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Chapter 13 & 14: Respiration

glycolysis
fermentation
the mitochondrion
the citric acid cycle
the electron transport chain and oxidative phosphorylation
ATP synthesis
4th ed: p. 417-435; 451-465
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(3rd ed: p. 427-433; 436-443; 456-464; 466-467)

Summary of
glycolysis

Summary of Glycolysis
1 Glucose (6C)
All reactions occur
initial energy investment
in the cytoplasm.
(-2 ATPs)
1 Fructose 1,6-bisphosphate (6C)
cleavage
2 Glyceraldehyde 3-phosphate (3C)
energy generation
- by the oxidation of 2 G3P => 2 NADH
- by substrate-level phosphorylation (the transfer of
a phosphate from a sugar intermediate to ADP)
=> 4 ATP
2 Pyruvate (3C)
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Glycolysis: step by step

Glucose (6C)
1) Phosphorylation
of glucose

ATP consumed
phosphorylation of C6

Glucose 6-phosphate (6C)


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Glycolysis: step by step

2) Isomerization of G6P
Fructose 6-phosphate (6C)
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Glycolysis: step by step

3) Phosphorylation
of F6P

ATP consumed
phosphorylation of C1

Fructose 1,6-bisphosphate (6C)


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Glycolysis: step by step

4) Cleavage of FBP
Dihydroxyacetone phosphate (3C)
+ Glyceraldehyde 3-phosphate (3C)
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Glycolysis: step by step

5) Isomerization of DHP
(2) Glyceraldehyde 3-phosphate (3C)
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Glycolysis: step by step

6) Oxidation of G3P

2NAD+ reduced to 2NADH


Pi attached to C1

(2) 1,3-bisphosphoglycerate (3C)


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Glycolysis: step by step

7) Transfer of 1P from
BPG to ADP

2ATP synthesized by
substrate-level
phosphorylation

(2) 3-Phosphoglycerate (3C)


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Glycolysis: step by step

8) Isomerization of 3PG
(2) 2-Phosphoglycerate (3C)
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Glycolysis: step by step

9) Dehydration of 2PG

2H2O removed

(2) Phosphoenolpyruvate (3C)


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Glycolysis: step by step

10) Transfer of P from


PEP to ADP

2ATP synthesized by
substrate-level
phosphorylation

(2) Pyruvate (3C)


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Glycolysis

Yield of glycolysis:
2 ATP consumed at the energy investment phase
4 ATP and 2 NADH produced at the energy generation phase
Net yield: 2 ATP + 2 NADH
Under anaerobic conditions (no O2 present):
pyruvate and NADH are consumed in fermentation
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+
NAD is regenerated and used in glycolysis (step 6)

Fermentation
In the absence of oxygen, pyruvate is reduced, driven by the
oxidation of the NADH generated from glycolysis.
In muscle cells

In yeast

2 Pyruvate (3C)

2 Pyruvate (3C)

NADH => NAD+


2 Lactic acid (3C)

CO2 removed
2 Acetaldehyde (2C)
NADH => NAD+
2 Ethanol (2C)

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Net yield of fermentation: 2 ATP

Fermentation leading to
excretion of lactate
In muscle cells

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Fermentation leading to
excretion of ethanol and CO2
In yeast

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Glycolysis
Under aerobic conditions (O2 present):
Pyruvate and NADH

to the mitochondria

pyruvate is converted to acetyl coenzyme A


and enters the citric acid cycle
NADH is used to power the oxidative phosphorylation
reactions (the electron transport chain)

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The mitochondrion

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The mitochondrion
The mitochondrion (plural mitochondria) is a membraneenclosed organelle of eukaryotic cells that generates most
of the cell's supply of ATP.
enclosed by a double membrane
the matrix is the site of the oxidation of pyruvate
and the citric acid cycle
the inner membrane is the site of the electron transport
chain
and oxidative phosphorylation

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Matrix

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The oxidation of pyruvate


Pyruvate + CoenzymeA Acetyl CoA + CO2

Pyruvate (3C) + CoA


CO2 is removed by hydrolysis, driving
NAD+ => NADH, and
the condensation with CoA
Acetyl CoA (2C)

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Citric Acid Cycle


(The Krebs Cycle)

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The Citric Acid Cycle: step by step


Acetyl CoA (2C) + Oxaloacetate (4C)
1) Condensation and
citrate synthesis

driven by the hydrolysis


that removes CoA

Citric acid (6C)


