Cellular Respiration
Cellular Respiration
Biology
Eighth Edition
Neil Campbell and Jane Reece
Lectures by Chris Romero, updated by Erin Barley with contributions from Joan Sharp
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Overview: Life Is Work
Light
energy
ECOSYSTEM
Photosynthesis
in chloroplasts
CO2 + H2O Organic + O
molecules 2
Cellular respiration
in mitochondria
ATP
Heat
energy
Catabolic Pathways and Production of ATP
becomes reduced
(gains electron)
The electron donor is called the reducing agent
The electron receptor is called the oxidizing
agent
Some redox reactions do not transfer electrons
but change the electron sharing in covalent bonds
Reactants Products
becomes oxidized
becomes reduced
becomes oxidized
becomes reduced
Dehydrogenase
Stepwise Energy Harvest via NAD+ and the Electron
Transport Chain
Electrons from organic compounds are usually
first transferred to NAD+ (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
2 e– + 2 H+
2 e– + H+
NADH H+
Dehydrogenase
Reduction of NAD+
NAD+ + 2[H] + H+
Oxidation of NADH
Nicotinamide
(reduced form)
Nicotinamide
(oxidized form)
NADH passes the electrons to the electron
transport chain…ETC
Unlike an uncontrolled reaction, the electron
transport chain passes electrons in a series of
steps instead of one explosive reaction
O2 (high electro negativity) pulls electrons down
the chain in an energy-yielding tumble
The energy yielded is used to regenerate ATP
from ADP and inorganic Phosphate
The Stages of Cellular Respiration: A Preview
Electrons
carried
via NADH
Glycolysis
Glucose Pyruvate
Cytosol
ATP
Substrate-level
phosphorylation
Fig. 9-6-2
Glycolysis Citric
acid
Glucose Pyruvate cycle
Mitochondrion
Cytosol
ATP ATP
Substrate-level Substrate-level
phosphorylation phosphorylation
Fig. 9-6-3
Oxidative
Glycolysis Citric phosphorylation:
acid electron transport
Glucose Pyruvate cycle and
chemiosmosis
Mitochondrion
Cytosol
Enzyme Enzyme
ADP
P
Substrate + ATP
Product
Concept 9.2: Glycolysis harvests chemical energy
by oxidizing glucose to pyruvate
• Glycolysis (“splitting of sugar”) breaks down
glucose into two molecules of pyruvate
• Glycolysis occurs in the cytoplasm and has two
major phases:
– Energy investment phase
– Energy payoff phase
Glucose
2 NAD+ + 4 e– + 4 H+ 2 NADH + 2 H+
2 Pyruvate + 2 H2O
Net
Glucose 2 Pyruvate + 2 H2O
4 ATP formed – 2 ATP used 2 ATP
2 NAD+ + 4 e– + 4 H+ 2 NADH + 2 H+
Fig. 9-9-1
Glucose
ATP
1
Hexokinase
ADP
Glucose
Glucose-6-phosphate
ATP
1
Hexokinase
ADP
Glucose-6-phosphate
Fig. 9-9-2
Glucose
ATP
1
Hexokinase
ADP
Glucose-6-phosphate
Glucose-6-phosphate
2
Phosphoglucoisomerase
2
Fructose-6-phosphate
Phosphogluco-
isomerase
Fructose-6-phosphate
Fig. 9-9-3
Glucose
ATP
1
Hexokinase
ADP
Fructose-6-phosphate
Glucose-6-phosphate
2
Phosphoglucoisomerase
ATP
3
Fructose-6-phosphate
Phosphofructo-
ATP
kinase
3
Phosphofructokinase
ADP
ADP
Fructose-
1, 6-bisphosphate
Fructose-
1, 6-bisphosphate
Fig. 9-9-4
Glucose
ATP
1
Hexokinase
ADP
Glucose-6-phosphate
2
Phosphoglucoisomerase
Fructose-
1, 6-bisphosphate
4
Fructose-6-phosphate
Aldolase
ATP
3
Phosphofructokinase
ADP
5
Isomerase
Fructose-
1, 6-bisphosphate
4
Aldolase
5
Isomerase
Dihydroxyacetone Glyceraldehyde-
phosphate 3-phosphate
Dihydroxyacetone Glyceraldehyde-
phosphate 3-phosphate
Fig. 9-9-5
2 NAD+ 6
Triose phosphate
