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RESPIRATION
8.1 ATP (Adenosine Triphosphate)
ADP + Pi → ATP
∆ G = + 31 kJ/mol (+7.3 kcal/mol)
Energy to drive endergonic regeneration
of ATP comes from exergonic process of
cellular respiration.
8.2 Aerobic Respiration
Stages of respiration:
i. Glycolysis
ii. Citric acid cycle
iii. Electron transport chain (ETC)
iv. Oxidative phosphorylation.
Glycolysis - in cytoplasm.
Glucose broken down into two
molecules of pyruvate.
Citric acid cycle - in mitochondrial
matrix.
Completes breakdown of glucose
by oxidizing a derivative of pyruvate
to CO2.
Several steps in glycolysis and citric
acid cycle are redox reactions -
dehydrogenase enzymes transfer
electrons from substrates to NAD+,
forming NADH.
NADH passes electrons to ETC.
Electrons then move from molecule to
molecule until they combine with
molecular O2 and H+ to form water.
As they passed along chain, energy
carried by electrons is used to
synthesize ATP in mitochondrion via
oxidative phosphorylation.
Inner membrane of mitochondrion is
site of electron transport chain (ETC)
and chemiosmosis
ETC + Chemiosmosis = Oxidative
Phosphorylation.
Produces almost 90% of ATP
generated by respiration.
Some ATP formed directly during
glycolysis and citric acid cycle by
substrate-level phosphorylation.
– Enzyme transfers phosphate group from
an organic substrate to ADP, forming
ATP.
Enzyme
Enzyme
ADP
P
Substrate + ATP
Product
For each molecule of glucose
degraded to CO2 & H2O, cell makes
up to 38 ATP
Each ATP ≈ 7.3 kcal/mol of free
energy.
8.2.1 Glycolysis
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+
Glucose split into two 3C sugars.
Sugars oxidized and rearranged to form
two molecules of pyruvate, ionized form
of pyruvic acid.
Two phases of glycolysis:
1. Energy investment phase
Cell invests ATP to provide activation
energy by phosphorylating glucose.
Requires 2 ATP per glucose.
2. Energy payoff phase
ATP produced by substrate-level
phosphorylation.
NAD+ reduced to NADH by electrons
released by oxidation of glucose.
Net yield from glycolysis = 2 ATP and 2
NADH per glucose.
No CO produced during glycolysis.
2
ATP
Hexokinase
ADP
Glucose-6-phosphate
Phosphoglucoisomerase
Fructose-6-phosphate
ATP
Phosphofructokinase
ADP
Fructose-
1, 6-bisphosphate
Aldolase
Isomerase
Dihydroxyacetone Glyceraldehyde-
phosphate 3-phosphate
2 NAD+
Triose phosphate
dehydrogenase
2 NADH
+ 2 H+
1, 3-Bisphosphoglycerate
2 ADP
Phosphoglycerokinase
2 ATP
3-Phosphoglycerate
Phosphoglyceromutase
2-Phosphoglycerate
Enolase
2 H2O
Phosphoenolpyruvate
2 ADP
Pyruvate kinase
2 ATP
Pyruvate
Overall reaction showing all reactants
and products resulting from
glycolysis:
NAD+ NADH + H+
Acetyl Co A
Pyruvate CO2 Coenzyme A
Transport protein
1) Carboxyl group removed as CO2.
2) Remaining 2C fragment oxidized to
acetate. An enzyme transfers 2
electrons to NAD+ to form NADH.
3) Acetate combines with coenzyme A,
forming the very reactive acetyl CoA.
Acetyl
CoA C C
CoA
C C CoA
Glycolysis Citric Oxidation
acid
cycle phosphorylation
Acetyl CoA
Acid Cycle
NADH
+ H+ H2 O
NAD+
Oxaloacetate
Malate Citrate
Isocitrate
CO2
Citric
acid
cycle NAD+
H2 O
NADH
Fumarate + H+
α -Ketoglutarate
FADH2
CO2
NAD+
FAD
Succinate NADH
Pi
GTP GDP Succinyl + H+
CoA
ADP
ATP
Acetyl group of acetyl CoA
combines with OAA, forming
citrate.
