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Cellular Respiration

What is Cellular Respiration?


The process of converting food energy
into ATP energy

C6H12O6 + 6 O2 → 6 CO2 + 6 H2O + 36 ATP


Why are both Photosynthesis and Cell
Respiration important to Ecosystems?
Light is the ultimate
source of energy for all
ecosystems
Chemicals cycle and
Energy flows
Photosynthesis and
cellular respiration are
opposite reactions
Why do plants need both
chloroplasts and mitochondria?
Chloroplasts use
energy from the
sun to make
glucose
Mitochondria
convert glucose to
ATP—the energy
currency of the cell
Definitions: Oxidation-Reduction =
transfer of electrons from electron donor
to electron acceptor

Example: A:H + B = A + B:H


donor acceptor
 
electron donor (reducing agent,
reductant) is itself oxidized

electron acceptor (oxidizing


agent, oxidant) is itself reduced
-- both oxidation and reduction
must occur simultaneously
 --one compound donates
electrons
to (reduces) a second compound;
the
second accepts electrons from
(oxidizes) the first.
-- electrons always move from compounds
Redox/oxidation-reduction
withRedox
lower reduction potential to
compounds with higher reduction potential
( more positive).
Biological redox = Two half-reactions
A:H A
Reductant  Oxidant + e-
  B B:H
Oxidant + e-  Reductant
(acceptor) (donor)
Standard reduction potential, E°
-- measure of the tendency of oxidant to
gain electrons, to become reduced, a
potential energy.
Biological redox = Two half-reactions
A:H A
Reductant  Oxidant + e-
B B:H
 
Oxidant + e-  Reductant
(acceptor) (donor)
Standard reduction potential, E°
-- Reduction potential (also known as redox
potential, oxidation / reduction potential or ORP) is
a measure of the tendency of a chemical species to
acquire electrons and thereby be reduced.
*********************************************************
**
So, the more negative the reduction potential
is, the easier a reductant can reduce an oxidant
and

The more positive the reductive potential is, the


easier an oxidant can oxidize a reductant

The difference in reduction potential must be


important
*********************************************************
***
Reduction Potential Difference =Eº

 Eº = E°(acceptor) - E (donor)

 measured in volts.
 The more positive the reduction potential
difference is, the easier the redox reaction
 Work can be derived from the transfer of
electrons and the ETS
can be used to synthesize ATP.
 The reduction potential can be
related to free energy change by:

Gº = -nFEº

where n = # electrons transferred =


1,2,3
F = 96.5 kJ/volt, called the Faraday constant
******************************************************************************
***

Table of Standard Reduction Potentials


Oxidant + e-  reductant
-- e.g., Lehninger, 5th ed., p. 511
Note:
oxidants can oxidize every
compound with less positive
voltage -- (below it in the Table)
reductants can reduce every compound with
a less negative
voltage -- (above it in the Table)
******************************************************************************
***********
-- Electrons can move through a chain
of donors and acceptors

-- In the electron transport chain, electrons


flow down a gradient.

-- Electrons move from a carrier


with low reduction potential (high tendency
to donate electrons)
toward carriers with higher
reduction potential (high tendency
to accept electrons).
-- The overall voltage drop from NADH
E = -(-0.32 V)
to O Eº = +0.82 V
is Eº = 1.14 V
-- This corresponds to a large free
energy change of
 
G = - nFE = -220 kJ/mole (n =2)
 
-- Since ATP requires 30.5 kJ/mole
to form from ADP, more than
enough energy is available to
synthesize 3 ATPs from the
oxidation of NADH.
What is ATP?
Adenosine Triphosphate
– 5-Carbon sugar (Ribose)
– Nitrogenous base (Adenine)
– 3 Phosphate groups

Energy currency of the


cell

The chemical bonds that


link the phosphate groups
together are high energy
bonds

When a phosphate group


is removed to form ADP
and P, small packets of
energy are released
How is ATP used?
As ATP is broken down, it
gives off usable energy to
power chemical work and
gives off some nonusable
energy as heat.

Synthesizing molecules for


growth and reproduction
Transport work – active
transport, endocytosis, and
exocytosis
Mechanical work – muscle
contraction, cilia and flagella
movement, organelle
movement
Why use ATP energy and not
energy from glucose?
Breaking down glucose yields too much energy
for cellular reactions and most of the energy
would be wasted as heat.