dehydrogenase
2 NADH 2 Pi
+ 2 H+
2 1, 3-Bisphosphoglycerate
Glyceraldehyde-
3-phosphate
2 NAD+ 6
Triose phosphate
dehydrogenase
2 NADH 2 Pi
+ 2 H+
2 1, 3-Bisphosphoglycerate
Fig. 9-9-6
2 NAD+ 6
Triose phosphate
dehydrogenase
2 NADH 2 Pi
+ 2 H+
2 1, 3-Bisphosphoglycerate
2 ADP
7
Phosphoglycerokinase
2 ATP
2 1, 3-Bisphosphoglycerate
2 ADP
2 3-Phosphoglycerate
7
Phosphoglycero-
2 ATP kinase
2 3-Phosphoglycerate
Fig. 9-9-7
2 NAD+ 6
Triose phosphate
dehydrogenase
2 NADH 2 Pi
+ 2 H+
2 1, 3-Bisphosphoglycerate
2 ADP
7 Phosphoglycerokinase
2 ATP
3-Phosphoglycerate
2 3-Phosphoglycerate
8
2
Phosphoglyceromutase
8
Phosphoglycero-
2 2-Phosphoglycerate mutase
2 2-Phosphoglycerate
Fig. 9-9-8
2 NAD+ 6
Triose phosphate
dehydrogenase
2 NADH 2 Pi
+ 2 H+
2 1, 3-Bisphosphoglycerate
2 ADP
7 Phosphoglycerokinase
2 ATP
2 3-Phosphoglycerate 2 2-Phosphoglycerate
8
Phosphoglyceromutase
9
Enolase
2 H2O
2 2-Phosphoglycerate
9
Enolase
2 H2O
2 Phosphoenolpyruvate
2 Phosphoenolpyruvate
Fig. 9-9-9
2 NAD+ 6
Triose phosphate
dehydrogenase
2 NADH 2 Pi
+ 2 H+
2 1, 3-Bisphosphoglycerate
2 ADP
7 Phosphoglycerokinase
2 ATP
2 Phosphoenolpyruvate
2 ADP
2 3-Phosphoglycerate
8
Phosphoglyceromutase 10
Pyruvate
2 ATP kinase
2 2-Phosphoglycerate
9
Enolase
2 H2O
2 Phosphoenolpyruvate
2 ADP
10
Pyruvate kinase
2 ATP
2 Pyruvate
2 Pyruvate
CITRIC ACID CYCLE
KREBS CYCLE /
TRICARBOXYLIC ACID/ TCA
CYCLE
35
Coenzyme A
37
SALIENT FEATURES OF CITRIC ACID
CYCLE/KREB’S CYCLE
▪ The final common oxidative pathway for carbohydrate, fats
and amino acids.
▪ The cycle operates only under aerobic conditions.
▪ The enzymes of TCA cycle are located in mitochondrial
matrix.
▪ It involves the combination of a two carbon acetyl CoA with
a four carbon oxaloacetate to produce a six carbon
tricarboxvlic acid, citrate
▪ Oxaloacetate is considered to play a catalytic role in. citric
acid cycle
Acetyl CoA + 3 NAD+ + FAD + GDP + Pi +2H2O 2CO2 + 3NADH +
3H+ + FADH2 + GTP + CoA
Citric Acid Cycle/Kreb’s
Cycle Process
Concept 9.3: The citric acid cycle completes the
energy-yielding oxidation of organic molecules
• In the presence of O2, pyruvate enters the
mitochondrion
• Before the citric acid cycle can begin, pyruvate
must be converted to acetyl CoA, which links
the cycle to glycolysis
CYTOSOL MITOCHONDRION
NAD+ NADH + H+
1 3
Acetyl CoA
Pyruvate CO2 Coenzyme A
Transport protein
• The citric acid cycle, also called the Krebs
cycle, takes place within the mitochondrial
matrix
• The cycle oxidizes organic fuel derived from
pyruvate, generating 1 ATP, 3 NADH, and 1
FADH2 per turn
Pyruvate
CO2
NAD+
CoA
NADH
+ H+ Acetyl CoA
CoA
CoA
Citric
acid
cycle 2 CO2
FADH2 3 NAD+
FAD 3 NADH
+ 3 H+
ADP + P i
ATP
• 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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Fig. 9-12-1
Acetyl CoA
CoA—SH
Oxaloacetate
Citrate
Citric
acid
cycle
Fig. 9-12-2
Acetyl CoA
CoA—SH
1 H2O
Oxaloacetate
2
Citrate
Isocitrate
Citric
acid
cycle
Fig. 9-12-3
Acetyl CoA
CoA—SH
1 H2O
Oxaloacetate
2
Citrate
Isocitrate
NAD+
Citric NADH
3
acid + H+
cycle
CO2
-Keto-
glutarate
Fig. 9-12-4
Acetyl CoA
CoA—SH
1 H2O
Oxaloacetate
2
Citrate
Isocitrate
NAD+
Citric NADH
acid 3
+ H+
cycle
CO2
CoA—SH
-Keto-
glutarate
4
CO2
NAD+
NADH
Succinyl + H+
CoA
Fig. 9-12-5
Acetyl CoA
CoA—SH
1 H2O
Oxaloacetate
2
Citrate
Isocitrate
NAD+
Citric NADH
3
acid + H+
