Lecture 9 – Cellular Respiration

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First…watch this video. Seriously.

http://www.khanacademy.org/video/introduction-
to-cellular-respiration?playlist=Biology

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 In this lecture
• Cellular respiration
• Redox reactions
• Glycolysis
– Pyruvate oxidation
• Krebs Cycle
• Electron Transport Chain

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 Why do we do respiration?
• Cellular respiration provides most of our ATP
• The components of our diet provides the
reactants for cellular respiration
– Glucose is what we’ll study today
– Lipids and protein breakdown will be briefly
covered, and studied more in depth in another
course

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Biochemical pathways are:

Exergonic and endergonic reactions

Oxidation and reduction reactions
Enzymatic reactions

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The Big Picture

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 From food to ATP
• Amylase in saliva starts to break down starches
to disaccharides
• Stomach acid breaks apart large structures such
as cells and intercellular structures
• Amylase in the small intestine completes the
breakdown of all carbohydrates to disaccharides
• Maltases, lactases, and sucrases break down
disaccharides into monosaccharides
• Glucose is brought to all the cells in the body
through the circulatory system
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Energy production sites in the cell
• Glucose is brought inside the cell by
cotransport with sodium
• The mitochondria are where ATP is produced

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 Electron energy levels
• An electron loses potential energy when it
shifts to a more electronegative atom
C6H12O6 + 6O2 6CO2 + 6H2O + energy

Here, electrons are transferred from carbon and hydrogen to

oxygen

The electrons in this reaction lose a LOT of potential energy

Carbohydrates and fats are high-energy foods because they

have a lot of electrons associated with hydrogen

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 Figure 9.5

H2  1/2 O2 2H  1/
2 O2
(from food via NADH)
Controlled
release of
2H  2e
+ 
energy for
synthesis of
ATP
ATP

Free energy, G
Free energy, G

Explosive ATP
release of
heat and light ATP
energy

2 e
1/ O2
2
2 H+

H2O H2O

(a) Uncontrolled reaction (b) Cellular respiration

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 The chemical reactions in respiration
The chemistry definition:

Oxidation and reduction

OIL RIG

Oxidation is “losing” Reduction is “gaining”

Oxygen is
highly
electro-
What is being lost and gained? Electrons!
negative
Electrons are usually lost to oxygen

Oxidation and reduction reactions often occur in a pair,

and together are called redox reactions

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The chemical reactions in respiration
The biology definition:

Oxidation and reduction

Losing a hydrogen atom Gaining a hydrogen atom

In biochemical reactions, hydrogen is what usually gets

swapped around

Hydrogen almost always bonds to an atom that is more

electronegative (C, O, N, P), and so loses its electron

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Redox Reactions

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 The chemical reactions in respiration
• Redox reactions
• Phosphorylation/dephosphorylation
– Carried out by kinases and phosphatases
– Phosphorylation increases chemical potential
energy and “primes” the molecule for work

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 New players in the enzyme game
NAD+ is derived
from niacin
• NAD+ and NADH NAD+ is a coenzyme
• FAD and FADH2 FAD is derived from riboflavin

NAD+ shuttles electrons through

the various stages of cellular
respiration

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 NAD+ and FAD
reduction Reduced
form
NAD+ NADH
Oxidized
oxidation
form

• NAD+ accepts electrons and becomes NADH

• NAD+ is reduced into NADH
• NADH is a reducing agent, and is recycled back to
NAD+ through oxidation
Each NADH (the reduced form of NAD+) represents
stored energy that is tapped to synthesize ATP.
Each NADH produces 3 ATPs in the e- transport chain
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 What is cellular respiration?
Glucose Energy - 686kcal/mol

C6H12O6 + 6O2 6CO2 + 6H2O + energy

Cellular respiration

38 ATP Heat
1 molecule of glucose produces 28 ATPs

Cellular respiration is the step-by-step

release and harness of the chemical
potential energy in glucose
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Redox Reactions in Cellular Respiration
During cellular respiration, the fuel (such as glucose) is
oxidized, and O2 is reduced

becomes oxidized

becomes reduced

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 Two types of cellular respiration:
The breakdown of organic
molecules is always exergonic

• Aerobic respiration – takes place in the

presence of oxygen
• Anaerobic respiration – takes place in the
absence of oxygen
– Fermentation is a type of anaerobic respiration
where sugars are partially degraded
– Consumes compounds other than oxygen

Cellular respiration includes both aerobic and anaerobic respiration but is

often used to refer to aerobic respiration
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 The stages of aerobic respiration
Consumes two ATP
Breaks glucose in Net 2 ATP
Glycolysis Generates 4 ATP
half

Rearranges the half-

glucose molecule Citric Acid Cycle* Generates 2 ATP

Electrons from Electron Transport Generates 34 ATP

glucose are passed Chain
around

*AKA the Krebs cycle and the TCA cycle

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The Big Picture

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 Figure 9.6-1

Glycolysis
Electrons
carried
via NADH

Glycolysis

Glucose Pyruvate

CYTOSOL MITOCHONDRION

ATP

Substrate-level
phosphorylation
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 Glycolysis
• The players:
The production of
– Glucose ATP from ADP by
– Pyruvate direct transfer of a
phosphate group
– ADP/ATP from a
phosphorylated
– Enzymes protein
• The processes:
– Substrate-level phosphorylation
• The locations
– Cytoplasm
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 Glycolysis
Beginning structure: End structures:

