

PSubstrate

Enzyme

ATP+

Product

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

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+

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

Fig. 9-9-2

Hexokinase

ATP

ADP

1

Phosphoglucoisomerase2

Phosphogluco-isomerase

2

Glucose

Glucose-6-phosphate

Fructose-6-phosphate

Glucose-6-phosphate


Glucose 6 phosphate is converted to its isomer fructose 6 phosphate by phosphoglucoisomerase

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



1

2

3


3

Phosphofructokinase transfers a phosphate group from ATP to the sugar investing another ATP, sugar is now ready to be split

Fig. 9-9-4

Glucose

ATP

ADP

Hexokinase

Glucose-6-phosphate

Phosphoglucoisomerase


ATP

ADP

Phosphofructokinase


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

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


NADH2

+ 2 H+

2 P i


6

2

2

Fig. 9-9-62 NAD+

NADH2


+ 2 H+

2 P i

2

2 ADP


Phosphoglycerokinase2 ATP

2 3-Phosphoglycerate

6

7

2

2 ADP

2 ATP


3-Phosphoglycerate

Phosphoglycero-kinase

2

7

Glycolysis produces some ATP by substrate-level phophorylation

Total of 2 ATP because there are 2 sugar molecules

Fig. 9-9-7

3-Phosphoglycerate


2 NAD+

2 NADH+ 2 H+

2 P i

2

2 ADP

Phosphoglycerokinase


2 ATP

3-Phosphoglycerate2

Phosphoglyceromutase

2-Phosphoglycerate2

2-Phosphoglycerate2

2

Phosphoglycero-mutase

6

7

8

8

Fig. 9-9-82 NAD+

NADH2

2

2

2

2

+ 2 H+

Triose phosphatedehydrogenase2 P i



2 ADP

2 ATP

3-Phosphoglycerate


Enolase

2-Phosphoglycerate

2 H2O

Phosphoenolpyruvate

9

8

7

6

2 2-Phosphoglycerate

Enolase

2

2 H2O

Phosphoenolpyruvate

9

Fig. 9-9-9


2 NAD+

NADH2

2

2

2

2

2

2 ADP

2 ATP

Pyruvate

Pyruvate kinase

Phosphoenolpyruvate

Enolase2 H2O

2-Phosphoglycerate


3-Phosphoglycerate


2 ATP

2 ADP


+ 2 H+

6

7

8

9

10

2

2 ADP

2 ATP

Phosphoenolpyruvate

Pyruvate kinase

2 Pyruvate

10

2 P i

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

Fig. 9-10

CYTOSOL MITOCHONDRION

NAD+ NADH + H+

2

1 3

Pyruvate

Transport protein

CO2Coenzyme A

Acetyl CoA

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

Fig. 9-12-1

Acetyl CoA

Oxaloacetate

CoA—SH

1

Citrate

Citricacidcycle

Fig. 9-12-2

Acetyl CoA

Oxaloacetate

Citrate

CoA—SH

Citricacidcycle

1

2

H2O

Isocitrate

Fig. 9-12-3

Acetyl CoA

CoA—SH

Oxaloacetate

Citrate

H2O

Citricacidcycle

Isocitrate

1

2

3

NAD+

NADH

+ H+

-Keto-glutarate

CO2

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

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

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

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

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

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

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–

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




• 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


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


Fig. 9-14

INTERMEMBRANE SPACE

Rotor

H+

Stator

Internalrod

Cata-lyticknob

ADP+P ATP

iMITOCHONDRIAL MATRIX

• 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

• 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


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

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


• 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


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

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

• 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


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

• 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


Fig. 9-19Glucose

Glycolysis

Pyruvate

CYTOSOL

No O2 present:Fermentation

O2 present:

Aerobic cellular respiration

MITOCHONDRION

Acetyl CoAEthanolor

lactateCitricacidcycle

10/18/11 Chapter 9: Cellular Respiration. The Principle of Redox Chemical reactions that transfer electrons between reactants are called oxidation- reduction.

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