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CAMPBELL BIOLOGY IN FOCUS
© 2014 Pearson Education, Inc.
Urry • Cain • Wasserman • Minorsky • Jackson • Reece
Lecture Presentations by
Kathleen Fitzpatrick and Nicole Tunbridge
7
Cellular
Respiration
and Fermentation
Overview: 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
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Figure 7.1
 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
© 2014 Pearson Education, Inc.
Animation: Carbon Cycle
© 2014 Pearson Education, Inc.
Figure 7.2
Light
energy
ECOSYSTEM
Photosynthesis
in chloroplasts
CO2 + H2O
Cellular respiration
in mitochondria
Organic
molecules
+ O2
ATP
ATP powers
most cellular work
Heat
energy
Concept 7.1: Catabolic pathways yield energy by
oxidizing organic fuels
 Several processes are central to cellular respiration
and related pathways
© 2014 Pearson Education, Inc.
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
© 2014 Pearson Education, Inc.
 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)
© 2014 Pearson Education, Inc.
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
© 2014 Pearson Education, Inc.
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)
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Figure 7.UN01
becomes oxidized
(loses electron)
becomes reduced
(gains electron)
© 2014 Pearson Education, Inc.
Figure 7.UN02
becomes oxidized
becomes reduced
 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
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Figure 7.3
Reactants Products
Methane
(reducing
agent)
Oxygen
(oxidizing
agent)
Carbon dioxide Water
becomes reduced
becomes oxidized
 Redox reactions that move electrons closer to
electronegative atoms, like oxygen, release chemical
energy that can be put to work
© 2014 Pearson Education, Inc.
Oxidation of Organic Fuel Molecules During
Cellular Respiration
 During cellular respiration, the 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
sythesis
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Figure 7.UN03
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
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Figure 7.4
NAD+
Nicotinamide
(oxidized form)
Nicotinamide
(reduced form)
Oxidation of NADH
Reduction of NAD+
Dehydrogenase
NADH
+ 2[H]
(from food)
2 e−
+ 2 H+
2 e−
+ H+
H+
H+
+
© 2014 Pearson Education, Inc.
Figure 7.4a
NAD+
Nicotinamide
(oxidized form)
© 2014 Pearson Education, Inc.
Figure 7.4b
Nicotinamide
(reduced form)
Oxidation of NADH
Reduction of NAD+
Dehydrogenase
NADH
2 e−
+ 2 H+
2 e−
+ H+
H+
H++ 2[H]
(from food)
+
© 2014 Pearson Education, Inc.
Figure 7.UN04
 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
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Figure 7.5
Explosive
release
(a) Uncontrolled reaction (b) Cellular respiration
H2O
Freeenergy,G
Freeenergy,G
Electrontransport
chain
Controlled
release of
energy
H2O
2 H+
2 e−
2 H+
+ 2 e−
ATP
ATP
ATP
½
½
½H2 + O2 O2
O2
2 H +
The Stages of Cellular Respiration: A Preview
 Harvesting of energy from glucose has three stages
 Glycolysis (breaks down glucose into two molecules
of pyruvate)
 Pyruvate oxidation and the citric acid cycle
(completes the breakdown of glucose)
 Oxidative phosphorylation (accounts for most of
the ATP synthesis)
© 2014 Pearson Education, Inc.
Animation: Cellular Respiration
© 2014 Pearson Education, Inc.
Figure 7.UN05
Glycolysis (color-coded teal throughout the chapter)
Pyruvate oxidation and the citric acid cycle
(color-coded salmon)
1.
Oxidative phosphorylation: electron transport and
chemiosmosis (color-coded violet)
2.
3.
© 2014 Pearson Education, Inc.
Figure 7.6-1
Electrons
via NADH
Glycolysis
Glucose Pyruvate
CYTOSOL
ATP
Substrate-level
MITOCHONDRION
© 2014 Pearson Education, Inc.
Figure 7.6-2
Electrons
via NADH
Glycolysis
Glucose Pyruvate
Pyruvate
oxidation
Acetyl CoA
Citric
acid
cycle
Electrons
via NADH and
FADH2
CYTOSOL
ATP
Substrate-level
ATP
Substrate-level
MITOCHONDRION
© 2014 Pearson Education, Inc.
Figure 7.6-3
Electrons
via NADH
Glycolysis
Glucose Pyruvate
Pyruvate
oxidation
Acetyl CoA
Citric
acid
cycle
Electrons
via NADH and
FADH2
Oxidative
phosphorylation:
electron transport
and
chemiosmosis
CYTOSOL
ATP
Substrate-level
ATP
Substrate-level
MITOCHONDRION
ATP
Oxidative
 The process that generates most of the ATP is
called oxidative phosphorylation because it is
powered by redox reactions
© 2014 Pearson Education, Inc.
 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
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Figure 7.7
Substrate
P
ADP
Product
ATP+
Enzyme Enzyme
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
 Glycolysis occurs whether or not O2 is present
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Figure 7.UN06
Glycolysis Pyruvate
oxidation
Citric
acid
cycle
Oxidative
phosphorylation
ATP ATP ATP
© 2014 Pearson Education, Inc.
Figure 7.8
Energy Investment Phase
Energy Payoff Phase
Net
Glucose
Glucose
2 ADP + 2 P
4 ADP + 4 P
2 NAD+
+ 4 e−
+ 4 H+
2 NAD+
+ 4 e−
+ 4 H+
4 ATP formed − 2 ATP used
2 ATP
4 ATP
used
formed
2 NADH + 2 H+
2 Pyruvate + 2 H2O
2 Pyruvate + 2 H2O
2 NADH + 2 H+
2 ATP
© 2014 Pearson Education, Inc.
Figure 7.9a
Glycolysis: Energy Investment Phase
Glucose
ATP
ADP
Glucose
6-phosphate
Phosphogluco-
isomerase
Hexokinase
1
2 3
4
ATP
ADP
Fructose
6-phosphate
Phospho-
fructokinase
Fructose
1,6-bisphosphate
Aldolase
Isomerase
5
Glyceraldehyde
3-phosphate (G3P)
Dihydroxyacetone
phosphate (DHAP)
© 2014 Pearson Education, Inc.
Figure 7.9aa-1
Glycolysis: Energy Investment Phase
Glucose
© 2014 Pearson Education, Inc.
Figure 7.9aa-2
Glycolysis: Energy Investment Phase
Glucose
Glucose
6-phosphate
ADP
ATP
Hexokinase
1
© 2014 Pearson Education, Inc.
Figure 7.9aa-3
Glycolysis: Energy Investment Phase
Glucose
Glucose
6-phosphate
ADP
ATP
Hexokinase
1
Fructose
6-phosphate
Phosphogluco-
isomerase
2
© 2014 Pearson Education, Inc.
Figure 7.9ab-1
Glycolysis: Energy Investment Phase
Fructose
6-phosphate
© 2014 Pearson Education, Inc.
