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CAMPBELL BIOLOGY IN FOCUS © 2016 Pearson Education, Inc. URRY CAIN WASSERMAN MINORSKY REECE Lecture Presentations by Kathleen Fitzpatrick and Nicole Tunbridge, Simon Fraser University SECOND EDITION 7 Cellular Respiration and Fermentation
Transcript
Page 1: URRY &$,1 :$66(50$1 MINORSKY 5((&( - anderson.k12.ky.us 1.pdfLife Is Work Living cells require energy from outside sources Some animals, such as the giraffe, obtain energy by eating

CAMPBELL BIOLOGY IN FOCUS

© 2016 Pearson Education, Inc.

URRY • CAIN • WASSERMAN • MINORSKY • REECE

Lecture Presentations by

Kathleen Fitzpatrick and

Nicole Tunbridge,

Simon Fraser University

SECOND EDITION

7 Cellular

Respiration and

Fermentation

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Life Is Work

Living cells require energy from outside sources

Some animals, such as the giraffe, obtain energy by

eating plants, and some animals feed on other

organisms that eat plants

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Figure 7.1

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Energy flows into an ecosystem as sunlight and

leaves as heat

Photosynthesis generates O2 and organic

molecules, which are used as fuel for cellular

respiration

Cells use chemical energy stored in organic

molecules to regenerate ATP, which powers work

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Animation: Carbon Cycle

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Figure 7.2

© 2016 Pearson Education, Inc.

Light energy

ECOSYSTEM

Photosynthesis in chloroplasts

CO2 + H2O Organic

molecules + O2

Cellular respiration in mitochondria

ATP

Heat energy

ATP powers most cellular work

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Concept 7.1: Catabolic pathways yield energy by oxidizing organic fuels

Catabolic pathways involving electron transfer are

central processes to cellular respiration

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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 processes 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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Figure 7.UN01

© 2016 Pearson Education, Inc.

becomes oxidized (loses electron)

becomes reduced (gains electron)

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Figure 7.UN02

© 2016 Pearson Education, Inc.

becomes oxidized

becomes reduced

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The electron donor is called the reducing agent

The electron acceptor 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 7.3

© 2016 Pearson Education, Inc.

Reactants Products

Carbon dioxide Water Methane (reducing

agent)

Oxygen (oxidizing

agent)

becomes oxidized

becomes reduced

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Redox reactions that move electrons closer to

electronegative atoms, like oxygen, release

chemical energy that can be put to work

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Oxidation of Organic Fuel Molecules During Cellular Respiration

During cellular respiration, fuel (such as glucose) is

oxidized, and O2 is reduced

Organic molecules with an abundance of hydrogen,

like carbohydrates and fats, are excellent fuels

As hydrogen (with its electron) is transferred to

oxygen, energy is released that can be used in ATP

synthesis

© 2016 Pearson Education, Inc.

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Figure 7.UN03

© 2016 Pearson Education, Inc.

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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Enzymes called dehydrogenases facilitate the

transfer of two electrons and one hydrogen ion to

NAD+

One hydrogen ion is released in this process

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Figure 7.4

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2 e + 2 H+

NAD+ 2 e + H+

NADH Dehydrogenase

2[H] (from food)

Nicotinamide Nicotinamide (reduced form) (oxidized form)

H+

H+ Oxidation of NADH

Reduction of NAD+

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Figure 7.4-1

© 2016 Pearson Education, Inc.

NAD+

Nicotinamide (oxidized form)

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Figure 7.4-2

© 2016 Pearson Education, Inc.

2 e + 2 H+

Dehydrogenase

2 e + H+ NADH H+

2[H] (from food)

H+

Nicotinamide (reduced form)

Reduction of NAD+

Oxidation of NADH

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Figure 7.UN04

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Dehydrogenase

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NADH passes the electrons to the electron

transport chain

Electrons are passed to increasingly

electronegative carrier molecules down the chain

through a series of redox reactions

Electron transfer to oxygen occurs in a series of

energy-releasing steps instead of one explosive

reaction

The energy yielded is used to regenerate ATP

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Figure 7.5

© 2016 Pearson Education, Inc.