Citrate regenerated back to OAA.
3 CO2 molecules released,
including one released during
conversion of pyruvate to acetyl
CoA.
Cycle generates one ATP per turn by
substrate-level phosphorylation.
GTP is formed by substrate-level
phosphorylation.
GTP is used to synthesize an ATP
directly.
Most of chemical energy
transferred to NAD+ and FAD
during redox reactions.
Reduced coenzymes NADH and
FADH2 then transfer high-energy
electrons to ETC.
For every acetyl CoA, each cycle
produces:
i. 1 ATP by substrate-level
phosphorylation
ii. 3 NADH, and
iii. 1 FADH2
Pyruvate
(from glycolysis,
2 molecules per glucose) Citric
CO2 Glycolysis
acid
Oxidation
phosphorylation
NAD+ cycle
CoA
NADH
ATP ATP ATP
+ H+ Acetyl CoA
CoA
CoA
Citric
acid 2 CO2
cycle
FADH2 3 NAD+
FAD 3 NADH
+ 3 H+
ADP + P i
ATP
Summary:
Acetyl CoA + 3 NAD+ + FAD + ADP + Pi
+2H2O
FADH2
Multiprotein
I FAD complexes
40 FMN
Free energy (G) relative to O2 (kcal/mol)
Fe•S Fe•S II
Q
III
Cyt b Oxidative
Glycolysis Citric phosphorylation:
Fe•S acid electron transport
30 cycle
and chemiosmosis
Cyt c1 IV
Cyt c
Cyt a ATP ATP ATP
Cyt a3
20
10
0 2 H+ + 1/2 O2
H2O
During electron transport along ETC,
electron carriers alternate between reduced
and oxidized states as they accept and
donate electrons.
NADH
FAD
(reduced)
(oxidized)
CoQH2
Cyto. Oxidase
(reduced)
NAD (oxidized) H2O
FADH2
(oxidized)
(reduced)
CoQ Cyto. Oxidase H2
(oxidized) (reduced) ½ O2
Each component of chain becomes
reduced when it accepts electrons
from its “uphill” neighbor, which is
less electronegative.
It then returns to its oxidized form as
it passes electrons to its more
electronegative “downhill” neighbor.
Electrons carried by NADH are
transferred to the first molecule in
ETC, a flavoprotein.
Electrons continue along chain that
includes several cytochrome
proteins and one lipid carrier.
Prosthetic group of each
cytochrome is a heme group with
an iron atom that accepts and
donates electrons.
Last cytochrome of chain, cyt a3,
passes its electrons to oxygen, which
is very electronegative.
Each oxygen atom also picks up a
pair of H+ from aqueous solution to
form water.
For every two electron carriers (four
electrons), one O2 molecule is
reduced to two molecules of water.
Electrons carried by FADH2 have
lower free energy and are added
at a lower energy level than those
carried by NADH.
–ETC provides about one-third
less energy for ATP synthesis
when electron donor is FADH2
rather than NADH.
NADH
FADH2
ETC generates no ATP directly.
Its function is to break the large
free energy drop from food to
oxygen into a series of smaller
steps that release energy in
manageable amounts.
Chemiosmosis: Energy-Coupling
Mechanism
OXIDATIVE PHOSPHORYLATION: ETC &
CHEMIOSMOSIS : THE MECHANISM……
1. NADH delivers two electrons and two
protons to the first protein complex (I)
located in cristae.
2. Energy is released as the electrons
pass down the ETC (electrons move
from high energy to low energy level).
3. Energy is used to actively pump H+
across the membrane..
One of Its
A Mitochondrion Mitochondri
a a
A b
Cell
A
Crista
Outer
& Inner
Membrane
s c
ntermembran
e Matrix
4. Protons move from matrix to
intermembrane space
5. Protons are moved across by three (of
the four) complexes/complex I, III and IV
6. Proton gradient provide energy for ATP
synthesis.