Figure 7.9ab-2
Glycolysis: Energy Investment Phase
Fructose
6-phosphate
Phospho-
fructokinase
3
Fructose
1,6-bisphosphate
ATP
ADP
© 2014 Pearson Education, Inc.
Figure 7.9ab-3
Glycolysis: Energy Investment Phase
Fructose
6-phosphate
Phospho-
fructokinase
3
Aldolase
Isomerase
4
5
Fructose
1,6-bisphosphate
Glyceraldehyde
3-phosphate (G3P)
ATP
ADP
Dihydroxyacetone
phosphate (DHAP)
© 2014 Pearson Education, Inc.
Figure 7.9b
Glycolysis: Energy Payoff Phase
2 NAD+
Glyceraldehyde
3-phosphate (G3P)
Triose
phosphate
dehydrogenase
6
+ 2 H+
2 NADH
2
2 P
i
1,3-Bisphospho-
glycerate
3-Phospho-
glycerate
2-Phospho-
glycerate
Phosphoenol-
pyruvate (PEP)
Pyruvate
Phospho-
glycerokinase
Phospho-
glyceromutase
Enolase Pyruvate
kinase
2 ADP 2 2 2 2
2 ADP
2 ATP
2 H2O
2 ATP
9
1087
© 2014 Pearson Education, Inc.
Figure 7.9ba-1
Isomerase
4
Glyceraldehyde
3-phosphate (G3P)
Dihydroxyacetone
phosphate (DHAP)
Glycolysis: Energy Payoff Phase
Aldolase
5
© 2014 Pearson Education, Inc.
Figure 7.9ba-2
Isomerase
Glyceraldehyde
3-phosphate (G3P)
Dihydroxyacetone
phosphate (DHAP)
Glycolysis: Energy Payoff Phase
2 NAD+
Triose
phosphate
dehydrogenase
+ 2 H+
2 NADH
2
1,3-Bisphospho-
glycerate
2
Aldolase
P
i
5 6
4
© 2014 Pearson Education, Inc.
Figure 7.9ba-3
Isomerase
Glyceraldehyde
3-phosphate (G3P)
Dihydroxyacetone
phosphate (DHAP)
Glycolysis: Energy Payoff Phase
2 NAD+
Triose
phosphate
dehydrogenase
+ 2 H+
2 NADH
2
1,3-Bisphospho-
glycerate
3-Phospho-
glycerate
Phospho-
glycerokinase
2 ADP
2 ATP
2
Aldolase
P
i
2
5 7
6
4
© 2014 Pearson Education, Inc.
Figure 7.9bb-1
3-Phospho-
glycerate
Glycolysis: Energy Payoff Phase
2
© 2014 Pearson Education, Inc.
Figure 7.9bb-2
83-Phospho-
glycerate
Glycolysis: Energy Payoff Phase
Phospho-
glyceromutase
222
2 H2O
2-Phospho-
glycerate
Phosphoenol-
pyruvate (PEP)
Enolase
9
© 2014 Pearson Education, Inc.
Figure 7.9bb-3
3-Phospho-
glycerate
Glycolysis: Energy Payoff Phase
2 ATP
Phospho-
glyceromutase
2222
2 ADP
2 H2O
2-Phospho-
glycerate
Phosphoenol-
pyruvate (PEP)
Pyruvate
Enolase Pyruvate
kinase
9
108
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
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Figure 7.UN07
Glycolysis Pyruvate
oxidation
Citric
acid
cycle
Oxidative
phosphorylation
ATP ATP ATP
© 2014 Pearson Education, Inc.
Figure 7.10
CYTOSOLPyruvate
(from glycolysis,
2 molecules per glucose)
CO2
CoA
NAD+
NADH
MITOCHONDRION CoA
CoA
Acetyl CoA+ H+
Citric
acid
cycle
FADH2
FAD
ADP + P
i
ATP
NADH
3 NAD+
3
+ 3 H+
2 CO2
© 2014 Pearson Education, Inc.
Figure 7.10a
CYTOSOLPyruvate
(from glycolysis,
2 molecules per glucose)
CO2
CoA
NAD+
NADH
MITOCHONDRION CoA
Acetyl CoA+ H+
© 2014 Pearson Education, Inc.
Figure 7.10b
CoA
Citric
acid
cycle
FADH2
FAD
ADP + P
i
ATP
NADH
3 NAD+
3
+ 3 H+
2 CO2
CoA
Acetyl CoA
 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
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Figure 7.UN08
Glycolysis Pyruvate
oxidation
Oxidative
phosphorylation
ATP ATP ATP
Citric
acid
cycle
© 2014 Pearson Education, Inc.
Figure 7.11-1
Acetyl CoA
Oxaloacetate
CoA-SH
Citrate
H2O
Isocitrate
Citric
acid
cycle
2
1
© 2014 Pearson Education, Inc.
Figure 7.11-2
Acetyl CoA
Oxaloacetate
Citrate
H2O
Isocitrate
NADH
NAD+
+ H+
CO2
α-Ketoglutarate
Citric
acid
cycle
3
1
CoA-SH
2
© 2014 Pearson Education, Inc.
Figure 7.11-3
Acetyl CoA
Oxaloacetate
Citrate
H2O
Isocitrate
NADH
NAD+
+ H+
CO2
α-Ketoglutarate
Citric
acid
cycle
CoA-SH
CO2
NAD+
NADH
+ H+
Succinyl
CoA
4
1
3
CoA-SH
2
© 2014 Pearson Education, Inc.
Figure 7.11-4
Acetyl CoA
Oxaloacetate
Citrate
H2O
Isocitrate
NADH
NAD+
+ H+
CO2
α-Ketoglutarate
Citric
acid
cycle
CoA-SH
CO2
NAD+
NADH
+ H+
ATP formation
Succinyl
CoA
ADP
GDPGTP
P
i
ATP
Succinate
5
4
1
CoA-SH
3
CoA-SH
2
© 2014 Pearson Education, Inc.
Figure 7.11-5
Malate
Succinate
FAD
FADH2
Fumarate
H2O
7
6
Acetyl CoA
Oxaloacetate
Citrate
H2O
Isocitrate
NADH
NAD+
+ H+
CO2
α-Ketoglutarate
Citric
acid
cycle
CoA-SH
CO2
NAD+
NADH
+ H+
ATP formation
Succinyl
CoA
ADP
GDPGTP
P
i
ATP
5
4
1
CoA-SH
3
CoA-SH
2
© 2014 Pearson Education, Inc.
Figure 7.11-6
NADH
NAD+
+ H+
8
Malate
Succinate
FAD
FADH2
Fumarate
H2O
7
6
Acetyl CoA
Oxaloacetate
Citrate
H2O
Isocitrate
NADH
NAD+
+ H+
CO2
α-Ketoglutarate
Citric
acid
cycle
CoA-SH
CO2
NAD+
NADH
+ H+
ATP formation
Succinyl
CoA
ADP
GDPGTP
P
i
ATP
5
4
1
CoA-SH
3
CoA-SH
2
© 2014 Pearson Education, Inc.