H2 + 1/2 O2 2 H +

Controlled release of

energy

ATP

ATP

ATP

2 e

2 H+

H2O

1 /2 O2

2 H+ + 2 e

Explosive release

H2O

(a) Uncontrolled reaction (b) Cellular respiration

Fre

e e

nerg

y,

G

Fre

e e

nerg

y,

G

1/2 O2

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The Stages of Cellular Respiration: A Preview

Harvesting of energy from glucose has three stages

Glycolysis breaks down glucose into two molecules

of pyruvate in the cytosol

Pyruvate oxidation and the citric acid cycle

completes the breakdown of glucose in the

mitochondrial matrix

Oxidative phosphorylation accounts for most of the

ATP synthesis and occurs in the inner membrane of

the mitochondria

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Figure 7.UN05

© 2016 Pearson Education, Inc.

1. GLYCOLYSIS (color-coded blue throughout the chapter)

2. PYRUVATE OXIDATION and the CITRIC ACID CYCLE

(color-coded orange)

3. OXIDATIVE PHOSPHORYLATION: Electron transport and

chemiosmosis (color-coded purple)

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Figure 7.6-s1

© 2016 Pearson Education, Inc.

Electrons via NADH

GLYCOLYSIS

Glucose Pyruvate

CYTOSOL MITOCHONDRION

ATP

Substrate-level

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Figure 7.6-s2

© 2016 Pearson Education, Inc.

Electrons via NADH

GLYCOLYSIS

Glucose Pyruvate

PYRUVATE OXIDATION

Acetyl CoA

CITRIC ACID

CYCLE

CYTOSOL MITOCHONDRION

ATP

Substrate-level

ATP

Substrate-level

Electrons via NADH and FADH2

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Figure 7.6-s3

© 2016 Pearson Education, Inc.

Electrons via NADH

GLYCOLYSIS

Glucose Pyruvate

PYRUVATE OXIDATION

Acetyl CoA

CITRIC ACID

CYCLE

CYTOSOL MITOCHONDRION

ATP

Substrate-level

ATP

Substrate-level

OXIDATIVE PHOSPHORYLATION

(Electron transport and chemiosmosis)

Electrons via NADH and FADH2

ATP

Oxidative

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Oxidative phosphorylation accounts for almost 90%

of the ATP generated by cellular respiration

This process involves the transfer of inorganic

phosphates to ADP

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A smaller amount of ATP is formed in glycolysis and

the citric acid cycle by substrate-level

phosphorylation

In this process, an enzyme transfers a phosphate

group directly from a substrate molecule to ADP

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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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Figure 7.7

© 2016 Pearson Education, Inc.

Enzyme

ADP

P

Substrate

Enzyme

Product

ATP

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Concept 7.2: Glycolysis harvests chemical energy by oxidizing glucose to pyruvate

Glycolysis (“sugar splitting”) breaks down glucose into two molecules of pyruvate

Glycolysis occurs in the cytoplasm and has two

major phases

Energy investment phase

Energy payoff phase

The net energy yield is 2 ATP plus 2 NADH per

glucose molecule

Glycolysis occurs whether or not O2 is present

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Figure 7.UN06

© 2016 Pearson Education, Inc.