7. Diffusion of protons from intermembrane
space to the matrix of mitochondrion
through ATP synthase (H+ only
permeable to ATP synthase)
8. As protons pass through it, energy is
obtained to phosphorylate ADP into ATP
ADP + Pi ATP.
9. After passing through three protein complexes,
electrons combine with one oxygen atom and two H+ to
form water (from matrix of mitochondrion)
2H+ + 1/2O2 + 2e H2 O
Glycolysis Oxidative
2 2 Citric phosphorylation:
Glucose Pyruvate Acetyl acid electron transport
CoA cycle and
chemiosmosis
About
Maximum per glucose: 36 or 38 ATP
Summary of
Glucose Metabolism
(2)
Products generated when a molecule of
glucose is oxidized to 6 CO2 molecules:
Conversions
NADH in cytoplasm produces 2 or 3 ATP
by oxidative phosphorylation depending
on shuttle system used to transport
electrons from cytosol into mitochondrion:
If electrons are passed to FAD, e.g.
brain cells - 2 ATP.
If electrons are passed to NAD+,
e.g., liver cells & heart cells = 3ATP
In mitochondria:
NADH - 3 ATP
FADH – 2ATP
2
FAD
NADH GP ETC
(glycolysis) FADH2
Malate-
NAD
NADH aspartate
ETC
(glycolysis) NADH
Glycolysis
Substrate-level phosphorylation = 2
ATP
2 NADH = 2 x 2 ATP = 4 ATP; or
= 2 x 3 ATP = 6 ATP
(Inner
Membrane) Electron Water
Transport 32
Oxygen System ATP
Efficiency of respiration
2 ADP + 2 P i 2 ATP
Glucose Glycolysis
2 Pyruvate
2 Ethanol 2 Acetaldehyde
2 ADP + 2 P i 2 ATP
Glucose Glycolysis
2 NAD+ 2 NADH
+ 2 H+
2 Pyruvate
2 Lactate
Lactic acid fermentation
Pyruvate reduced by NADH to lactate
without release of CO2.
Lactic acid fermentation by fungi and
bacteria - cheese and yogurt.
Human muscle cells switch from aerobic
respiration to lactic acid fermentation to
generate ATP when O2 is scarce.
Waste product, lactate, causes muscle
fatigue - converted back to pyruvate in
liver.
GLUCOSE (6C)
2NAD+
2NADH 2ETHANAL /
+ 2H+ ASETALDEHYDE (2C)
muscle cell/bacteria
2ATP CO2
2Pyruvates converted into
2lactate (3C) Yeast cell
2NADH are used 2Pyruvates converted into
2 ATP are produced 2ethanol (2C) &2CO2
2NADH are used
2PYRUVATES (3C) 2 ATP are produced
Fermentation and Cellular Respiration
Compared
Anaerobic : 2 ATP.
Facultative Anaerobes
CYTOSOL
Pyruvate
No O2 present O2 present
Fermentation Cellular respiration
MITOCHONDRION
Glycolysis
Glucose
Glyceraldehyde-3-P
NH3 Pyruvate
Acetyl CoA
Citric
acid
cycle
Oxidative
phosphorylation
Catabolism
Carbohydrates:
Polysaccharides
(starch/glycogen) hydrolyzed to
glucose monomers that enter
glycolysis.
Hexose sugars (galactose and
fructose) - modified to undergo
glycolysis.
Protein:
Proteins digested to individual amino
acids.
– Amino groups removed via
deamination.
– Nitrogenous waste excreted as
ammonia, urea, or another waste
product.
Carbon skeletons modified by enzymes
and enter as intermediaries into
glycolysis or citric acid cycle.
Fats:
Fats digested to glycerol and fatty
acids.
Glycerol converted to G3P –
enters glycolysis.
Fatty acids split into 2C fragments
via beta oxidation.
Enter citric acid cycle as acetyl
CoA.
Biosynthesis (Anabolic Pathways)