Figure 7.11a
CoA-SH
Acetyl CoA
Start: Acetyl CoA adds its
two-carbon group to
oxaloacetate, producing
citrate; this is a highly
exergonic reaction.
Oxaloacetate
Citrate
Isocitrate
H2O1
2
© 2014 Pearson Education, Inc.
Figure 7.11b
Isocitrate 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.
CO2 release
CO2 release
α-Ketoglutarate
Succinyl
CoA
NAD+
NADH
+ H+
CO2
CO2
CoA-SH
NAD+
NADH
+ H+
3
4
© 2014 Pearson Education, Inc.
Figure 7.11c
CoA-SH
Redox reaction:
Succinate is oxidized;
FAD is reduced.
Fumarate
Succinate
Succinyl
CoA
ATP formation
ATP
ADP
GDPGTP
FAD
FADH2
P
i
5
6
© 2014 Pearson Education, Inc.
Figure 7.11d
Redox reaction:
Malate is oxidized;
NAD+
is reduced.
Fumarate
Malate
Oxaloacetate
H2O
NAD+
+ H+
NADH
7
8
 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
© 2014 Pearson Education, Inc.
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
© 2014 Pearson Education, Inc.
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
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Figure 7.UN09
Glycolysis
Pyruvate
oxidation
Citric
acid
cycle
Oxidative
phosphorylation:
electron transport
and chemiosmosis
ATP ATP ATP
© 2014 Pearson Education, Inc.
Figure 7.12
Multiprotein
complexes
(originally from
NADH or FADH2)
Freeenergy(G)relativetoO2(kcal/mol)
50
40
30
20
10
0
NADH
NAD+
FADH2
FAD
2
2
e−
e−
FMN
Fe•S Fe•S
Q
I
II
III
Cyt b
Cyt c1
Fe•S
Cyt c
IV
Cyt a
Cyt a3
2 e−
O22 H+
+ ½
H2O
© 2014 Pearson Education, Inc.
Figure 7.12a
Multiprotein
complexes
Freeenergy(G)relativetoO2(kcal/mol)
50
40
30
20
10
NADH
NAD+
FADH2
FAD
2
2
e−
e−
FMN
Fe•S Fe•S
Q
I
II
III
Cyt b
Cyt c1
Fe•S
Cyt c
IV
Cyt a
Cyt a3
2 e−
© 2014 Pearson Education, Inc.
Figure 7.12b
30
20
10
0
Cyt c1
Cyt c
IV
Cyt a
Cyt a3
2 e−
Freeenergy(G)relativetoO2(kcal/mol)
(originally from
NADH or FADH2)
2 H+
+ ½ O2
H2O
 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
© 2014 Pearson Education, Inc.
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
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Video: ATP Synthase 3-D Side View
Video: ATP Synthase 3-D Top View
© 2014 Pearson Education, Inc.
Figure 7.13
INTERMEMBRANE SPACE
MITOCHONDRIAL MATRIX
Rotor
Internal rod
Catalytic knob
Stator
H+
ATP
ADP
P
i
+
 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
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Figure 7.UN09
Glycolysis
Pyruvate
oxidation
Citric
acid
cycle
Oxidative
phosphorylation:
electron transport
and chemiosmosis
ATP ATP ATP
© 2014 Pearson Education, Inc.
Figure 7.14
Protein
complex
of electron
carriers
H+
H+
H+
H+
Q
I
II
III
FADH2
FAD
NAD+
NADH
(carrying electrons
from food)
Electron transport chain
Oxidative phosphorylation
Chemiosmosis
ATP
synthase
H+
ADP + ATPP
i
H2O2 H+
+ ½ O2
IV
Cyt c
1 2
© 2014 Pearson Education, Inc.
Figure 7.14a
Protein
complex
of electron
carriers
H+
Q
I
II
III
FADH2
FAD
NAD+
NADH
(carrying electrons
from food)
Electron transport chain
H2O2 H+
+ ½ O2
Cyt c
1
IV
H+
H+
© 2014 Pearson Education, Inc.
Figure 7.14b
ATP
synthase
Chemiosmosis2
H+
H+
ADP + P
i
ATP
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
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Figure 7.15
Electron shuttles
span membrane
CYTOSOL
2 NADH
2 NADH
2 FADH2
or
2 NADH
Glycolysis
Glucose 2
Pyruvate
Pyruvate
oxidation
2 Acetyl CoA
Citric
acid
cycle
6 NADH 2 FADH2
Oxidative
phosphorylation:
electron transport
and
chemiosmosis
+ about 26 or 28 ATP+ 2 ATP + 2 ATP
About
30 or 32 ATP
Maximum per glucose:
MITOCHONDRION
© 2014 Pearson Education, Inc.
Figure 7.15a
Electron shuttles
span membrane
2 NADH
2 FADH2
or
2 NADH
Glycolysis
Glucose 2
Pyruvate
+ 2 ATP
© 2014 Pearson Education, Inc.
Figure 7.15b
2 NADH 6 NADH 2 FADH2
Citric
acid
cycle
Pyruvate
oxidation
2 Acetyl CoA
+ 2 ATP
© 2014 Pearson Education, Inc.
Figure 7.15c
2 NADH
2 NADH 6 NADH 2 FADH2
2 FADH2
or
Oxidative
phosphorylation:
electron transport
and
chemiosmosis
+ about 26 or 28 ATP
© 2014 Pearson Education, Inc.
Figure 7.15d
Maximum per glucose:
About
30 or 32 ATP
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
© 2014 Pearson Education, Inc.
 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
© 2014 Pearson Education, Inc.
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
© 2014 Pearson Education, Inc.
 In alcohol fermentation, pyruvate is converted to
ethanol in two steps
 The first step releases CO2 from pyruvate, and the
second step reduces acetaldehyde to ethanol
 Alcohol fermentation by yeast is used in brewing,
winemaking, and baking
© 2014 Pearson Education, Inc.
Animation: Fermentation Overview
© 2014 Pearson Education, Inc.
Figure 7.16
2 ADP + 2 2 ATPP
i
Glucose Glycolysis
2 Pyruvate
2 CO2
2 NADH
+ 2 H+
2 NAD+
2 Ethanol
(a) Alcohol fermentation
2 Acetaldehyde
(b) Lactic acid fermentation
2 Lactate
2 NADH
+ 2 H+
2 NAD+
2 Pyruvate
Glycolysis
2 ATP2 ADP + 2 P
i
Glucose
© 2014 Pearson Education, Inc.
Figure 7.16a
2 ADP + 2 2 ATPP
i
Glucose Glycolysis
2 Pyruvate
2 CO2
2 NADH
+ 2 H+
2 NAD+
2 Ethanol
(a) Alcohol fermentation
2 Acetaldehyde
© 2014 Pearson Education, Inc.
