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Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Chapter 9
Cellular Respiration: Harvesting Chemical Energy
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Life Is Work!• What kind of work is done by living cells?
– synthesis, active transport, movement, growth, reproduction, maintaining homeostasis
• Work requires energy – where does the energy come from?
– Cells harvest energy by using various metabolic pathways:
• Some pathways convert light energy to organic compounds = photosynthesis (that’s the next chapter!)
• Other pathways convert organic compounds to usable energy (ATP) and heat = respiration
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
The flow of energy• Energy flows into an ecosystem as sunlight
and leaves as heat
Light energy
ECOSYSTEM
CO2 + H2O
Photosynthesisin chloroplasts
Cellular respiration
in mitochondria
Organicmolecules+ O2
ATP
powers most cellular work
HeatenergyFigure 9.2
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Harvesting stored energy• Organic compounds store energy in their arrangement
of atoms.
• To harvest that energy, a cell systematically breaks down complex organic molecules that are rich in potential energy to simpler waste products that have less energy
• Some of this energy can be used to do work (= usable energy), the rest is lost as heat
Efficiency = ___amount of usable energy___ x 100total amount of energy released
(usable energy + heat)
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Harvesting stored energy• The chemical reactions involved are
catabolic pathways (metabolic pathways that release stored energy by breaking down complex molecules)
• Two major catabolic pathways:
o Fermentation – partial breakdown of sugar without oxygen
o Cellular respiration – complete breakdownof sugar with oxygen
o This is far more prevalent and efficient!
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Harvesting stored energy• Respiration operates on the same principle as
combustion (using different mechanism at much lower temperature)Summary of overall process:organic compounds + oxygen carbon dioxide + water + energy
• Carbohydrates, fats, and proteins can all be processed and used as fuel, but main pathways use carbohydrates (glucose):
C6H12O6 + 6O2 6CO2 + 6H2O + energy (ATP + heat)
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Harvesting stored energy• What is the important difference between
combustion and respiration?
– In combustion, all of the energy is released in one step (high heat & much energy is lost)
– In respiration, energy is released in many controlled steps to make ATP (less heat & less energy loss)
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Catabolic Pathways and Production of ATP• The breakdown of glucose is exergonic
– G = – 686 kcal/mole)
– remember that negative value means that the products have less energy than the reactants.
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Cellular respiration• Just as the energy released by combustion
of petroleum is transferred by the engine and drive shaft to make the wheels turn, respiration is linked to cellular work by a chemical drive shaft: ATP
• So what is the point of cellular respiration??
To make ATP!!
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The ATP Cycle• To keep working cells must regenerate ATP
• A working muscle cell recycles its ATP at a rate of about 10 million molecules per second!
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
ATP Review• Recall that the three negatively charged
phosphate groups is an unstable, energy-storing arrangement (like charges repel)
• The chemical “spring” tends to “relax” by losing the terminal phosphate
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Redox Reactions: Oxidation and Reduction• Catabolic pathways yield energy due to the
transfer of electrons
• Energy is released when electrons move closer to electronegative atoms
• Oxygen is the the one of the most electronegative elements around!
• Oxidation is the loss of electrons (becomes more positive)
• Reduction is the gain of electrons (becomes more negative)
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Redox Reactions: Oxidation and Reduction• In a chemical reaction, when one substance is
oxidized (loses electrons), another must be reduced (gains electrons)!
– the electrons have to go somewhere!