GLYCOLYSIS PYRUVATE OXIDATION

CITRIC ACID

CYCLE

OXIDATIVE PHOSPHORYL-

ATION

ATP

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Figure 7.8

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Energy Investment Phase

Glucose

2 ATP used 2 ADP + 2 P

Energy Payoff Phase

4 ADP + 4 P 4 ATP formed

2 NAD+ + 4 e + 4 H+ 2 NADH + 2 H+

2 Pyruvate + 2 H2O

Net Glucose 2 Pyruvate + 2 H2O

2 ATP

2 NADH + 2 H+

4 ATP formed 2 ATP used

2 NAD+ + 4 e + 4 H+

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Figure 7.9-1

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GLYCOLYSIS: Energy Investment Phase

Glyceraldehyde

3-phosphate (G3P)

ATP Glucose

Glucose

6-phosphate

Fructose

6-phosphate ATP Fructose

1,6-bisphosphate

ADP ADP

Phosphogluco-

isomerase

Hexokinase Phospho-

fructokinase

Isomerase

Aldolase Dihydroxyacetone

phosphate (DHAP)

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Figure 7.9-1a-s1

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Glucose

GLYCOLYSIS: Energy Investment Phase

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Figure 7.9-1a-s2

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ATP Glucose

Glucose

6-phosphate

ADP

Hexokinase

GLYCOLYSIS: Energy Investment Phase

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Figure 7.9-1a-s3

© 2016 Pearson Education, Inc.

ATP Glucose

Glucose

6-phosphate

ADP

Fructose

6-phosphate

Hexokinase Phosphogluco-

isomerase

GLYCOLYSIS: Energy Investment Phase

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Figure 7.9-1b-s1

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GLYCOLYSIS: Energy Investment Phase

Fructose 6-phosphate

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Figure 7.9-1b-s2

© 2016 Pearson Education, Inc.

Fructose 6-phosphate

ATP Fructose 1,6-bisphosphate

ADP

Phospho-

fructokinase

GLYCOLYSIS: Energy Investment Phase

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Figure 7.9-1b-s3

© 2016 Pearson Education, Inc.

Glyceraldehyde 3-phosphate (G3P)

Fructose 6-phosphate

ATP Fructose 1,6-bisphosphate

ADP

Isomerase

Phospho-

fructokinase

Aldolase Dihydroxyacetone phosphate (DHAP)

GLYCOLYSIS: Energy Investment Phase

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Figure 7.9-2

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GLYCOLYSIS: Energy Payoff Phase

2 ATP

2 ADP

2

Glyceraldehyde 3-phosphate (G3P)

2 NADH

2 NA D+ + 2 H+

2 H2O 2

2 ADP

ATP

2 2 2 2

Triose phosphate

dehydrogenase 2 P i

Phospho- glycerokinase

Phospho- glyceromutase

Enolase

1,3-Bisphospho- glycerate

3-Phospho- glycerate

Pyruvate kinase

2-Phospho- glycerate

Phosphoenol- pyruvate (PEP)

Pyruvate

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Figure 7.9-2a-s1

© 2016 Pearson Education, Inc.

GLYCOLYSIS: Energy Payoff Phase

Glyceraldehyde 3-phosphate (G3P)

Isomerase

Aldolase Dihydroxyacetone phosphate (DHAP)

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Figure 7.9-2a-s2

© 2016 Pearson Education, Inc.

GLYCOLYSIS: Energy Payoff Phase

2

Glyceraldehyde 3-phosphate (G3P)

2 NADH

2 NAD+ + 2 H+

Isomerase

Triose phosphate

dehydrogenase 2 P i

Aldolase Dihydroxyacetone phosphate (DHAP)

1,3-Bisphospho- glycerate

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Figure 7.9-2a-s3

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GLYCOLYSIS: Energy Payoff Phase

2 ATP

2 ADP

2

Glyceraldehyde 3-phosphate (G3P)

2 NADH

2 NAD+ + 2 H+ 2

Isomerase

Triose phosphate

dehydrogenase

Phospho- glycerokinase

Aldolase Dihydroxyacetone phosphate (DHAP)

1,3-Bisphospho- glycerate

3-Phospho- glycerate

2 P i

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Figure 7.9-2b-s1

© 2016 Pearson Education, Inc.