Figure 7.16b
2 ADP + 2 2 ATPP
i
Glucose Glycolysis
2 Pyruvate
2 NADH
+ 2 H+
2 NAD+
(b) Lactic acid fermentation
2 Lactate
 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
© 2014 Pearson Education, Inc.
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
© 2014 Pearson Education, Inc.
 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
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Figure 7.17
Glucose
CYTOSOL
Glycolysis
Pyruvate
O2 present:
Aerobic cellular
respiration
No O2 present:
Fermentation
Ethanol,
lactate, or
other products
Acetyl CoA
Citric
acid
cycle
MITOCHONDRION
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
© 2014 Pearson Education, Inc.
Concept 7.6: 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
© 2014 Pearson Education, Inc.
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
© 2014 Pearson Education, Inc.
 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
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Figure 7.18-1
Proteins
Amino
acids
Carbohydrates
Sugars
Fats
Glycerol Fatty
acids
© 2014 Pearson Education, Inc.
Figure 7.18-2
Proteins
Amino
acids
Carbohydrates
Sugars
Glucose
Glycolysis
Glyceraldehyde 3-
Pyruvate
P
NH3
Fats
Glycerol Fatty
acids
© 2014 Pearson Education, Inc.
Figure 7.18-3
Proteins
Amino
acids
Carbohydrates
Sugars
Glucose
Glycolysis
Glyceraldehyde 3-
Pyruvate
P
Acetyl CoA
NH3
Fats
Glycerol Fatty
acids
© 2014 Pearson Education, Inc.
Figure 7.18-4
Proteins
Amino
acids
Carbohydrates
Sugars
Glucose
Glycolysis
Glyceraldehyde 3-
Pyruvate
P
Acetyl CoA
Citric
acid
cycle
NH3
Fats
Glycerol Fatty
acids
© 2014 Pearson Education, Inc.
Figure 7.18-5
Proteins
Amino
acids
Carbohydrates
Sugars
Glucose
Glycolysis
Glyceraldehyde 3-
Pyruvate
P
Acetyl CoA
Citric
acid
cycle
NH3
Fats
Glycerol Fatty
acids
Oxidative
phosphorylation
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
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Figure 7.UN10a
© 2014 Pearson Education, Inc.
Figure 7.UN10b
© 2014 Pearson Education, Inc.
Figure 7.UN11
Inputs
Glucose
Glycolysis
2 Pyruvate + 2
Outputs
ATP NADH+ 2
© 2014 Pearson Education, Inc.
Figure 7.UN12
Inputs
2 Pyruvate 2 Acetyl CoA
2 Oxaloacetate
Citric
acid
cycle
Outputs
ATP
CO2
2
6 2
8 NADH
FADH2
© 2014 Pearson Education, Inc.
Figure 7.UN13a
Protein
complex
of electron
carriers
INTERMEMBRANE
SPACE
MITOCHONDRIAL MATRIX
(carrying electrons from food)
NADH NAD+
FADH2
FAD
Cyt c
Q
I
II
III
IV
2 H+
+
½O2
H2O
H+
H+
H+
© 2014 Pearson Education, Inc.
Figure 7.UN13b
INTER-
MEMBRANE
SPACE
MITO-
CHONDRIAL
MATRIX
ATP
synthase
ATPADP + H+
H+
P
i
© 2014 Pearson Education, Inc.
Figure 7.UN14
Time
pHdifference
acrossmembrane

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Biology in Focus - Chapter 7

  • 1. CAMPBELL BIOLOGY IN FOCUS © 2014 Pearson Education, Inc. Urry • Cain • Wasserman • Minorsky • Jackson • Reece Lecture Presentations by Kathleen Fitzpatrick and Nicole Tunbridge 7 Cellular Respiration and Fermentation
  • 2. Overview: 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 © 2014 Pearson Education, Inc.
  • 3. © 2014 Pearson Education, Inc. Figure 7.1
  • 4.  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 © 2014 Pearson Education, Inc. Animation: Carbon Cycle
  • 5. © 2014 Pearson Education, Inc. Figure 7.2 Light energy ECOSYSTEM Photosynthesis in chloroplasts CO2 + H2O Cellular respiration in mitochondria Organic molecules + O2 ATP ATP powers most cellular work Heat energy
  • 6. Concept 7.1: Catabolic pathways yield energy by oxidizing organic fuels  Several processes are central to cellular respiration and related pathways © 2014 Pearson Education, Inc.
  • 7. 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 © 2014 Pearson Education, Inc.
  • 8.  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) © 2014 Pearson Education, Inc.
  • 9. 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 © 2014 Pearson Education, Inc.
  • 10. 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) © 2014 Pearson Education, Inc.
  • 11. © 2014 Pearson Education, Inc. Figure 7.UN01 becomes oxidized (loses electron) becomes reduced (gains electron)
  • 12. © 2014 Pearson Education, Inc. Figure 7.UN02 becomes oxidized becomes reduced
  • 13.  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 © 2014 Pearson Education, Inc.
  • 14. © 2014 Pearson Education, Inc. Figure 7.3 Reactants Products Methane (reducing agent) Oxygen (oxidizing agent) Carbon dioxide Water becomes reduced becomes oxidized
  • 15.  Redox reactions that move electrons closer to electronegative atoms, like oxygen, release chemical energy that can be put to work © 2014 Pearson Education, Inc.
  • 16. Oxidation of Organic Fuel Molecules During Cellular Respiration  During cellular respiration, the 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 sythesis © 2014 Pearson Education, Inc.
  • 17. © 2014 Pearson Education, Inc. Figure 7.UN03 becomes oxidized becomes reduced
  • 18. 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 © 2014 Pearson Education, Inc.
  • 19. © 2014 Pearson Education, Inc. Figure 7.4 NAD+ Nicotinamide (oxidized form) Nicotinamide (reduced form) Oxidation of NADH Reduction of NAD+ Dehydrogenase NADH + 2[H] (from food) 2 e− + 2 H+ 2 e− + H+ H+ H+ +
  • 20. © 2014 Pearson Education, Inc. Figure 7.4a NAD+ Nicotinamide (oxidized form)
  • 21. © 2014 Pearson Education, Inc. Figure 7.4b Nicotinamide (reduced form) Oxidation of NADH Reduction of NAD+ Dehydrogenase NADH 2 e− + 2 H+ 2 e− + H+ H+ H++ 2[H] (from food) +
  • 22. © 2014 Pearson Education, Inc. Figure 7.UN04
  • 23.  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 © 2014 Pearson Education, Inc.
  • 24. © 2014 Pearson Education, Inc. Figure 7.5 Explosive release (a) Uncontrolled reaction (b) Cellular respiration H2O Freeenergy,G Freeenergy,G Electrontransport chain Controlled release of energy H2O 2 H+ 2 e− 2 H+ + 2 e− ATP ATP ATP ½ ½ ½H2 + O2 O2 O2 2 H +
  • 25. The Stages of Cellular Respiration: A Preview  Harvesting of energy from glucose has three stages  Glycolysis (breaks down glucose into two molecules of pyruvate)  Pyruvate oxidation and the citric acid cycle (completes the breakdown of glucose)  Oxidative phosphorylation (accounts for most of the ATP synthesis) © 2014 Pearson Education, Inc. Animation: Cellular Respiration
  • 26. © 2014 Pearson Education, Inc. Figure 7.UN05 Glycolysis (color-coded teal throughout the chapter) Pyruvate oxidation and the citric acid cycle (color-coded salmon) 1. Oxidative phosphorylation: electron transport and chemiosmosis (color-coded violet) 2. 3.