• Redox reactions are the transfer electrons from one reactant to another by oxidation and reduction
– The substance that is oxidized is the reducing agent (by giving its electrons to something)
– The substance that is reduced is the oxidizing agent (by taking the electrons from something)
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Redox Reactions: Oxidation and Reduction• Example of a redox reaction
Na + Cl Na+ + Cl–
becomes oxidized(loses electron)
becomes reduced(gains electron)
Oxidizing agent (takes electrons)
Reducing agent (gives up electrons)
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Redox Reactions: Oxidation and Reduction• Some redox reactions
– Do not completely exchange electrons
– Change the degree of electron sharing in covalent bonds of molecules
CH4
H
H
HH
C O O O O OC
H H
Methane(reducing
agent)
Oxygen(oxidizing
agent)
Carbon dioxide Water
+ 2O2CO
2+ Energy + 2 H2O
becomes oxidized
becomes reduced
Reactants Products
Figure 9.3
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Oxidation vs. Reduction
• Oxidation
– losing electrons
– removing H
– adding O2
– releases energy
– exergonic
– catabolic
• Reduction
– gaining electrons
– adding H
– removing O2
– stores energy
– endergonic
– anabolic
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Oxidation of Organic Fuel Molecules During Cellular Respiration
• During cellular respiration
– Glucose is oxidized and oxygen is reduced
C6H12O6 + 6O2 6CO2 + 6H2O + Energybecomes oxidized
becomes reduced
C6H12O6 O2 CO2 H2O0 +1 -2 0 +4 -2 +1 -2
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Stepwise energy harvest• During cellular respiration, electrons “fall” from
organic molecules to oxygen in a series of steps(= redox reactions)
• Organic molecules store energy because they have many C-H bonds which are a source of “hilltop” electrons which have the potential to “fall” closer to oxygen.
– Only the barrier of activation energy holds back the flood of electrons to a lower energy state. Without this barrier, glucose would combine spontaneously with O2
Body temperature is not enough to overcome this activation energy!
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Stepwise electron transfer: overview
• As hydrogen atoms are stripped from glucose during respiration, electrons are transferred by:
– NAD+ (“taxi”)
– Electron Transport Chain (ETC) (“water slide”)
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NAD: taxi to the electron transport chain• NAD+ (nicotinamide adenine dinucleotide) is a
coenzyme
• Since it accepts electrons, it functions as an oxidizing agent
NAD+
H
O
OO O–
OO O–
O
O
O
P
P
CH2
CH2
HO OHH
HHO OH
HO
H
H
N+
C NH2
HN
H
NH2
N
N
Nicotinamide(oxidized form)
NH2+ 2[H](from food)
Dehydrogenase
Reduction of NAD+
Oxidation of NADH
2 e– + 2 H+
2 e– + H+
NADH
OH H
N
C +
Nicotinamide(reduced form)
N
Figure 9.4
• Dehydrogenase transfers 2 electronsand 1 H+ to NAD+, forming NADH
• The other H+ is released to the cytosol
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Electron Transport Chain• NADH passes the
electrons to the electron transport chain
• The electron transport chain passes electrons in a series of stepsinstead of in one explosive reaction
• The ETC uses the energy from the electron transfer to make ATP!
2 H 1/2 O2
(from food via NADH)
2 H+ + 2 e–
2 H+
2 e–
H2O
1/2 O2
Controlled release of energy for
synthesis ofATP ATP
ATP
ATP
Free
ene
rgy,
G
(b) Cellular respiration
+
Figure 9.5 B
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Electron Transport Chain: Overview• The ETC consists of several carrier molecules,
mostly proteins, built into the inner membrane of amitochondrion.
• Electrons removed from food are “shuttled” by NADH (taxi!) to the “top” end of the chain.
• Electrons “fall” down the chain because each carrier molecule in the chain is more electronegative than the one before it.
• At the bottom end, oxygen (the most electronegative) captures these electrons along with hydrogen & forms water (water slide!).