GLYCOLYSIS: Energy Payoff Phase

2

3-Phospho- glycerate

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Figure 7.9-2b-s2

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GLYCOLYSIS: Energy Payoff Phase

2 H2O

2 2 2

Enolase Phospho-

glyceromutase

3-Phospho- glycerate

2-Phospho- glycerate

Phosphoenol-

pyruvate (PEP)

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Figure 7.9-2b-s3

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GLYCOLYSIS: Energy Payoff Phase

2 H2O

2

2

2 ADP

ATP

2 2 2

Enolase Pyruvate

kinase

Phospho-

glyceromutase

3-Phospho- glycerate

2-Phospho- glycerate

Phosphoenol-

pyruvate (PEP) Pyruvate

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Concept 7.3: 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

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

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Figure 7.UN07

© 2016 Pearson Education, Inc.

GLYCOLYSIS PYRUVATE OXIDATION

CITRIC ACID

CYCLE

OXIDATIVE PHOSPHORYL-

ATION

ATP

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Figure 7.10

© 2016 Pearson Education, Inc.

Pyruvate (from glycolysis, 2 molecules per glucose)

CYTOSOL

PYRUVATE OXIDATION

NAD+

NADH

+ H+ Acetyl CoA

CoA

CITRIC ACID

CYCLE

FADH2

FAD

ADP + P i

ATP MITOCHONDRION

3 NAD+

3 NADH

+ 3 H+

CO2

CoA

CoA

2 CO2

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Figure 7.10-1

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Pyruvate (from glycolysis, 2 molecules per glucose)

CYTOSOL

PYRUVATE OXIDATION

NAD+

NADH

+ H+

CO2

CoA

Acetyl CoA

CoA MITOCHONDRION

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Figure 7.10-2

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Acetyl CoA

CoA

CoA

CITRIC ACID

CYCLE

FADH2

FAD

ADP + P i

ATP MITOCHONDRION

2 CO2

3 NAD+

3 NADH

+ 3 H+

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The citric acid cycle, also called the Krebs cycle,

completes the breakdown 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 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 7.UN08

© 2016 Pearson Education, Inc.

GLYCOLYSIS PYRUVATE OXIDATION

CITRIC ACID

CYCLE

OXIDATIVE PHOSPHORYL-

ATION

ATP

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Figure 7.11-s1

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Acetyl CoA

Oxaloacetate

Citrate

CITRIC ACID

CYCLE

Isocitrate

CoA-SH

H2O

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Figure 7.11-s2

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Acetyl CoA

Oxaloacetate

Citrate

CITRIC ACID

CYCLE

Isocitrate

a-Ketoglutarate

CoA-SH

H2O

NAD+

NADH

+ H+

CO2

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Figure 7.11-s3

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Acetyl CoA

Oxaloacetate

Citrate

CITRIC ACID

CYCLE

CoA-SH

Isocitrate

a-Ketoglutarate

Succinyl CoA

CoA-SH

H2O

NAD+

NADH

+ H+

CO2

NAD+

NADH

+ H+

CO2

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Figure 7.11-s4

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Acetyl CoA

Oxaloacetate

Citrate

CITRIC ACID

CYCLE

CoA-SH

Isocitrate

a-Ketoglutarate

CoA-SH

Succinate

ADP

Succinyl CoA

CoA-SH

H2O

NAD+

NADH

+ H+

CO2

NAD+

NADH

+ H+

CO2

P

GTP GDP

i

ATP

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Figure 7.11-s5

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Acetyl CoA

Oxaloacetate

Citrate

CITRIC ACID

CYCLE

Fumarate CoA-SH

Isocitrate

a-Ketoglutarate

FAD

CoA-SH

Succinate

ADP

ATP

Succinyl CoA

CoA-SH

H2O

NAD+

NADH

+ H+

CO2

NAD+

NADH

+ H+

CO2 FADH2

P

GTP GDP

i

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Figure 7.11-s6

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Acetyl CoA

NADH

NAD+

Oxaloacetate

Malate Citrate

CITRIC ACID

CYCLE

Fumarate CoA-SH

Isocitrate

a-Ketoglutarate

FAD

CoA-SH

Succinate

ADP

ATP

Succinyl CoA

CoA-SH

H2O

NAD+

NADH

+ H+

CO2

NAD+

NADH

+ H+

CO2

+ H+

H2O

FADH2

P

GTP GDP

i

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Figure 7.11-1

© 2016 Pearson Education, Inc.