  • 27. © 2014 Pearson Education, Inc. Figure 7.6-1 Electrons via NADH Glycolysis Glucose Pyruvate CYTOSOL ATP Substrate-level MITOCHONDRION
  • 28. © 2014 Pearson Education, Inc. Figure 7.6-2 Electrons via NADH Glycolysis Glucose Pyruvate Pyruvate oxidation Acetyl CoA Citric acid cycle Electrons via NADH and FADH2 CYTOSOL ATP Substrate-level ATP Substrate-level MITOCHONDRION
  • 29. © 2014 Pearson Education, Inc. Figure 7.6-3 Electrons via NADH Glycolysis Glucose Pyruvate Pyruvate oxidation Acetyl CoA Citric acid cycle Electrons via NADH and FADH2 Oxidative phosphorylation: electron transport and chemiosmosis CYTOSOL ATP Substrate-level ATP Substrate-level MITOCHONDRION ATP Oxidative
  • 30.  The process that generates most of the ATP is called oxidative phosphorylation because it is powered by redox reactions © 2014 Pearson Education, Inc.
  • 31.  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 © 2014 Pearson Education, Inc.
  • 32. © 2014 Pearson Education, Inc. Figure 7.7 Substrate P ADP Product ATP+ Enzyme Enzyme
  • 33. 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  Glycolysis occurs whether or not O2 is present © 2014 Pearson Education, Inc.
  • 34. © 2014 Pearson Education, Inc. Figure 7.UN06 Glycolysis Pyruvate oxidation Citric acid cycle Oxidative phosphorylation ATP ATP ATP
  • 35. © 2014 Pearson Education, Inc. Figure 7.8 Energy Investment Phase Energy Payoff Phase Net Glucose Glucose 2 ADP + 2 P 4 ADP + 4 P 2 NAD+ + 4 e− + 4 H+ 2 NAD+ + 4 e− + 4 H+ 4 ATP formed − 2 ATP used 2 ATP 4 ATP used formed 2 NADH + 2 H+ 2 Pyruvate + 2 H2O 2 Pyruvate + 2 H2O 2 NADH + 2 H+ 2 ATP
  • 36. © 2014 Pearson Education, Inc. Figure 7.9a Glycolysis: Energy Investment Phase Glucose ATP ADP Glucose 6-phosphate Phosphogluco- isomerase Hexokinase 1 2 3 4 ATP ADP Fructose 6-phosphate Phospho- fructokinase Fructose 1,6-bisphosphate Aldolase Isomerase 5 Glyceraldehyde 3-phosphate (G3P) Dihydroxyacetone phosphate (DHAP)
  • 37. © 2014 Pearson Education, Inc. Figure 7.9aa-1 Glycolysis: Energy Investment Phase Glucose
  • 38. © 2014 Pearson Education, Inc. Figure 7.9aa-2 Glycolysis: Energy Investment Phase Glucose Glucose 6-phosphate ADP ATP Hexokinase 1
  • 39. © 2014 Pearson Education, Inc. Figure 7.9aa-3 Glycolysis: Energy Investment Phase Glucose Glucose 6-phosphate ADP ATP Hexokinase 1 Fructose 6-phosphate Phosphogluco- isomerase 2
  • 40. © 2014 Pearson Education, Inc. Figure 7.9ab-1 Glycolysis: Energy Investment Phase Fructose 6-phosphate
  • 41. © 2014 Pearson Education, Inc. Figure 7.9ab-2 Glycolysis: Energy Investment Phase Fructose 6-phosphate Phospho- fructokinase 3 Fructose 1,6-bisphosphate ATP ADP
  • 42. © 2014 Pearson Education, Inc. Figure 7.9ab-3 Glycolysis: Energy Investment Phase Fructose 6-phosphate Phospho- fructokinase 3 Aldolase Isomerase 4 5 Fructose 1,6-bisphosphate Glyceraldehyde 3-phosphate (G3P) ATP ADP Dihydroxyacetone phosphate (DHAP)
  • 43. © 2014 Pearson Education, Inc. Figure 7.9b Glycolysis: Energy Payoff Phase 2 NAD+ Glyceraldehyde 3-phosphate (G3P) Triose phosphate dehydrogenase 6 + 2 H+ 2 NADH 2 2 P i 1,3-Bisphospho- glycerate 3-Phospho- glycerate 2-Phospho- glycerate Phosphoenol- pyruvate (PEP) Pyruvate Phospho- glycerokinase Phospho- glyceromutase Enolase Pyruvate kinase 2 ADP 2 2 2 2 2 ADP 2 ATP 2 H2O 2 ATP 9 1087
  • 44. © 2014 Pearson Education, Inc. Figure 7.9ba-1 Isomerase 4 Glyceraldehyde 3-phosphate (G3P) Dihydroxyacetone phosphate (DHAP) Glycolysis: Energy Payoff Phase Aldolase 5
  • 45. © 2014 Pearson Education, Inc. Figure 7.9ba-2 Isomerase Glyceraldehyde 3-phosphate (G3P) Dihydroxyacetone phosphate (DHAP) Glycolysis: Energy Payoff Phase 2 NAD+ Triose phosphate dehydrogenase + 2 H+ 2 NADH 2 1,3-Bisphospho- glycerate 2 Aldolase P i 5 6 4
  • 46. © 2014 Pearson Education, Inc. Figure 7.9ba-3 Isomerase Glyceraldehyde 3-phosphate (G3P) Dihydroxyacetone phosphate (DHAP) Glycolysis: Energy Payoff Phase 2 NAD+ Triose phosphate dehydrogenase + 2 H+ 2 NADH 2 1,3-Bisphospho- glycerate 3-Phospho- glycerate Phospho- glycerokinase 2 ADP 2 ATP 2 Aldolase P i 2 5 7 6 4
  • 47. © 2014 Pearson Education, Inc. Figure 7.9bb-1 3-Phospho- glycerate Glycolysis: Energy Payoff Phase 2
  • 48. © 2014 Pearson Education, Inc. Figure 7.9bb-2 83-Phospho- glycerate Glycolysis: Energy Payoff Phase Phospho- glyceromutase 222 2 H2O 2-Phospho- glycerate Phosphoenol- pyruvate (PEP) Enolase 9
  • 49. © 2014 Pearson Education, Inc. Figure 7.9bb-3 3-Phospho- glycerate Glycolysis: Energy Payoff Phase 2 ATP Phospho- glyceromutase 2222 2 ADP 2 H2O 2-Phospho- glycerate Phosphoenol- pyruvate (PEP) Pyruvate Enolase Pyruvate kinase 9 108
  • 50. 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 © 2014 Pearson Education, Inc.