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
General “flow” of electrons
Food NADH ETC O2
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
What if it’s not stepwise?• If electron transfer is not stepwise
• A large (explosive!) release of energy occurs
• Example: reaction of hydrogen and oxygen to form water
(a) Uncontrolled reaction
Free
ene
rgy,
G
H2O
Explosiverelease of
heat and lightenergy
Figure 9.5 A
H2 + 1/2 O2
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Overview of Cellular Respiration:
• Glycolysis
– Breaks down glucose into two molecules of pyruvate
• Krebs Cycle (citric acid cycle)
– Completes the breakdown of glucose
• Electron transport chain
– Series of redox reactions releases energy
– Energy is used to generate most of the ATP (by oxidative phosphorylation & chemiosmosis)
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Overview of Cellular Respiration
Figure 9.6
Electronscarried
via NADH
GlycolsisGlucos
ePyruvate
ATP
Substrate-levelphosphorylation
Electrons carried via NADH and
FADH2
Citric acid cycle
Oxidativephosphorylation
:electron
transport andchemiosmosis
ATPATP
Substrate-levelphosphorylation
Oxidativephosphorylation
MitochondrionCytosol
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Two ways to make ATP:
1. Substrate-level phosphorylation
2. Oxidative phosphorylation
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Substrate-level phosphorylation occurs during glycolysis and Krebs Cycle enzyme transfers a phosphate group from a
substrate molecule to ADP (instead of getting it as free Pi from the surroundings)
Figure 9.7
Enzyme Enzyme
ATP
ADP
Product
SubstrateP
+
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Oxidative phosphorylation
• almost 90% of ATP generated this way
• powered by series of electron transfers(= redox reactions) through NADH and FADH2 and electron transport chain
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
A closer look: Glycolysis• Glycolysis
– Means “splitting of sugar”
– Breaks down glucose (6-C) into pyruvate (3-C)
– Occurs in the cytoplasm of the cell
– Pathway consists of ten steps, each catalyzed by a specific enzyme
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A closer look: Glycolysis
• Glycolysis consists of two major phases:
Glycolysis Citricacidcycle
Oxidativephosphorylation
ATP ATP ATP
2 ATP
4 ATP
used
formed
Glucose
2 ATP + 2 P
4 ADP + 4 P
2 NAD+ + 4 e- + 4 H +
2 NADH + 2 H+
2 Pyruvate + 2 H2O
Energy investment phase
Energy payoff phase
Glucose 2 Pyruvate + 2 H2O
4 ATP formed – 2 ATP used 2 ATP
2 NAD+ + 4 e– + 4 H +
2 NADH
+ 2 H+
Figure 9.8
Energy investment phase
Energy payoff phase
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
A closer look at the energy investment phase
Dihydroxyacetonephosphate
Glyceraldehyde-3-phosphate
HH
H
HH
OHOH
HO HO
CH2OHH H
H
HO H
OHHO OH
P
CH2O P
H
OH
HO
HO
HHOCH2OH
P O CH2O CH2 O P
HOH HO
HOH
OP CH2
C OCH2OH
HCCHOHCH2
O
O P
ATP
ADPHexokinase
Glucose
Glucose-6-phosphate
Fructose-6-phosphate
ATP
ADP
Phosphoglucoisomerase
Phosphofructokinase
Fructose-1, 6-bisphosphate
Aldolase
Isomerase
Glycolysis
1
2
3
4
5
CH2OHOxidative
phosphorylation
Citricacidcycle
Figure 9.9 A
Cell “spends” 2 ATPs to phosphorylatesugars (enzymes are kinases!)
Other enzymes rearrange sugar molecules to make them more reactive
This is where sugar is actually split!
This is an important intermediatecompound called Glyceraldehyde-3-phosphate (a.k.a. PGAL)
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
A closer look at the energy investment phase
• Why phosphorylate the sugars first?
– Accomplishes 2 things:
• Traps sugar inside cell (charged phosphate can’t easily go through membrane)
• Makes sugar molecules more reactive (makes them easier to rearrange and split)
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A closer look at the energy payoff phase2 NAD+
NADH2+ 2 H+
Triose phosphatedehydrogenase
2 P i
2P C
CHOHO
P
O
CH2 O
2 O–
1, 3-Bisphosphoglycerate2 ADP
2 ATP
Phosphoglycerokinase
CH2 O P
2
CCHOH
3-Phosphoglycerate
Phosphoglyceromutase
O–
CC
CH2OH
H O P
2-Phosphoglycerate
2 H2O2 O–
Enolase
CC
OPO
CH2Phosphoenolpyruvate
2 ADP
2 ATPPyruvate kinase
O–
CC
OO
CH3
2
6
8
7
9
10
Pyruvate
O
Figure 9.8 B
Sugar is oxidizedas electrons and H+
are transferred to NAD+, forming NADH (a high energy compound). Very exergonic!