1 Glucose = 686 kcal


1 ATP = 7.3 kcal
1 Glucose → 36 ATP

How efficient are cells at converting glucose into


ATP?
– 38% of the energy from glucose yields ATP,
therefore 62% wasted as heat.
Cellular Respiration is a Redox Reaction
(Oxidation)

C6H12O6 + 6 O2 → 6 CO2 + 6 H2O


(Reduction)

Oxidation is the loss of electrons or H+


Reduction is the gain of electrons or H+

Glucose is oxidized when electrons and H+ are passed


to coenzymes NAD+ and FAD before reducing or
passing them to oxygen.
Glucose is oxidized by a series of smaller steps so that
smaller packets of energy are released to make ATP,
rather than one large explosion of energy.
Cell Respiration can be divided into 4 Parts:
1) Glycolysis
2) Oxidation of Pyruvate / Transition Reaction
3) The Krebs Cycle
4) The Electron Transport Chain and
Chemiosmotic Phosphorylation
Where do the 4 parts of Cellular
Respiration take place?
Glycolysis:
– Cytosol
Oxidation of
Pyruvate:
– Matrix
The Krebs Cycled:
– Matrix
Electron Transport
Chain and
Cheimiosmotic
Phosphorylation:
– Cristae
Parts of the Mitochondria
Anaerobic Respiration (no oxygen required, cytoplasm)

1. Glycolysis Glucose  4 ATP (Net 2 ATP)


(substrate level) 2 ATP 2 NADH
2 Pyruvate

Aerobic Respiration (oxygen required, mitochondria)

2. Oxidation 2 Pyruvate  2 CO2


of 2 NADH
Pyruvate 2 Acetyl CoA

3. Krebs Cycle 2 Acetyl CoA  4 CO2


(substrate level)
2 ATP
6 NADH
2 FADH2
4. Electron
10 NADH  32 ATP
Transport
2 FADH2 6 H2O
Chain
(chemiosmotic) 6 O2

Total: 36 ATP produced


ATP is made in two ways:
1) Substrate Level
Phosphorylation (glycolysis
& Krebs cycle)
2) Chemiosmotic
Phosphorylation (electron
transport chain)

Substrate-Level
Phosphorylation:
Energy and phosphate are
transferred to ADP using an
enzyme, to form ATP.
Phosphate comes from one
of the intermediate
molecules produced from
the breakdown of glucose.
Glycolysis
Glucose  2 Pyruvate
2 ATP 4 ATP (Net 2 ATP)
2 NADH

Glucose (C6) is split to make


2 Pyruvates (C3)
– 1st: ATP energy used to phosphorylate
glucose (stored energy)
– 2nd: phosphorylated glucose broken
down into two C3 sugar phosphates
– 3rd: the sugar phosphates are oxidized
to yield electrons and H+ ions which are
donated to 2 NAD+ → 2 NADH (stored
electron and hydrogen for the Electron
Transport Chain)
– 4th: The energy from oxidation is used
to make 4 ATP molecules (net 2 ATP)

This is substrate level phosphorylation


because an enzyme transfers
phosphate to ADP making ATP

Glycolysis produces very little ATP


energy, most energy is still stored in
Pyruvate molecules.
Oxidation of Pyruvate /Transition Reaction
2 Pyruvate  2 CO2
2 NADH
2 Acetyl CoA
When Oxygen is present,
2 Pyruvates go to the
matrix where they are
converted into 2 Acetyl
CoA (C2).
Multienzyme complex:
– 1st: each Pyruvate releases
CO2 to form Acetate.
– 2nd: Acetate is oxidized and
gives electrons and H+ ions
to 2 NAD+ → 2 NADH.
– 3rd Acetate is combined with
Coenzyme A to produce 2
Acetyl CoA molecules.
2 NADH’s carry electrons
and hydrogens to the
Electron Transport Chain.
The Krebs Cycle / Citric Acid Cycle
2 Acetyl CoA  4 CO2
2 ATP
6 NADH
2 FADH2
8 Enzymatic Steps in Matrix of
Mitochondria: Break down and Oxidize
each Acetyl CoA (2-C’s) to release 2 CO2
and yield electrons and H+ ions to
3 NAD+ + 1 FAD → 3 NADH + FADH2.
This yields energy to produce ATP by
substrate level phosphorylation.
The first step of the Krebs cycle combines
Oxaloacetate (4 C’s) with Acetyl CoA to
form Citric Acid, then the remaining 7
steps ultimately recycle oxalacetate.
Two Turns of the Krebs Cycle are required
to break down both Acetyl Coenzyme A
molecules.
The Krebs cycle produces some chemical
energy in the form of ATP but most of
the chemical energy is in the form of
NADH and FADH2 which then go on to
the Electron Transport Chain.
The Electron Transport Chain
10 NADH  32 ATP
2 FADH2 H2O
Oxygen
NADH and FADH2 produced
earlier, go to the Electron
Transport Chain.
NADH and FADH2 release
electrons to carriers/proteins
embedded in the membrane
of the cristae. As the
electrons are transferred, H+
ions are pumped from the
matrix to the intermembrane
space up the concentration
gradient. Electrons are
passed along a series of 9
carriers until they are
ultimately donated to an
Oxygen molecule.
½ O2 + 2 electrons + 2 H+
(from NADH and FADH2) →
H2O.
http://vcell.ndsu.nodak.edu/animations/etc/movie.htm
Sequence of
Respiratory
Electron
Carriers