Start: Acetyl CoA adds its two-carbon group to oxaloacetate, producing citrate; this is a highly exergonic reaction.

Acetyl CoA

H2O

Oxaloacetate

Citrate

Isocitrate

CoA-SH

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Figure 7.11-2

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CO2 release

a-Ketoglutarate

CO2 CO2 release

Redox reaction: Isocitrate is oxidized; NAD+ is reduced.

Redox reaction: After CO2 release, the resulting four-carbon molecule is oxidized (reducing NAD+), then made reactive by addition of CoA.

Isocitrate NAD+

NADH

+ H+

CoA-SH

NAD+

NADH

+ H+

Succinyl CoA

CO2

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Figure 7.11-3

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Fumarate

FADH2

Redox reaction: Succinate is oxidized; FAD is reduced.

FAD

Succinate

GTP

ADP

Succinyl CoA

ATP formation ATP

P

GDP

i

CoA-SH

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Figure 7.11-4

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Redox reaction: Malate is oxidized; NAD+ is reduced.

NADH

+ H+

NAD+

Oxaloacetate

Malate

H2O

Fumarate

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Concept 7.4: 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 located 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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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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Figure 7.UN09

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GLYCOLYSIS PYRUVATE OXIDATION

CITRIC ACID

CYCLE

OXIDATIVE PHOSPHORYL-

ATION

ATP

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Figure 7.12

© 2016 Pearson Education, Inc.

50

40

30

20

10

0

(least electronegative)

FAD Complexes I-IV

FMN

Electron transport

chain

(most electronegative)

NADH

NAD+

FADH2

2

2

Fe•S

Cyt c1

Cyt c

Cyt a

Fe•S

Q

Cyt b

Cyt a3

II

III

IV

I

Fe•S

2 H+ + ½

H2O

O2

2 e

e

e

Fre

e e

ne

rgy (

G)

rela

tive

to

O2 (k

ca

l/m

ol)

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Figure 7.12-1

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II

III

IV

I

Electron transport

chain

50

40

30

20

10

NADH

FADH2

FAD

FMN

Fe•S Fe•S

Q

(least electronegative)

NAD+

Fe•S

Cyt c1

Cyt c

Cyt a

Complexes I-IV

Cyt b

Cyt a3

2

2

2

e

e

e

Fre

e e

ne

rgy

(G

) re

lati

ve

to

O2

(k

ca

l/m

ol)

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Figure 7.12-2

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2 H+ + ½

30

Electron transport chain

20

10

0 (most electronegative)

Fe•S

Cyt c1

Cyt c

Cyt a

Cyt a3

IV

2 e

O2

H2O

Fre

e e

ne

rgy

(G

) re

lati

ve

to

O2

(k

ca

l/m

ol)

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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 protein complex, 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 7.13

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Intermembrane

space

Mitochondrial

matrix

INTERMEMBRANE

SPACE

Inner mitochondrial membrane

Rotor

Stator

Internal

rod

ADP

Catalytic

knob

ATP

MITOCHONDRIAL MATRIX

(a) The ATP synthase protein complex

H+

+ P

i

(b) Computer model of ATP synthase

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Figure 7.13-1

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INTERMEMBRANE SPACE

Rotor

H+ Stator

Internal rod

Catalytic knob

ADP +

MITOCHONDRIAL MATRIX P ATP

(a) The ATP synthase protein complex

i

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Figure 7.13-2

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(b) Computer model of ATP synthase

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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

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Figure 7.UN09

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GLYCOLYSIS PYRUVATE OXIDATION

CITRIC ACID

CYCLE

OXIDATIVE PHOSPHORYL-

ATION

ATP

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Figure 7.14

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Electron transport chain Chemiosmosis