  • 51. © 2014 Pearson Education, Inc. Figure 7.UN07 Glycolysis Pyruvate oxidation Citric acid cycle Oxidative phosphorylation ATP ATP ATP
  • 52. © 2014 Pearson Education, Inc. Figure 7.10 CYTOSOLPyruvate (from glycolysis, 2 molecules per glucose) CO2 CoA NAD+ NADH MITOCHONDRION CoA CoA Acetyl CoA+ H+ Citric acid cycle FADH2 FAD ADP + P i ATP NADH 3 NAD+ 3 + 3 H+ 2 CO2
  • 53. © 2014 Pearson Education, Inc. Figure 7.10a CYTOSOLPyruvate (from glycolysis, 2 molecules per glucose) CO2 CoA NAD+ NADH MITOCHONDRION CoA Acetyl CoA+ H+
  • 54. © 2014 Pearson Education, Inc. Figure 7.10b CoA Citric acid cycle FADH2 FAD ADP + P i ATP NADH 3 NAD+ 3 + 3 H+ 2 CO2 CoA Acetyl CoA
  • 55.  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 © 2014 Pearson Education, Inc.
  • 56. © 2014 Pearson Education, Inc. Figure 7.UN08 Glycolysis Pyruvate oxidation Oxidative phosphorylation ATP ATP ATP Citric acid cycle
  • 57. © 2014 Pearson Education, Inc. Figure 7.11-1 Acetyl CoA Oxaloacetate CoA-SH Citrate H2O Isocitrate Citric acid cycle 2 1
  • 58. © 2014 Pearson Education, Inc. Figure 7.11-2 Acetyl CoA Oxaloacetate Citrate H2O Isocitrate NADH NAD+ + H+ CO2 α-Ketoglutarate Citric acid cycle 3 1 CoA-SH 2
  • 59. © 2014 Pearson Education, Inc. Figure 7.11-3 Acetyl CoA Oxaloacetate Citrate H2O Isocitrate NADH NAD+ + H+ CO2 α-Ketoglutarate Citric acid cycle CoA-SH CO2 NAD+ NADH + H+ Succinyl CoA 4 1 3 CoA-SH 2
  • 60. © 2014 Pearson Education, Inc. Figure 7.11-4 Acetyl CoA Oxaloacetate Citrate H2O Isocitrate NADH NAD+ + H+ CO2 α-Ketoglutarate Citric acid cycle CoA-SH CO2 NAD+ NADH + H+ ATP formation Succinyl CoA ADP GDPGTP P i ATP Succinate 5 4 1 CoA-SH 3 CoA-SH 2
  • 61. © 2014 Pearson Education, Inc. Figure 7.11-5 Malate Succinate FAD FADH2 Fumarate H2O 7 6 Acetyl CoA Oxaloacetate Citrate H2O Isocitrate NADH NAD+ + H+ CO2 α-Ketoglutarate Citric acid cycle CoA-SH CO2 NAD+ NADH + H+ ATP formation Succinyl CoA ADP GDPGTP P i ATP 5 4 1 CoA-SH 3 CoA-SH 2
  • 62. © 2014 Pearson Education, Inc. Figure 7.11-6 NADH NAD+ + H+ 8 Malate Succinate FAD FADH2 Fumarate H2O 7 6 Acetyl CoA Oxaloacetate Citrate H2O Isocitrate NADH NAD+ + H+ CO2 α-Ketoglutarate Citric acid cycle CoA-SH CO2 NAD+ NADH + H+ ATP formation Succinyl CoA ADP GDPGTP P i ATP 5 4 1 CoA-SH 3 CoA-SH 2
  • 63. © 2014 Pearson Education, Inc. Figure 7.11a CoA-SH Acetyl CoA Start: Acetyl CoA adds its two-carbon group to oxaloacetate, producing citrate; this is a highly exergonic reaction. Oxaloacetate Citrate Isocitrate H2O1 2
  • 64. © 2014 Pearson Education, Inc. Figure 7.11b Isocitrate 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. CO2 release CO2 release α-Ketoglutarate Succinyl CoA NAD+ NADH + H+ CO2 CO2 CoA-SH NAD+ NADH + H+ 3 4
  • 65. © 2014 Pearson Education, Inc. Figure 7.11c CoA-SH Redox reaction: Succinate is oxidized; FAD is reduced. Fumarate Succinate Succinyl CoA ATP formation ATP ADP GDPGTP FAD FADH2 P i 5 6
  • 66. © 2014 Pearson Education, Inc. Figure 7.11d Redox reaction: Malate is oxidized; NAD+ is reduced. Fumarate Malate Oxaloacetate H2O NAD+ + H+ NADH 7 8
  • 67.  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 © 2014 Pearson Education, Inc.
  • 68. 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 © 2014 Pearson Education, Inc.
  • 69. 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 © 2014 Pearson Education, Inc.
  • 70. © 2014 Pearson Education, Inc. Figure 7.UN09 Glycolysis Pyruvate oxidation Citric acid cycle Oxidative phosphorylation: electron transport and chemiosmosis ATP ATP ATP
  • 71. © 2014 Pearson Education, Inc. Figure 7.12 Multiprotein complexes (originally from NADH or FADH2) Freeenergy(G)relativetoO2(kcal/mol) 50 40 30 20 10 0 NADH NAD+ FADH2 FAD 2 2 e− e− FMN Fe•S Fe•S Q I II III Cyt b Cyt c1 Fe•S Cyt c IV Cyt a Cyt a3 2 e− O22 H+ + ½ H2O
  • 72. © 2014 Pearson Education, Inc. Figure 7.12a Multiprotein complexes Freeenergy(G)relativetoO2(kcal/mol) 50 40 30 20 10 NADH NAD+ FADH2 FAD 2 2 e− e− FMN Fe•S Fe•S Q I II III Cyt b Cyt c1 Fe•S Cyt c IV Cyt a Cyt a3 2 e−
  • 73. © 2014 Pearson Education, Inc. Figure 7.12b 30 20 10 0 Cyt c1 Cyt c IV Cyt a Cyt a3 2 e− Freeenergy(G)relativetoO2(kcal/mol) (originally from NADH or FADH2) 2 H+ + ½ O2 H2O
  • 74.  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 © 2014 Pearson Education, Inc.
  • 75. 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 © 2014 Pearson Education, Inc.
  • 76. © 2014 Pearson Education, Inc. Video: ATP Synthase 3-D Side View Video: ATP Synthase 3-D Top View
  • 77. © 2014 Pearson Education, Inc. Figure 7.13 INTERMEMBRANE SPACE MITOCHONDRIAL MATRIX Rotor Internal rod Catalytic knob Stator H+ ATP ADP P i +
  • 78.  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 © 2014 Pearson Education, Inc.