Inhibitors
in green
In-depth Summary of the Site
Components
Complex 1
Has NADH binding site
– NADH reductase activity
NADH - NAD+
– NADH ---> FMN--->FeS---> ubiquinone
– ubiquinone ---> ubiquinone H2
– 4 H+ pumped/NADH
Nicotinamide

NAD+/NADH
NADP+/NADPH


Never covalently bound- freely diffusible
Reaction at Enzyme
+
reduced substrate + NAD(P) 
+
oxidized substrate + NAD(P)H + H

substrate relieved of two H atoms


= hydride to NAD+ & H+ into solution
-
Hydride = H = H:
+
= H + 2e-
two electrons
transferred reduced
NADH Dehydrogenase transfers
electrons from NADH to CoQ,
an electron carrier

Uses two bound cofactors to


accomplish this.

flavin and iron-sulfur center


Complex II

succinate ---FAD—ubiquinone
– Contains coenzyme Q
– FADH2 binding site
FAD reductase activity
FADH2 -- FAD
For NADH, one of two entry points
into the electron transport chain:

-- So the oxidation of one NADH results in the


reduction of one CoQ
-- Another important function
of the enzyme will be mentioned later.
Succinate Dehydrogenase

-- similar reaction can be written


yielding CoQH2
-- second entry into electron transport
-- substrate is succinate
-- FAD is reduced, not FMN
Flavin-linked
Dehydrogenases

Flavin mononucleotide Isoalloxazine


= FMN ring
Flavin adenine
dinucleotide = FAD ribitol

Riboflavin = ring
+ ribitol
Reduced form = FADH2
Transfer 2 H atoms with 2e-
or 2H

Reaction
AH2 + E-FMN = A +E-FMNH2
Coenzyme Q = Ubiquinone

a lipid in inner membrane


 carries electrons
 polyisoprene tail
 moves freely within membrane

2H++2e

Coenzyme Q
Coenzyme Q
Complex III
ubiquinone - ubiquinone ox
while cyt C gets reduced
Also contains cytochromes b
– proton pump 4H+
Adds to gradient
– 8 H+ / NADH
– 4 H+ / FADH2
Cytochromes - proteins in ETS
Carry electrons
Contain heme
or heme-like group
Heme is based on
porphyrins with iron
in center, usually
as Fe(II), and is tightly
bound at sides,
sometimes covalently
Contrast heme in cytochromes &
hemoglobin
--For cytochrome heme,

  a) all 6 sites of Fe are filled


(4 from porphyrin, 2 above and
below from protein) = no molecule
can approach
  b) carries electrons only:
Fe(III) + e-  Fe(II)
 
Note: Only one electron is transferred
at a time.
 COMPLEX III = b, an Fe-S and c1.
 Cytochrome c is mobile.
 COMPLEX IV = a+a3 =
cytochrome a-a3 =
cytochrome c oxidase -- large protein.
-- both a and a3 contain heme A and Cu
-- a3 Cu binds to oxygen and donates
electrons to oxygen
cytochrome a3 - only component of
ETS that can interact with O2
Complex IV
reduction of oxygen
cytochrome oxidase
cyt a+a3 red ---> oxidized state
oxygen ---> water
– 2 H+ + 2 e- + ½ O2 -- 2 H2O
– transfers e- one at a time to oxygen
Pumps 2H+ out
– Total of 10 H+ / NADH
– Total of 6 H+ / FADH2
What about NADH from glycolysis?
NADH made in cytosol
Can’t get into matrix of
mitochondrion
2 mechanisms
– In muscle and brain
Glycerol phosphate shuttle
– In liver and heart
Malate / aspartate shuttle
Glycerol Phosphate shuttle

http://courses.cm.utexas.edu/jrobertus/ch339k/overheads-3/ch19_glycerol-shuttle.jpg
Glycerol phosphate shuttle
In muscle and brain
Each NADH converted to FADH2
inside mitochondrion
– FADH2 enters later in the electron
transport chain
– Produces 1.5 ATP
Total ATP per glucose in muscle
and brain
Gycerol phosphate shuttle
– 2 NADH per glucose - 2 FADH2
– 2 FADH2 X 1.5 ATP / FADH2……….3.0 ATP
– 2 ATP in glycoysis ……………………2.0
ATP
– From pyruvate and Krebs
12.5 ATP X 2 per glucose ……………..25.0 ATP