Oxidative phosphorylation

H+ H+

H+ ATP synthase

Protein complex of electron carriers

NADH NAD+

(carrying electrons

from food) H+

ATP

H+

I III

II

IV

Cyt c

Q

FADH2 2 H+ + ½ H2O

FAD O2

ADP + P i

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Figure 7.14-1

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Protein complex of electron carriers

NADH

(carrying electrons from food)

Electron transport chain

Q

I III

II

IV

H+ H+

Cyt c

H+

FADH2

NAD+

2 H+ + ½ H2O FAD

O2

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Figure 7.14-2

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ATP synthase

H+

ADP +

H+

ATP

Chemiosmosis

P i

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An Accounting of ATP Production by Cellular Respiration

During cellular respiration, most energy flows in the

following 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

molecules is not known exactly

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Figure 7.15

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CYTOSOL Electron shuttles span membrane

MITOCHONDRION

2 NADH

or

2 FADH2

2 NADH 6 NADH 2 FADH2 2 NADH

GLYCOLYSIS

Glucose 2 Pyruvate

PYRUVATE OXIDATION

2 Acetyl CoA

OXIDATIVE

PHOSPHORYLATION

(Electron transport

and chemiosmosis)

+ 2 ATP + 2 ATP + about 26 or 28 ATP

Maximum per glucose: About

30 or 32 ATP

CITRIC ACID

CYCLE

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Figure 7.15-1

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Electron shuttles

span membrane 2 NADH

or

2 FADH2

2 NADH

GLYCOLYSIS

Glucose 2 Pyruvate

+ 2 ATP

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Figure 7.15-2

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2 NADH 6 NADH 2 FADH2

PYRUVATE OXIDATION

2 Acetyl CoA

CITRIC

ACID

CYCLE

+ 2 ATP

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Figure 7.15-3

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2 NADH

or

2 FADH2

2 NADH 6 NADH 2 FADH2

OXIDATIVE

PHOSPHORYLATION

(Electron transport

and chemiosmosis)

+ about 26 or 28 ATP

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Figure 7.15-4

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Maximum per glucose: About

30 or 32 ATP

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Concept 7.5: 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

The first step releases CO2 from pyruvate, and the

second step reduces the resulting acetaldehyde to

ethanol

Alcohol fermentation by yeast is used in brewing,

winemaking, and baking

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Animation: Fermentation Overview

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Figure 7.16-1

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2 ADP + 2 P 2 ATP

Glucose GLYCOLYSIS

2 Pyruvate

2 2

+ 2 H+ 2 CO2

2 Ethanol

(a) Alcohol fermentation

2 Acetaldehyde

NAD+ NADH

i

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In lactic acid fermentation, pyruvate is reduced by

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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Figure 7.16-2

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2 Lactate

(b) Lactic acid fermentation

Glucose GLYCOLYSIS

NAD+ NADH

2 Pyruvate

2 2

+ 2 H+

2 ATP 2 ADP + 2 P i

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Figure 7.16

© 2016 Pearson Education, Inc.

2 Ethanol

(a) Alcohol fermentation

2 Acetaldehyde 2 Lactate

(b) Lactic acid fermentation

2 ADP + 2 P 2 ATP 2 ADP 2 ATP

Glucose GLYCOLYSIS Glucose GLYCOLYSIS

2 Pyruvate

2 2

+ 2 H+ 2 CO2

2 2

+ 2 H+

2 Pyruvate

NADH NAD+ NADH NAD+

i i + 2 P

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Comparing Fermentation with Anaerobic and Aerobic Respiration

All use glycolysis (net ATP = 2) to oxidize glucose

and other organic fuels to pyruvate

In all three, NAD+ is the oxidizing agent that accepts

electrons from food during glycolysis

The mechanism of NADH oxidation differs

In fermentation the final electron acceptor is an

organic molecule such as pyruvate or acetaldehyde

Cellular respiration transfers electrons from NADH to

a carrier molecule in the electron transport chain

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Cellular respiration produces about 32 ATP per

glucose molecule; fermentation produces 2 ATP per

glucose molecule

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Obligate anaerobes carry out only 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 7.17

© 2016 Pearson Education, Inc.