cycle
CO2
CoA—SH
-Keto-
glutarate
4
CoA—SH
5
CO2
NAD+
Succinate Pi NADH
GTP GDP Succinyl + H+
CoA
ADP
ATP
Fig. 9-12-6
Acetyl CoA
CoA—SH
1 H2O
Oxaloacetate
2
Citrate
Isocitrate
NAD+
Citric NADH
3
acid + H+
cycle
CO2
Fumarate CoA—SH
-Keto-
glutarate
6 4
CoA—SH
FADH2 5
CO 2
NAD+
FAD
Succinate Pi NADH
GTP GDP Succinyl + H+
CoA
ADP
ATP
Fig. 9-12-7
Acetyl CoA
CoA—SH
1 H2O
Oxaloacetate
2
Malate Citrate
Isocitrate
NAD+
Citric 3
NADH
7
acid + H+
H2O cycle
CO2
Fumarate CoA—SH
-Keto-
glutarate
6 4
CoA—SH
FADH2 5
CO 2
NAD+
FAD
Succinate Pi NADH
GTP GDP Succinyl + H+
CoA
ADP
ATP
Fig. 9-12-8
Acetyl CoA
CoA—SH
NADH
+H+ 1 H2O
NAD+
8 Oxaloacetate
2
Malate Citrate
Isocitrate
NAD+
Citric 3
NADH
acid + H+
7
H2O cycle
CO2
Fumarate CoA—SH
-Keto-
glutarate
6 4
CoA—SH
FADH2 5
CO 2
NAD+
FAD
Succinate Pi NADH
GTP GDP Succinyl + H+
CoA
ADP
ATP
Concept 9.4: During oxidative phosphorylation,
chemiosmosis couples electron transport to ATP
synthesis
NADH
50
2 e– NAD+
FADH2
2 e– FAD
Multiprotein
40 FAD complexes
FMN
Fe•S Fe•S
Q
Cyt b
Fe•S
30
Cyt c1 I
Cyt c V
Cyt a
Cyt a3
20
10 2 e–
(from NADH
or FADH2)
0 2 H+ + 1/2 O2
H2O
• Electrons are transferred from NADH or FADH2
to the electron transport chain
• Electrons are passed through a number of
proteins to O2
• The electron transport chain generates no ATP
• The chain’s function is to break the large free-
energy drop from food to O2 into smaller steps
that release energy in manageable amounts
H+
Stator
Rotor
Internal
rod
Cata-
lytic
knob
ADP
+
P ATP
i
MITOCHONDRIAL MATRIX
• The energy stored in a H+ gradient across a
membrane couples the redox reactions of the
electron transport chain to ATP synthesis
• The H+ gradient is referred to as a proton-
motive force, emphasizing its capacity to do
work
H+
H+
H+
H+
Protein complex Cyt c
of electron
carriers
V
Q
ATP
synthase
2 H+ + 1/2O2 H2O
FADH2
FAD
NADH NAD+
ADP + P i ATP
(carrying electrons
from food)
H+
Oxidative phosphorylation
An Accounting of ATP Production by Cellular
Respiration
• During cellular respiration, most energy flows in
this sequence:
glucose → NADH → electron transport chain
→ proton-motive force → ATP
Glycolysis Oxidative
2 2 Citric phosphorylation:
Glucose Pyruvate Acetyl acid electron transport
CoA cycle and
chemiosmosis
2 ADP + 2 Pi 2 ATP
Glucose Glycolysis
2 Pyruvate
2 Ethanol 2 Acetaldehyde
2 ADP + 2 Pi 2 ATP
Glucose Glycolysis
2 NAD+ 2 NADH
+ 2 H+
2 Pyruvate
2 Lactate
2 ADP + 2 P i 2 ATP
Glucose Glycolysis
2 Pyruvate
2 NAD+ 2 NADH 2 CO 2
+ 2 H+
2 Ethanol 2 Acetaldehyde
2 ADP + 2 P i 2 ATP
Glucose Glycolysis
2 NAD+ 2 NADH
+ 2 H+
2 Pyruvate
2 Lactate
Glycolysis
CYTOSOL
Pyruvate
No O2 present: O2 present:
Fermentation Aerobic cellular
respiration
MITOCHONDRION
Ethanol Acetyl CoA
or
lactate
Citric
acid
cycle
The Evolutionary Significance of Glycolysis
Glycolysis
Glucose
Glyceraldehyde-3- P
NH3 Pyruvate
Acetyl CoA
Citric
acid
cycle
Oxidative
phosphorylation
Biosynthesis (Anabolic Pathways)
AMP
Glycolysis
Fructose-6-phosphate Stimulates
+
Phosphofructokinase
–
–
Fructose-1,6-bisphosphate
Inhibits Inhibits
Pyruvate
ATP Citrate
Acetyl CoA
Citric
acid
cycle
Oxidative
phosphorylation
Fig. 9-UN5
Inputs Outputs
2 ATP
Glycolysis
+
2 NADH
Glucose 2 Pyruvate
Fig. 9-UN6
Inputs Outputs
S—CoA
2 ATP
C O
CH3
2 Acetyl CoA
6 NADH
O C COO
Citric acid
CH2 cycle
COO 2 FADH2
2 Oxaloacetate