4 ATPs are produced by substrate-level phosphorylation
Net energy gain is 2 ATPs!
Glycolysis produces 2 pyruvate (3C
sugar) molecules –still contain much
energy!
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acetyl CoA• Before the citric acid cycle can begin,
pyruvate must first be converted to acetyl CoA, which links the cycle to glycolysis
CYTOSOL MITOCHONDRION
NADH + H+NAD+
2
31
CO2 Coenzyme APyruvate
Acetyle CoA
S CoA
C
CH3
O
Transport protein
O–
O
O
C
C
CH3
Figure 9.10
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Formation of acetyl CoA • One carbon is removed (carboxyl group; no
energy left in it), releasing CO2
• Remaining 2-carbon fragment is oxidized to form acetate (ionized form of acetic acid)
• Coenzyme A (sulfur containing; derived from a B vitamin) is attached, forming an unstable molecule with a very reactive acetyl group
• Coenzyme A acts as a shuttle that feeds the 2-carbon acetyl group into the Krebs cycle
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An overview of the citric acid cycle
Glycolysis is linked to Krebs cycle by formation of Acetyl CoA!
ATP
2 CO2
3 NAD+
3 NADH
+ 3 H+
ADP + P i
FAD
FADH2
Citricacidcycle
CoA
CoAAcetyle CoA
NADH+ 3 H+
CoA
CO2
Pyruvate(from glycolysis,2 molecules per glucose)
ATP ATP ATP
Glycolysis Citricacidcycle
Oxidativephosphorylatio
n
Figure 9.11
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A closer look at Citric Acid (Krebs) Cycle• The citric acid cycle
– takes place in the matrix of the mitochondrion
– completes the energy-yielding oxidation of organic molecules
– has 8 steps, each catalyzed by a specific enzyme
• Each “turn” of the cycle:
• Releases 2 CO2
• Generates 1 ATP, 3 NADH, and 1 FADH2
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Hans Krebs
(1900-1981)
In case you’re curious . . .
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Figure 9.12
Acetyl CoA
NADH
Oxaloacetate
CitrateMalate
Fumarate
SuccinateSuccinyl
CoA
-Ketoglutarate
Isocitrate
Citricacidcycle
S CoA
CoA SH
NADH
NADH
FADH2
FAD
GTP GDP
NAD+
ADP
P i
NAD+
CO2
CO2
CoA SH
CoA SH
CoAS
H2O
+ H+
+ H+ H2O
C
CH3
O
O C COO–
CH2
COO–
COO–
CH2
HO C COO–
CH2
COO–
COO–
COO–
CH2
HC COO–
HO CHCOO–
CHCH2
COO–
HO
COO–
CH
HCCOO–
COO–
CH2
CH2
COO–
COO–
CH2
CH2
C O
COO–
CH2
CH2
C O
COO–
1
2
3
4
5
6
7
8
Glycolysis Oxidativephosphorylation
NAD+
+ H+
ATP
Citricacidcycle
Figure 9.12
A closer look at Citric Acid (Krebs) Cycle
Acetyl CoA adds its 2-carbon fragment to a 4-carbon molecule called oxaloacetateto form a 6-carbon molecule called citrate (ionized form of citric acid – gives the cycle its name!)