  • 79. © 2014 Pearson Education, Inc. Figure 7.UN09 Glycolysis Pyruvate oxidation Citric acid cycle Oxidative phosphorylation: electron transport and chemiosmosis ATP ATP ATP
  • 80. © 2014 Pearson Education, Inc. Figure 7.14 Protein complex of electron carriers H+ H+ H+ H+ Q I II III FADH2 FAD NAD+ NADH (carrying electrons from food) Electron transport chain Oxidative phosphorylation Chemiosmosis ATP synthase H+ ADP + ATPP i H2O2 H+ + ½ O2 IV Cyt c 1 2
  • 81. © 2014 Pearson Education, Inc. Figure 7.14a Protein complex of electron carriers H+ Q I II III FADH2 FAD NAD+ NADH (carrying electrons from food) Electron transport chain H2O2 H+ + ½ O2 Cyt c 1 IV H+ H+
  • 82. © 2014 Pearson Education, Inc. Figure 7.14b ATP synthase Chemiosmosis2 H+ H+ ADP + P i ATP
  • 83. 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 © 2014 Pearson Education, Inc.
  • 84. © 2014 Pearson Education, Inc. Figure 7.15 Electron shuttles span membrane CYTOSOL 2 NADH 2 NADH 2 FADH2 or 2 NADH Glycolysis Glucose 2 Pyruvate Pyruvate oxidation 2 Acetyl CoA Citric acid cycle 6 NADH 2 FADH2 Oxidative phosphorylation: electron transport and chemiosmosis + about 26 or 28 ATP+ 2 ATP + 2 ATP About 30 or 32 ATP Maximum per glucose: MITOCHONDRION
  • 85. © 2014 Pearson Education, Inc. Figure 7.15a Electron shuttles span membrane 2 NADH 2 FADH2 or 2 NADH Glycolysis Glucose 2 Pyruvate + 2 ATP
  • 86. © 2014 Pearson Education, Inc. Figure 7.15b 2 NADH 6 NADH 2 FADH2 Citric acid cycle Pyruvate oxidation 2 Acetyl CoA + 2 ATP
  • 87. © 2014 Pearson Education, Inc. Figure 7.15c 2 NADH 2 NADH 6 NADH 2 FADH2 2 FADH2 or Oxidative phosphorylation: electron transport and chemiosmosis + about 26 or 28 ATP
  • 88. © 2014 Pearson Education, Inc. Figure 7.15d Maximum per glucose: About 30 or 32 ATP
  • 89. 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 © 2014 Pearson Education, Inc.
  • 90.  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 © 2014 Pearson Education, Inc.
  • 91. 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 © 2014 Pearson Education, Inc.
  • 92.  In alcohol fermentation, pyruvate is converted to ethanol in two steps  The first step releases CO2 from pyruvate, and the second step reduces acetaldehyde to ethanol  Alcohol fermentation by yeast is used in brewing, winemaking, and baking © 2014 Pearson Education, Inc. Animation: Fermentation Overview
  • 93. © 2014 Pearson Education, Inc. Figure 7.16 2 ADP + 2 2 ATPP i Glucose Glycolysis 2 Pyruvate 2 CO2 2 NADH + 2 H+ 2 NAD+ 2 Ethanol (a) Alcohol fermentation 2 Acetaldehyde (b) Lactic acid fermentation 2 Lactate 2 NADH + 2 H+ 2 NAD+ 2 Pyruvate Glycolysis 2 ATP2 ADP + 2 P i Glucose
  • 94. © 2014 Pearson Education, Inc. Figure 7.16a 2 ADP + 2 2 ATPP i Glucose Glycolysis 2 Pyruvate 2 CO2 2 NADH + 2 H+ 2 NAD+ 2 Ethanol (a) Alcohol fermentation 2 Acetaldehyde
  • 95. © 2014 Pearson Education, Inc. Figure 7.16b 2 ADP + 2 2 ATPP i Glucose Glycolysis 2 Pyruvate 2 NADH + 2 H+ 2 NAD+ (b) Lactic acid fermentation 2 Lactate
  • 96.  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 © 2014 Pearson Education, Inc.
  • 97. 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 © 2014 Pearson Education, Inc.
  • 98.  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 © 2014 Pearson Education, Inc.
  • 99. © 2014 Pearson Education, Inc. Figure 7.17 Glucose CYTOSOL Glycolysis Pyruvate O2 present: Aerobic cellular respiration No O2 present: Fermentation Ethanol, lactate, or other products Acetyl CoA Citric acid cycle MITOCHONDRION
  • 100. 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 © 2014 Pearson Education, Inc.
  • 101. Concept 7.6: 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 © 2014 Pearson Education, Inc.
  • 102. 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 © 2014 Pearson Education, Inc.
  • 103.  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 © 2014 Pearson Education, Inc.
  • 104. © 2014 Pearson Education, Inc. Figure 7.18-1 Proteins Amino acids Carbohydrates Sugars Fats Glycerol Fatty acids
  • 105. © 2014 Pearson Education, Inc. Figure 7.18-2 Proteins Amino acids Carbohydrates Sugars Glucose Glycolysis Glyceraldehyde 3- Pyruvate P NH3 Fats Glycerol Fatty acids
  • 106. © 2014 Pearson Education, Inc. Figure 7.18-3 Proteins Amino acids Carbohydrates Sugars Glucose Glycolysis Glyceraldehyde 3- Pyruvate P Acetyl CoA NH3 Fats Glycerol Fatty acids
  • 107. © 2014 Pearson Education, Inc. Figure 7.18-4 Proteins Amino acids Carbohydrates Sugars Glucose Glycolysis Glyceraldehyde 3- Pyruvate P Acetyl CoA Citric acid cycle NH3 Fats Glycerol Fatty acids
  • 108. © 2014 Pearson Education, Inc. Figure 7.18-5 Proteins Amino acids Carbohydrates Sugars Glucose Glycolysis Glyceraldehyde 3- Pyruvate P Acetyl CoA Citric acid cycle NH3 Fats Glycerol Fatty acids Oxidative phosphorylation
  • 109. 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 © 2014 Pearson Education, Inc.