Glucose

Glycolysis

Pyruvate

No O2 present:

Fermentation

O2 present:

Aerobic cellular

respiration

CYTOSOL

MITOCHONDRION

Ethanol, lactate, or

other products

Acetyl CoA

CITRIC ACID

CYCLE

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The Evolutionary Significance of Glycolysis

Glycolysis is the most common metabolic pathway

among organisms on Earth, indicating that it

evolved early in the history of life

Early prokaryotes may have generated ATP

exclusively through glycolysis due to the low oxygen

content in the atmosphere

The location of glycolysis in the cytosol also

indicates its ancient origins; eukaryotic cells with

mitochondria evolved much later than prokaryotic

cells

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Concept 7.6: Glycolysis and the citric acid cycle connect to many other metabolic pathways

Glycolysis 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 and

amino groups must be removed before amino acids

can feed glycolysis or the citric acid cycle

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Fats are digested to glycerol (used in glycolysis)

and fatty acids

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 7.18-s1

© 2016 Pearson Education, Inc.

Proteins

Amino acids

Carbohydrates

Sugars

Fats

Glycerol Fatty acids

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Figure 7.18-s2

© 2016 Pearson Education, Inc.

Proteins

Amino acids

Carbohydrates

Sugars

Fats

Glycerol Fatty acids

GLYCOLYSIS

Glucose

NH3 Pyruvate

Glyceraldehyde 3- P

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Figure 7.18-s3

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Proteins

Amino acids

Carbohydrates

Sugars

Fats

Glycerol Fatty acids

GLYCOLYSIS

Glucose

NH3 Pyruvate

Acetyl CoA

Glyceraldehyde 3- P

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Figure 7.18-s4

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Proteins

Amino acids

Carbohydrates

Sugars

Fats

Glycerol Fatty acids

GLYCOLYSIS

Glucose

NH3 Pyruvate

Acetyl CoA

CITRIC

ACID CYCLE

Glyceraldehyde 3- P

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Figure 7.18-s5

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Proteins

Amino acids

Carbohydrates

Sugars

Fats

Glycerol Fatty acids

GLYCOLYSIS

Glucose

NH3 Pyruvate

Acetyl CoA

OXIDATIVE

PHOSPHORYLATION

Glyceraldehyde 3- P

CITRIC

ACID CYCLE

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Biosynthesis (Anabolic Pathways)

The body uses small molecules to build other

substances

Some of these small molecules come directly from

food; others can be produced during glycolysis or

the citric acid cycle

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Figure 7.UN10-1

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Figure 7.UN11

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Inputs

GLYCOLYSIS

Glucose

Outputs

2 Pyruvate ATP NADH 2 2

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Figure 7.UN12

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Inputs

2 Pyruvate 2 Acetyl CoA

2 Oxaloacetate CITRIC ACID

CYCLE

Outputs

2 ATP NADH 8

6 F A DH2 CO2 2

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Figure 7.UN13

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Cyt c

Q

I

IV

III

MITOCHONDRIAL MATRIX NAD+

NA DH

(carrying electrons from food)

FA DH2

II

FAD

H+

H+

H+

Protein complex of electron carriers

INTERMEMBRANE SPACE

H2O 2 H+ O2 + ½

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Figure 7.UN14

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INTER- MEMBRANE SPACE H+

MITO CHONDRIAL MATRIX ATP

synthase

ADP H+ ATP + P i

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Figure 7.UN15

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pH

dif

fere

nce

acro

ss m

em

bra

ne

Time

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Figure 7.UN16

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Ph

os

ph

ofr

ucto

kin

ase

a

cti

vit

y

Low ATP concentration

High ATP concentration

Fructose 6-phosphate concentration


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