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
A closer look at Citric Acid (Krebs) Cycle
Acetyl CoA
NADH
Oxaloacetate
CitrateMalate
Fumarate
SuccinateSuccinyl
CoA
-Ketoglutarate
Isocitrate
Citricacidcycle
S CoA
CoA SH
NADH
NADH
FADH2
FAD
GTP GDP
NAD+
ADP
P i
NAD+
CO2
CO2
CoA SH
CoA SH
CoAS
H2O
+ H+
+ H+ H2O
C
CH3
O
O C COO–
CH2
COO–
COO–
CH2
HO C COO–
CH2
COO–
COO–
COO–
CH2
HC COO–
HO CHCOO–
CHCH2
COO–
HO
COO–
CH
HCCOO–
COO–
CH2
CH2
COO–
COO–
CH2
CH2
C O
COO–
CH2
CH2
C O
COO–
1
2
3
4
5
6
7
8
Glycolysis Oxidativephosphorylation
NAD+
+ H+
ATP
Citricacidcycle
Figure 9.12
Carbon dioxide is released
High energy molecules are generated:
3 NADH
1 FADH2
1 ATP
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• It’s called a cycle because at the end of the series of reactions, oxaloacetate is regenerated, allowing the series to repeat as a new acetyl-CoA enters
Another closer look at Citric Acid Cycle
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
A closer look at oxidative phosphorylation
• Oxidative phosphorylation
– is ATP synthesis powered by redox reactions that transfer electrons from food to oxygen.
– involves:
• transfer of electrons by NADH and FADH2
• to the electron transport chain
• coupled with chemiosmosis (by which ATP is actually generated)
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
A closer look at the Electron Transport Chain• The electron transport chain
• is a collection of molecules embedded in the inner mitochondrial membrane.
• How does this provide an example of STF?
Folding of the inner membrane to form cristae increases its surface area, providing space for thousands of copies of the chain in each mitochondrion.
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A closer look at the Electron Transport Chain
Consists of 5 major components:
• 3 complexes of integral membrane proteins
• 2 freely-diffusible molecules (that shuttle electrons from one complex to the next):
o ubiquinone (a lipid – only member of the ETC that is not a protein!)
o cytochrome c
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A closer look at the Electron Transport Chain
H2O
O2
NADH
FADH2
FMN
Fe•S Fe•S
Fe•S
O
FAD
Cyt b
Cyt c1
Cyt c
Cyt a
Cyt a3
2 H + + 12
I
II
III
IV
Multiproteincomplexes
0
10
20
30
40
50
Free
ene
rgy
(G) r
elat
ive
to O
2 (k
cl/m
ol)
Figure 9.13
Electrons from NADH and FADH2“fall’ through a series of energy releasing steps
3 integral protein complexes
2 “free floating”molecules
At the end of the chain electrons are passed to oxygen, forming water
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Electron Transport Chain – cytochromes
• Cytochromes are key “players” in the ETC!
– are membrane-bound hemoproteins(contain heme groups) that carry out electron transport
– held in membranes in an organized way so redox reactions occur in correct sequence for maximum efficiency!
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A closer look at the Electron Transport Chain
• At certain steps along the electron transport chain electron transfer causes protein complexes to pump H+ from the mitochondrial matrix to the intermembrane space
• The resulting H+ gradient
– Stores energy
– Drives chemiosmosis in ATP synthase
– Is referred to as a proton-motive force
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oxidative phosphorylation = ETC + chemiosmosis
Oxidativephosphorylation.electron transport
and chemiosmosis
Glycolysis
ATP ATP ATP
InnerMitochondrialmembrane
H+
H+H+
H+
H+
ATPP i
Protein complexof electron carners
Cyt c
I
II
III
IV
(Carrying electronsfrom, food)
NADH+
FADH2
NAD+
FAD+ 2 H+ + 1/2 O2
H2O
ADP +
Electron transport chain (ETC) Chemiosmosis
ATPsynthase
Q
Oxidative phosphorylation!