- Breaking down of glucose

- Can be done with or without oxygen “glyco” = glucose
“lysis” = breaking
apart
Broken down into two stages:
• The “investment” phase (uses 2 ATP)
• The “payoff” phase (produces 2 ATP)
“You have toNSCC
spend money to make money”
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Glycolysis

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Glycolysis

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Glycolysis

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Glycolysis

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Glycolysis

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Glycolysis

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Glycolysis

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Glycolysis

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Glycolysis

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Glycolysis

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 Glycolysis
• Begin:
– Glucose
– NAD+
– ADP
• End: Most of glycose’s original energy
is still present in pyruvate!
– 2 pyruvate
– 2 NADH
– 2 ATP

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 Glycolysis
• In the presence of O2, pyruvate enters the
mitochondrion where the oxidation of glucose
is completed during TCA cycle
• Without O2, pyruvate undergoes fermentation
into either ethanol or lactic acid

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Glycolysis

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 Pyruvate Oxidation
Glycolysis feeds into TCA cycle ONLY when oxygen is present!!

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 Pyruvate Oxidation
• Before pyruvate can be fed into TCA cycle, it
must become acetyl-CoA (acetyl-coenzyme A)
• It does this through pyruvate oxidation
– Produces one NADH from NAD+
– Three-carbon pyruvate is converted into two-
carbons + acetyl-CoA

Think of acetyl-
CoA as a
transporter for
the carbon
atoms from
pyruvate
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Pyruvate Oxidation

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 Pyruvate Oxidation

• Acetyl-CoA couples with oxaloacetate, the

first molecule in TCA cycle
• Acetyl-CoA + oxaloacetate = citrate

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 The Citric Acid Cycle
• The players:
– Acetyl-CoA
– Oxaloacetate
– Plus many more carbon skeleton intermediates
– Enzymes
• The processes:
– Hydrolysis
– Redox reactions
• The locations:
– Mitochondrial matrix

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 The Citric Acid Cycle

Begin: End:
Oxaloacetate Oxaloacetate
1 ADP 1 ATP
3 NAD+ 3 NADH
1 FAD 1 FADH2

The citric acid cycle, also called the Krebs cycle,

completes the break down of pyruvate to CO2
The citric acid cycle has eight steps, each catalyzed by a specific enzyme

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The Citric Acid Cycle

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 The Citric Acid Cycle
Step 1 and 2: Overview

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 The Citric Acid Cycle
Step 1 and 2: In detail

CoA is recycled
here to go back
to pyruvate
oxidation

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 The Citric Acid Cycle
Step 3 and 4: In detail and overview

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 The Citric Acid Cycle
Step 5 and 6: Overview

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 The Citric Acid Cycle
Step 5 and 6: In detail

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 The Citric Acid Cycle
Step 7 and 8: Overview

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 The Citric Acid Cycle

Step 7 and 8:
In detail

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 The Citric Acid Cycle
• The citric acid cycle is the entry point for other
catabolic pathways
• Acetyl-CoA can be derived from carbohydrates,
proteins, and fats

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 The Citric Acid Cycle
This is for one pyruvate. Remember, one glucose molecule produces two pyruvates!

• Begin: • End:
– Acetyl-CoA – 3 CO2s
– Oxaloacetate – Oxaloacetate
– 3NAD+ – 3 NADH
– 2 FAD – 2 FADH2
– 1ADP – 1 ATP
The whole point of
TCA cycle is to
produce NADH
and FADH2

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The Citric Acid Cycle

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 The electron transport chain
• The whole cellular process is about producing
ATP. Why do we care about NADH and
FADH2?
– These molecules then get oxidized in the electron
transport chain
– Every NADH will produce 3 ATP
– Every FADH2 will produce 2 ATP

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 The electron transport chain
Begin: End:
ADP ATP
10 NADH 10 NAD+
2 FADH2 2 FAD
Electrons are passed along at lower and lower energy levels to release their energy

The electron transport chain breaks the large free-energy drop from food to O2
into smaller steps that release energy in manageable amounts

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 The electron transport chain
• The players:
– FADH2, NADH, ADP
– ATP Synthase
– Cytochromes and membrane proteins
• The processes:
– Chemiosmosis
• The location:
– Intermembrane space of the mitochondria

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 The electron transport chain
• The electron transport chain is a series of
proteins that pass along electrons
– Electrons come from NADH and FADH2
– Proteins are embedded in the matrix membrane
– Each time an electron is passed, it releases energy
– That energy is used to drive protons across the
membrane into the intermembrane space
– This creates an electrochemical gradient

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 The electron transport chain
NADH  NAD+ + H+ + 2e-
CoQ, CytC, and
CytB are all
membrane proteins
CoQ on the inner matrix
Energy increases