Total = 30.0 ATP/ glucose


Malate – Aspartate Shuttle
in cytosol
In liver and heart
NADH oxidized while reducing
oxaloacetate to malate
– Malated dehydrogenase
Malate crosses membrane
Total ATP per glucose in liver and
heart
Malate – Aspartate Shuttle
– 2 NADH per glucose - 2 NADH
– 2 NADH X 2.5 ATP / NADH…………5.0 ATP
– 2 ATP from glycolysis………………..2.0 ATP
– From pyruvate and Krebs
12.5 ATP X 2 per glucose ……………..25.0 ATP

Total = 32.0 ATP/ glucose


Summary
Total ATP / glucose
– Muscle and brain 30.0 ATP
Uses glycerol phosphate shuttle

– Heart and liver 32.0 ATP


Uses malate aspartate shuttle
Chemiosmotic Theory --Peter Mitchell
 
-- A proton gradient is
generated with energy
from electron transport
by the vectorial transport
of protons (proton
pumping)
pumping by Complexes I,
III, IV from the matrix to
intermembrane space
of the mitochondrion.
-- The protons have a thermodynamic
tendency to return to the matrix =
Proton-motive force
The proton move back into the matrix
through the
FoF1ATP
synthase
driving
ATP synthesis.
The proton pumps are Complexes I,
III and IV.

Protons return thru ATP synthase


Chemiosmotic Phosphorylation

Hydrogen ions travel down their concentration gradient through a channel


protein coupled with an enzyme called ATP Synthase.
As H+ ions move into the matrix, energy is released and used to combine
ADP + P → ATP.
Hydrogens are recycled and pumped back across the cristae using the
Electron Transport Chain.
ATP diffuses out of the mitochondria through channel proteins to be used
by the cell.
http://vcell.ndsu.nodak.edu/animations/atpgradient/movie.htm
ATP Synthase
Multisubunit complex
with 4 parts:
– Rotor – spins as H+ ions flow
– Stator – holds the rotor and
knob complex together in the
cristae
– Internal Rod – extends
between rotor and knob, spins
when rotor spins which then
turns the knob
– Knob – contains 3 catalytic
sites that when turned change
shape and activate the enzyme
used to make ATP
The return of protons “downhill”
through Fo rotates Fo
relative to F1,
driving ATP
synthesis.
Note: Subunit 
rotates
through F1.
All Types of Molecules can be used
to form ATP by Cell Respiration:
Proteins, Carbohydrates,
and Lipids must first be
broken down into their
monomers and absorbed
in the small intestine.

Monomers may be
further broken down into
intermediate molecules
before entering different
parts of Cell respiration
to ultimately form ATP.
Review ATP Production:
1) Glycolysis → 2 ATP
2) Oxidation of Pyruvate → No ATP
3) The Krebs Cycle → 2 ATP
4) The Electron Transport Chain and
Chemiosmotic Phosphorylation:
– Each NADH produces 2-3 ATP so 10
NADH → 28 ATP
– Each FADH2 produces 2 ATP so 2
FADH2 → 4 ATP
Total = 36 ATP

1 Glucose = 686 kcal


1 ATP = 7.3 kcal
1 Glucose → 36 ATP
How efficient are cells at converting
glucose into ATP?
– 38% of the energy from glucose
yields ATP, therefore 62% wasted as
heat (used to maintain body
temperature or is dissipated)
– Ex. Most efficient Cars: only 25% of
the energy from gasoline is used to
move the car, 75% heat.
How is amount ATP synthesized
measured?

Quantify P/O ratio


 
Definition: # Pi taken up in
phosphorylating ADP per atom
oxygen (½O2), in other words
per 2e-.
NADH 3
FADH2 2
Anaerobic Respiration: Fermentation
If there is NO oxygen, then cells can make ATP by Fermentation
Without oxygen, Oxidation of Pyruvate and the Electron
Transport Chain do not operate.

Glucose → Pyruvate → Lactate


NAD+ Glycolysis 2 NADH Reduction Rxn or
2 ATP Alcohol + CO2

Fermentation yields a net gain of 2 ATP by substrate level phosphorylation


for every 1 Glucose. (Inefficient)

Two Forms of Fermentation:


Lactic Acid Fermentation (animals)
Alcohol Fermentation (yeast)
Inhibitors and Uncouplers: The
Difference is Important
Table 19-4 Lehninger POB 3rd Ed.

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