Intermembranespace
Innermitochondrial
membrane
Mitochondrialmatrix
Figure 9.15
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• Electron transfer in ETC provides energy needed to pump H+ into intermembrane space
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Chemiosmosis
– Chemiosmosis is an energy-coupling mechanism that uses the energy from an H+ gradient generated across a membrane to drive cellular work
– in this case, the cellular work is to make ATP
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Chemiosmosis: The Energy-Coupling Mechanism
• ATP synthase is the enzyme (& molecular motor!) that actually makes ATP
INTERMEMBRANE SPACE
H+
H+
H+
H+
H+
H+ H+
H+
P i+
ADP
ATP
rotor spins clockwise when H+ flows past it down H+ gradient.
stator is anchoredin membrane to hold knobstationary.
rod (“stalk”) extends into knob & also spins, activatingcatalytic sites in knob.
stationary knob contains 3 catalytic sites that join Pi to ADP to make ATP.
MITOCHONDRIAL MATRIX
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Chemiosmosis & ATP synthesis• return flow of H+ turns the molecular motor
(enzyme complex called ATP synthase) which provides energy to make ATP
Motor is driven by
proton motive force!
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Summary of energy flow during cellular respiration
• During respiration, most energy flows in this sequence:
Glucose NADH (FADH2) electron transport chain proton-motive force ATP
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Review of three main metabolic phases
Electron shuttlesspan membraneCYTOSOL 2 NADH
2 FADH2
2 NADH 6 NADH 2 FADH22 NADH
Glycolysis
Glucose2
Pyruvate
2AcetylCoA
Citricacidcycle
Oxidativephosphorylation:electron transport
andchemiosmosis
MITOCHONDRION
substrate-levelphosphorylation
substrate-levelphosphorylation
oxidative phosphorylation (# depends on which shuttle
transports electronsfrom NADH in cytosol)
Maximum per glucose:About
36 or 38 ATP
+ 2 ATP + 2 ATP + about 32 or 34 ATP
or
Figure 9.16
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Efficiency• About 40% of the energy in a glucose
molecule is transferred to ATP during cellular respiration, making approximately 38 ATP (a car has an efficiency of about 25%)
– Most of this ATP (30-34) is generated by oxidative phosphorylation
– Oxygen is required to harvest the maximum amount of energy from food
Oxygen is the final electron and hydrogen acceptor!
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Glycolysis review
• Glycolysis oxidizes glucose to 2 molecules of pyruvate.
• The oxidizing agent is NAD+, not oxygen!
• Therefore, glycolysis generates 2 ATPs whether oxygen is present (aerobic) or not (anaerobic)
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Fermentation• In the absence of oxygen cells can still
produce ATP through fermentation
• Fermentation is an extension of glycolysisthat can generate ATP solely by substrate-level phosphorylation – as long as there is enough NAD+!
• For this to work, there must be a way to recycle NAD+ from NADH
– NADH must be able to donate its electrons to something!
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Fermentation
• Remember that under aerobic conditions, NADH transfers electrons to the ETC
• Under anaerobic conditions, the only choice is to transfer electrons to pyruvate
• Therefore, fermentation consists of glycolysis plus reactions in which NADH transfers electrons to pyruvate or its derivatives
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Types of Fermentation• Types of fermentation differ in the waste
products formed from pyruvate:
– In alcohol fermentation, pyruvate is converted to ethanol in 2 steps, releasing CO2
• Occurs in yeast and many bacteria
– In lactic acid fermentation, pyruvate is reduced directly to form lactate, with no release of CO2
• Occurs in bacteria, certain fungi, and human muscle cells (sore muscles!)
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2 ADP + 2 P1 2 ATP
GlycolysisGlucose
2 NAD+ 2 NADH
2 Pyruvate
2 Acetaldehyde���2 Ethanol(a) Alcohol fermentation
2 ADP + 2 P1 2 ATP
GlycolysisGlucose
2 NAD+ 2 NADH
2 Lactate
(b) Lactic acid fermentation
HH OH
CH3
C
O –
OCC O
CH3
HC O
CH3
O–
C OC O
CH3OC OC OHH
CH3
CO22
Figure 9.17
Types of Fermentation
1 carbon dioxide is removed and released as CO2
NADH reduces acetaldehyde to ethanol
NADH reduces pyruvate to lactate
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Products of Fermentation• Alcohol fermentation:
– Beer, wine
• Lactic acid fermentation:
– Cheese, yogurt
• Combination alcohol & lactic acid fermentation:
– Soy sauce!