2) Isomerization of CA
Isocitrate (6C)
3) Oxidation of IC

CO2 removed
NAD+ reduced to NADH

alpha-ketoglutarate (5C)
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The Citric Acid Cycle: step by step


4) Oxidation of KG

CO2 removed
NAD+ reduced to NADH
CoA added

Succinyl CoA (4C)


CoA removed
GTP synthesis
5) Hydrolysis of SCoA
GTP is hydrolyzed to GDP
ATP synthesis
Succinate (4C)
6) Oxidation of SA

FAD reduced to FADH2

Fumarate (4C)

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Succinate
HS-CoA

GTP
ADP

Succinyl CoA
GDP
ATP

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The Citric Acid Cycle: step by step


7) Hydrogenation of FA

H2O consumed

Malate (4C)
8) Oxidation of MA

NAD+ reduced to NADH

Oxaloacetate (4C)

Re-enters the cycle

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Yield of citric acid cycle


2 CO2
(per one turn)

4 CO2
(two pyruvates)

3 NADH
1 FADH2
1 ATP

6 NADH
2 FADH2
2 ATP

X2

ATP

NADH and FADH2


to the e- transport chain
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Matrix

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Oxidation of NADH transfers


high-energy electrons
to the electron transport chain

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Oxidative phosphorylation:
The electron transport chain

The respiratory enzyme complexes in the inner membrane


pump H+ from the matrix into the intermembrane space,
generating a H+ gradient across the inner 31membrane.

Oxidative phosphorylation:
The electron transport chain

the flow of H+ back into the matrix through ATP


synthase powers ATP synthesis
the energy that powers the H+ pumps is provided by the
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loss of electrons (oxidation) of NADH and FADH2

The electron transport chain


NADH is oxidized
H+ pumps are powered

e- lose energy
final e- acceptor:
1
/2O2 + 2H+
=> H2O

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The electron transport chain


The respiratory enzyme complexes in the inner membrane
contain a series of electron carriers (co-factors).

the electrons move through these carriers


each carrier becomes sequentially reduced and oxidized
H+ pumps are powered
e- lose energy

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The electron transport chain

e- transfer to A

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The electron transport chain

e- transfer from B
H+ are picked up from one side of the membrane (the matrix)
and released to the other (into the intermembrane
space).
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The electron transport chain


as an e- is transferred from
A to B (B is reduced),
a H+ is picked up from
the matrix
as an e- is transferred from
B to C (B is oxidized),
the H+ is released into
the intermembrane space

Result: a H+ gradient across


the inner membrane
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The electron transport chain:


Cytochrome oxidase complex

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The electron transport chain


Transport of NADH electrons
1- NADH dehydrogenase complex (I): accepts e2- Cytochrome b-c1 complex (III)
3- Cytochrome oxidase complex (IV)
Transport of FADH2 electrons
(not illustrated)
1- Fe-S complex (II): accepts e- from FADH2
2- Cytochrome b-c1 complex (III)
3- Cytochrome oxidase complex (IV)
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Electron transport chain:


Transport of NADH electrons

III

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IV

Synthesis of ATP
The H+ gradient generated by the e- transport chain powers
ATP synthase complex.
as H+ flow back across the inner mitochondrial
membrane into the matrix, the central rotor unit
rotates,
powering ATP synthesis (stator: stationary unit)
Result: oxidative phosphorylation of ADP to ATP
(coupled to redox reactions of ETC)

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Synthesis of ATP

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ATP yield of respiration


per molecule of glucose:
ATP from glycolysis and citric acid cycle: _
ATP from oxidative phosphorylation:
NADH from glycolysis: _
NADH from the oxidation of pyruvate: _
NADH from the citric acid cycle: _
FADH2 from the citric acid cycle: _
2.5 ATP / NADH: 25
1.5 ATP / FADH2: 3
Total: 32 ATP
In most cells: the 2 NADH generated during glycolysis shuttle
electrons to FADH2 in mitochondria: yield 1.5 ATP each
Actual net yield: 30 ATP 43

ATP yield of fermentation


per molecule of glucose:
ATP from glycolysis: _
NADH from glycolysis: _
But pyruvate is reduced in fermentation
and NADH is oxidized to regenerate NAD+ to
be re-used in another round of glycolysis
Total: _ ATP
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