Release of energy membrane

CytC
Release of energy
CytB

Release of energy
O2

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The electron transport chain
FADH2’s electrons are
lower energy than
NADH, and so enter
the electron transport
chain at a protein
further along in the
chain

The transport proteins alternate

reduced and oxidized states as they
accept and donate electrons

Oxygen accepts those now

very-low energy electrons.
Oxygen is the terminal
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 The electron transport chain
• The release of energy is used to pump H+
across the matrix membrane into the
intermembrane space

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 The electron transport chain
• H+ are pumped against their gradient using the
energy released from passing electrons to lower and
lower energy states
• This creates an electrochemical gradient
We can then
couple the
potential energy
in the
electrochemical
gradient to
another
biochemical
reaction
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 The electron transport chain
• A bigger picture:

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 The electron transport chain
What is that electrochemical gradient used for?
To create ATP!
How is that done?
Through a protein called ATP synthase
ATP synthase translates the potential energy in the electrochemical
gradient into the potential energy in the phosphate bonds of ATP

The flow of H+ with its electrochemical gradient is an exergonic reaction

ATP synthase couples an exergonic reaction with an endergonic reaction

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The electron transport chain
• ATP synthase is a turbine that
connects the flow of protons
to ADP  ATP
phosphorylation
• This is called chemiosmosis

Electrochemical energy  Kinetic energy  Chemical energy

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 The electron transport chain

As the turbine turns with

the “current” of protons
flowing past, it
phosphorylates ADP into
ATP
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 The electron transport chain
ATP Synthase: another view

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 The electron transport chain
All together:

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 The electron transport chain
• 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

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Energy flows in this direction:
glucose  NADH  electron transport chain  proton-motive force  ATP

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What happens without oxygen?

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 Fermentation
• Anaerobic respiration uses an electron
transport chain with a final electron acceptor
other than O2, for example sulfate
• Produces much less energy than aerobic
respiration
– Only source of ATP is substrate-level
phosphorylation

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 Fermentation
• Two common types of fermentation:
– Lactic acid fermentation
• Lactic acid fermentation by some fungi and bacteria is used to
make cheese and yogurt
• Human muscle cells use lactic acid fermentation to generate ATP
when O2 is scarce
– Alcohol fermentation
• Alcohol fermentation by yeast is used in brewing,
winemaking, and baking

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 Alcohol Fermentation

• Pyruvate is converted
to ethanol in two steps
– NADH produced in
glycolysis is oxidized
to NAD+
– Glucose is not
conpletely digested

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 Lactic Acid Fermentation

• Pyruvate is
converted to
lactate in one
step
– NADH produced
during glycolysis
is oxidized to
NAD+

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 Comparing Aerobic and Anaerobic
Respiration
Aerobic Respiration Anaerobic Respiration
Glycolysis Yes Yes
Krebs Cycle Yes No
Electron Transport Yes No
Chain
ATP Production 32 per glucose 2 per glucose
NADH production Yes Yes
FADH2 production Yes No
Terminal electron O2 Pyruvate or acetaldehyde
acceptor

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 Who uses what pathway?
• Obligate anaerobes carry out fermentation or
anaerobic respiration and cannot survive in the
presence of O2
• Yeast and many bacteria are facultative anaerobes,
meaning that they can survive using either
fermentation or cellular respiration
• We require oxygen to live, and are obligate aerobes
• In a facultative anaerobe, pyruvate is a fork in the
metabolic road that leads to two alternative catabolic
routes

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Figure 9.18
Glucose

Glycolysis
CYTOSOL

Pyruvate
No O2 present: O2 present:
Fermentation Aerobic cellular
respiration

MITOCHONDRION
Ethanol, Acetyl CoA
lactate, or
other products
Citric
acid
cycle
 Catabolism of other biomolecules
• Proteins must be digested to
amino acids; amino groups
can feed glycolysis or the
citric acid cycle

• Fats are digested to glycerol

(used in glycolysis) and fatty
acids (used in generating
acetyl CoA)

• Fatty acids are broken down

by beta oxidation and yield
acetyl CoA

• An oxidized gram of fat

produces more than twice as
much ATP as an oxidized
gram of carbohydrate

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Controlling Respiration
ATP and citrate inhibit
phosphofructokinase

AMP is a positive allosteric

regulator of phosphofructokinase

If you have a lot of ATP or citrate (that

means a lot of energy) glycolysis is shut
down

If you have a lot of AMP (very little

energy is present) glycolysis is
stimulated
If ATP concentration begins to drop,
respiration speeds up; when there is
plenty of ATP, respiration slows down

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 The Evolutionary Significance of
Glycolysis
• Ancient prokaryotes are thought to have used
glycolysis long before there was oxygen in the
atmosphere
• Very little O2 was available in the atmosphere until
about 2.7 billion years ago, so early prokaryotes likely
used only glycolysis to generate ATP
• Glycolysis is a very ancient process

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 Vocabulary
• Glycolysis
• Krebs/TCA cycle
• Redox reactions
• Terminal electron acceptor
• Chemiosmosis
• Oxidative phosphorylation
• Proton-motive force
• Fermentation

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