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Fermentation vs. Cellular Respiration
• Similarity:
– Both use glycolysis to oxidize glucose and other organic fuels to pyruvate
• Differences:
– final electron acceptor
• Cellular respiration oxygen
• Fermentation pyruvate (or derivative)
– ATP yield
• Cellular respiration about 38 ATP
• Fermentation 2 ATP
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Facultative anaerobes
• Facultative anaerobes are organisms that can use either fermentation or respiration, depending on conditions
– When fermenting, organisms must consume sugar at a faster rate (why?)
– Our muscle cells behave as facultative anaerobes (how?)
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Significance of pyruvate• Pyruvate is the fork in the metabolic road
Glucose
CYTOSOL
PyruvateNo O2 presentFermentation
O2 presentCellular respiration
Ethanolor
lactate
Acetyl CoAMITOCHONDRION
Citricacidcycle
Figure 9.18
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Evolutionary Significance of Glycolysis• Glycolysis
– Occurs in nearly all organisms
– Probably evolved in ancient prokaryotes before there was oxygen in the atmosphere
• Oldest fossils 3.5 billion ya
• Evidence of O2 2.7 billion ya
– Does not require membrane-enclosed organelles, which evolved later
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Catabolic pathways• We get calories (ATP) from various
carbohydrates, fats, and proteins (not just glucose!)
– Glycolysis can accept a wide range of carbohydrates for catabolism
– Amino acids can feed into glycolysis or Krebs cycle after the amino group is removed (deamination) urea/ammonia
– Glycerol (from fats) is converted to PGAL
– Beta oxidation breaks down fatty acids to form acetyl CoA
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Catabolic pathways• The catabolism of various molecules from food
Amino acids
Sugars Glycerol Fattyacids
GlycolysisGlucose
Glyceraldehyde-3- P
Pyruvate
Acetyl CoA
NH3
Citricacidcycle
Oxidativephosphorylation
FatsProteins Carbohydrates
Figure 9.19
Deamination produces nitrogenous wastes
After digestion, food molecules are converted to various intermediate compoundsthat can enter at various stages of respiration
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Anabolic Pathways (Biosynthesis)
• Cells need materials as well as energy! (Not all food molecules are oxidized as fuel to make ATP)
• Instead, intermediates of glycolysis and Krebs cycle can be diverted into anabolic pathways to synthesize needed molecules(this uses ATP!)– Humans can make about half of the 20 amino acids
from Krebs cycle intermediates
– Carbohydrates and proteins can be converted to fats through glycolysis & Krebs cycle intermediates (even fat-free diet can store fat!)
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Feedback Mechanisms in Cellular Respiration
• Cellular respiration is controlled by allosteric enzymes at key points in glycolysis and the citric acid cycle
– Principle of supply & demand
• When cell is working hard, ATP drops, respiration speeds up
• When there is plenty of ATP, respiration slows down
– Phosphofructokinase (allosteric enzyme) is inhibited by ATP and stimulated by AMP
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Control of cellular respiration
• Phosphofructokinase is an important “switch” (regulated by ATP)
Glucose
Glycolysis
Fructose-6-phosphate
Phosphofructokinase
Fructose-1,6-bisphosphateInhibits Inhibits
Pyruvate
ATPAcetyl CoA
Citricacidcycle
Citrate
Oxidativephosphorylation
Stimulates
AMP
+– –
Figure 9.20
Cell doesn’t waste valuable resources!
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
One final point . . .
• Remember, the energy that keeps us alive is released, not produced, by cellular respiration!
• That energy was stored in food by photosynthesis (next chapter . . . . ! )
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Any questions?