  • 110. © 2014 Pearson Education, Inc. Figure 7.UN10a
  • 111. © 2014 Pearson Education, Inc. Figure 7.UN10b
  • 112. © 2014 Pearson Education, Inc. Figure 7.UN11 Inputs Glucose Glycolysis 2 Pyruvate + 2 Outputs ATP NADH+ 2
  • 113. © 2014 Pearson Education, Inc. Figure 7.UN12 Inputs 2 Pyruvate 2 Acetyl CoA 2 Oxaloacetate Citric acid cycle Outputs ATP CO2 2 6 2 8 NADH FADH2
  • 114. © 2014 Pearson Education, Inc. Figure 7.UN13a Protein complex of electron carriers INTERMEMBRANE SPACE MITOCHONDRIAL MATRIX (carrying electrons from food) NADH NAD+ FADH2 FAD Cyt c Q I II III IV 2 H+ + ½O2 H2O H+ H+ H+
  • 115. © 2014 Pearson Education, Inc. Figure 7.UN13b INTER- MEMBRANE SPACE MITO- CHONDRIAL MATRIX ATP synthase ATPADP + H+ H+ P i
  • 116. © 2014 Pearson Education, Inc. Figure 7.UN14 Time pHdifference acrossmembrane

Notas del editor

  1. Figure 7.1 How do these leaves power the work of life for this giraffe?
  2. Figure 7.2 Energy flow and chemical recycling in ecosystems
  3. Figure 7.UN01 In-text figure, NaCl redox reaction, p. 136
  4. Figure 7.UN02 In-text figure, generalized redox reaction, p. 136
  5. Figure 7.3 Methane combustion as an energy-yielding redox reaction
  6. Figure 7.UN03 In-text figure, cellular respiration redox reaction, p. 137
  7. Figure 7.4 NAD+ as an electron shuttle
  8. Figure 7.4a NAD+ as an electron shuttle (part 1: NAD+ structure)
  9. Figure 7.4b NAD+ as an electron shuttle (part 2: NAD+ reaction)
  10. Figure 7.UN04 In-text figure, dehydrogenase reaction, p. 138
  11. Figure 7.5 An introduction to electron transport chains
  12. Figure 7.UN05 In-text figure, color code for stages of cellular respiration, p. 139
  13. Figure 7.6-1 An overview of cellular respiration (step 1)
  14. Figure 7.6-2 An overview of cellular respiration (step 2)
  15. Figure 7.6-3 An overview of cellular respiration (step 3)
  16. Figure 7.7 Substrate-level phosphorylation
  17. Figure 7.UN06 In-text figure, mini-map, glycolysis, p. 140
  18. Figure 7.8 The energy input and output of glycolysis
  19. Figure 7.9a A closer look at glycolysis (part 1: investment phase)
  20. Figure 7.9aa-1 A closer look at glycolysis (part 1a, step 1)
  21. Figure 7.9aa-2 A closer look at glycolysis (part 1a, step 2)
  22. Figure 7.9aa-3 A closer look at glycolysis (part 1a, step 3)
  23. Figure 7.9ab-1 A closer look at glycolysis (part 1b, step 1)
  24. Figure 7.9ab-2 A closer look at glycolysis (part 1b, step 2)
  25. Figure 7.9ab-3 A closer look at glycolysis (part 1b, step 3)
  26. Figure 7.9b A closer look at glycolysis (part 2: payoff phase)
  27. Figure 7.9ba-1 A closer look at glycolysis (part 2a, step 1)
  28. Figure 7.9ba-2 A closer look at glycolysis (part 2a, step 2)
  29. Figure 7.9ba-3 A closer look at glycolysis (part 2a, step 3)
  30. Figure 7.9bb-1 A closer look at glycolysis (part 2b, step 1)
  31. Figure 7.9bb-2 A closer look at glycolysis (part 2b, step 2)
  32. Figure 7.9bb-3 A closer look at glycolysis (part 2b, step 3)
  33. Figure 7.UN07 In-text figure, mini-map, pyruvate oxidation, p. 142
  34. Figure 7.10 An overview of pyruvate oxidation and the citric acid cycle
  35. Figure 7.10a An overview of pyruvate oxidation and the citric acid cycle (part 1: pyruvate oxidation)
  36. Figure 7.10b An overview of pyruvate oxidation and the citric acid cycle (part 2: citric acid cycle)
  37. Figure 7.UN08 In-text figure, mini-map, citric acid cycle, p. 143
  38. Figure 7.11-1 A closer look at the citric acid cycle (steps 1-2)
  39. Figure 7.11-2 A closer look at the citric acid cycle (step 3)
  40. Figure 7.11-3 A closer look at the citric acid cycle (step 4)
  41. Figure 7.11-4 A closer look at the citric acid cycle (step 5)
  42. Figure 7.11-5 A closer look at the citric acid cycle (steps 6-7)
  43. Figure 7.11-6 A closer look at the citric acid cycle (step 8)
  44. Figure 7.11a A closer look at the citric acid cycle (part 1)
  45. Figure 7.11b A closer look at the citric acid cycle (part 2)
  46. Figure 7.11c A closer look at the citric acid cycle (part 3)
  47. Figure 7.11d A closer look at the citric acid cycle (part 4)
  48. Figure 7.UN09 In-text figure, mini-map, oxidative phosphorylation, p. 144
  49. Figure 7.12 Free-energy change during electron transport
  50. Figure 7.12a Free-energy change during electron transport (part 1)
  51. Figure 7.12b Free-energy change during electron transport (part 2)
  52. Figure 7.13 ATP synthase, a molecular mill
  53. Figure 7.UN09 In-text figure, mini-map, oxidative phosphorylation, p. 146
  54. Figure 7.14 Chemiosmosis couples the electron transport chain to ATP synthesis
  55. Figure 7.14a Chemiosmosis couples the electron transport chain to ATP synthesis (part 1: electron transport chain)
  56. Figure 7.14b Chemiosmosis couples the electron transport chain to ATP synthesis (part 2: chemiosmosis)
  57. Figure 7.15 ATP yield per molecule of glucose at each stage of cellular respiration
  58. Figure 7.15a ATP yield per molecule of glucose at each stage of cellular respiration (part 1: glycolysis)
  59. Figure 7.15b ATP yield per molecule of glucose at each stage of cellular respiration (part 2: citric acid cycle)
  60. Figure 7.15c ATP yield per molecule of glucose at each stage of cellular respiration (part 3: oxidative phosphorylation)
  61. Figure 7.15d ATP yield per molecule of glucose at each stage of cellular respiration (part 4: yield per glucose)
  62. Figure 7.16 Fermentation
  63. Figure 7.16a Fermentation (part 1: alcohol)
  64. Figure 7.16b Fermentation (part 2: lactic acid)
  65. Figure 7.17 Pyruvate as a key juncture in catabolism
  66. Figure 7.18-1 The catabolism of various molecules from food (step 1)
  67. Figure 7.18-2 The catabolism of various molecules from food (step 2)
  68. Figure 7.18-3 The catabolism of various molecules from food (step 3)
  69. Figure 7.18-4 The catabolism of various molecules from food (step 4)
  70. Figure 7.18-5 The catabolism of various molecules from food (step 5)
  71. Figure 7.UN10a Skills exercise: making a bar graph and interpreting the data (part 1)
  72. Figure 7.UN10b Skills exercise: making a bar graph and interpreting the data (part 2)
  73. Figure 7.UN11 Summary of key concepts: pyruvate
  74. Figure 7.UN12 Summary of key concepts: citric acid cycle
  75. Figure 7.UN13a Summary of key concepts: oxidative phosphorylation (part 1)
  76. Figure 7.UN13b Summary of key concepts: oxidative phosphorylation (part 2)
  77. Figure 7.UN14 Test your understanding, question 8 (pH vs. time)