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ATP as Chief Energy Currency

ATP lies approximately in the middle of the energy spectrum for all
energy molecules.
This position is ideal because it can accept energy from high-energy
molecules and transfer energy to low-energy molecules.
Almost all of the high-energy molecules that have high energy of
hydrolysis, e.g., PEP, are formed during the oxidation of
carbohydrates, and almost all of the lower-energy molecules are
involved in metabolic processes that require energy to drive reactions.
ATP is in the middle as the carrier of energy from the high-energy
molecules to the low-energy molecules,
ATP distributes energy by passing it onto other nucleotides.
All reactions involve making and breaking of bonds so that
thermodynamic considerations as to whether a reaction proceeds or
not depends on the energy balance between making and breaking
bonds.

ATP as Chief Energy Currency

Physiological Importance of ATP


ATP is generated in the cell by energy-releasing processes and is broken
down by energy-consuming processes.
In this way, ATP transfers energy between spatially-separate metabolic
reactions.
ATP is the main energy source for the majority of cellular functions,
including synthesis of macromolecules such as DNA, RNA and proteins.
ATP also plays a critical role in the transport of macromolecules across
cell membranes, e.g., exocytosis and endocytosis.
ATP is critically involved in maintaining cell structure by facilitating
assembly and disassembly of elements of the cytoskeleton.

Stage Four: The Electron Transport Chain


As the electrons are transferred, some

electron energy is lost with each transfer.

This energy is used to pump protons (H +)

across the membrane from the matrix to the


inter membrane space.

A proton gradient is established.

Electron Transport Chain


ETC is a series of membrane-bound electron

carriers
Embedded in the inner mitochondrial
membrane
Electrons from NADH and FADH2 are
transferred to complexes of the ETC
Each complex

A proton pump creating proton gradient


Transfers electrons to next carrier

Electron Transport Chain and Chemiosmosis

Electron Transport Chain

Chemiosmosis

Chemiosmosis
Accumulation of protons in the

intermembrane space drives protons into the


matrix via diffusion
Membrane relatively impermeable to ions
Most protons can only reenter matrix through
ATP synthase
Uses energy of gradient to make ATP from ADP

+ Pi

Electron Transport Chain


The higher negative charge in the matrix

attracts the protons (H+) back from the


intermembrane space to the matrix.
The accumulation of protons in the

intermembrane space drives protons into the


matrix via diffusion.

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Energy Yield of
Theoretical energy yields
Respiration
- 2 ATPS for every FADH and 3 ATPs for every NADH

molecule are generated


- 38 ATP per glucose for bacteria
- 36 ATP per glucose for eukaryotes- NADH made in the
cytoplasm has to be transported to the mitochondria
( 1 NADH uses 1 ATP)
Actual energy yield
- 30 ATP per glucose for eukaryotes Why?
- 1.5 ATPs per molecule of FADH and 2.5 ATPs per

molecule of NADH

- reduced yield is due to leaky inner membrane and


use of the proton gradient for purposes other than
ATP synthesis

Theoretical ATP Yield

Regulating Aerobic Respiration


Control of glucose catabolism occurs at two key

points in the catabolic pathway.


glycolysis phosphofructokinase (ADP-stimulated;
ATP- and citrate-inhibited)
Krebs cycle - citrate synthetase (high ATPinhibited and low ATP-stimulated)
Other control mechanisms
low levels of citrate stimulate

phosphofructokinase
high levels of NADH inhibit pyruvate
dehydrogenase

Control of Glucose Catabolism

Oxidation Without O2
1. Anaerobic respiration
Use of inorganic molecules (other than O2) as
final electron acceptor
Many prokaryotes use sulfur, nitrate, carbon
dioxide or even inorganic metals
2. Fermentation
Use of organic molecules as final electron
acceptor

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Anaerobic respiration
Methanogens
CO2 is reduced to CH4 (methane)
Found in diverse organisms including cows

Sulfur bacteria
Inorganic sulphate (SO4) is reduced to hydrogen
sulfide (H2S)
Early sulfate reducers set the stage for
evolution of photosynthesis

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Fermentation
Reduces organic molecules in order to

regenerate NAD+
1.Ethanol fermentation occurs in yeast
CO2, ethanol, and NAD+ are produced

2.Lactic acid fermentation


Occurs in animal cells (especially muscles)
Electrons are transferred from NADH to
pyruvate to produce lactic acid

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ETHANOL FERMENTATION AND LACTIC ACID FORMATION


Alcohol fermentation in yeast
Glucose
2 ADP
2 ATP
O
C O
C O
CH3

G
L
Y
C
O
L
Y
S
I
S

H
H C OH
CH3
2 Ethanol

2 NAD+
2 NADH
CO2

H
C O
CH3
2 Acetaldehyde

2 Pyruvate
Lactic acid fermentation in muscle cells
Glucose
O
C O
G
2 ADP
L
H C OH
Y
CH3
C
2 NAD+ 2 Lactate
2 ATP
O
O
C O
C O
CH3

L
Y
S
I
S

2 NADH

Catabolism of Protein
Amino acids undergo deamination to remove

the amino group


Remainder of the amino acid is converted to a
molecule that enters glycolysis or the Krebs
cycle
Alanine is converted to pyruvate
Aspartate is converted to oxaloacetate

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Deamination
O

HO
C

Urea

H2N C H

HO

NH2

C
O

HC H

HC H

HC H

HC H

HO
Glutamate

HO

-Ketoglutarate

Catabolism of Fat
Fats are broken down to fatty acids and

glycerol
Fatty acids are converted to acetyl groups by -

oxidation
Oxygen-dependent process

The respiration of a 6-carbon fatty acid yields

20% more energy than 6-carbon glucose.

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-oxidation
H H O
Fatty acid
C C C OH
H H
CoA
ATP
AMP + PPi
O
Fatty acid H H
C C C
H H

CoA

FAD
FADH2
Fatty acid

H HO
C C C

CoA

CoA

H2O

HO H O
Fatty acid C C C CoA
HH
NAD+

Energy yield: 36
ATPs for one 6-C
fatty acid

NADH
OHO
Fatty acid C C C
H

Each round of oxidation:


consumes 1 ATP
and produces 1
FADH2 and NADH

CoA

Acetyl-CoA
Krebs
cycle

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Evolution of Cellular Respiration


1. degradation breakdown of organic molecules abiotically

produced

2. ability to store chemical energy in ATP


3. glycolysis initial breakdown of glucose
4. anaerobic photosynthesis use of light to pump protons out of

cells, the resulting proton gradient utilized to power ATP


production through chemiosmosis; H 2S is source of H+

5. oxygen-forming photosynthesis H 2O substituted for H2S


6. nitrogen fixation obtaining nitrogen atoms from N 2 gas which

requires breaking an NN triple bond

7. aerobic respiration energy harvested by stripping energetic

electrons from organic molecules

Evolution of Metabolism
Hypothetical timeline
1. Ability to store chemical energy in ATP
2. Evolution of glycolysis

Pathway found in all living organisms

Anaerobic photosynthesis (using H2S)


4. Use of H2O in photosynthesis (not H2S)
3.

Begins permanent change in Earths atmosphere

Evolution of nitrogen fixation


6. Aerobic respiration evolved most recently
5.

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