Biochemical Energy Production

BIOCHEMICAL
ENERGY
PRODUCTION
Stoker Chapter 23
Lippincott Chapter 6
Copyright ©2016 Cengage Learning. All Rights Reserved. 1

Sum total of all chemical reactions in a
living organism
Source of energy for the functioning of the

Metabolism human body
Also needed for many of the cellular

processes such as protein synthesis, DNA
replication, RNA transcription, and
membrane transport

Catabolism: All metabolic reactions
in which large biochemical molecules
Subtypes are broken down to smaller ones
• Usually energy is released in these reactions
of • Example: Oxidation of glucose
Metabolic Anabolism: All metabolic reactions in

which small biochemical molecules
Reactions are joined to form larger ones
• Usually require energy
• Example: Synthesis of proteins

FIGURE 23.1 -
THE
PROCESSES
OF
CATABOLISM
AND
ANABOLISM

• Series of consecutive biochemical
reactions used to convert a starting
material into an end product
• There are two types of metabolic
pathways:
• Linear
Metabolic • Cyclic
• Major pathways for all forms of life
Pathway are similar

• Knowledge of the cell structure is
essential for understanding metabolism
• Prokaryotic cell:
• No nucleus and found only in bacteria
• Presence of a single circular DNA molecule
near the center of the cell called nucleoid
• Eukaryotic cell: Cell where the DNA is
found in a membrane-enclosed nucleus
• About 1000 times larger than bacterial cells

FIGURE 23.3 - SCHEMATIC
REPRESENTATION OF A EUKARYOTIC
CELL

Plasma membrane - Cellular boundary
Cytoplasm: Water-based material of a eukaryotic

Eukaryotic cell
Cell Mitochondrion: Generates most of the energy

needed for a cell
Organelles Lysosome: Contains hydrolytic enzymes needed
and Their for cell rebuilding, repair, and degradation
Function Ribosome - Site for protein synthesis
Nucleus - Site where DNA is found

Mitochondrion
• An organelle that is
responsible for the generation
of most of the energy for a cell
• Outer membrane - Permeable
to small molecules
• 50% lipid and 50% protein
• Inner membrane - Highly
impermeable to most
substances
• 20% lipid and 80% protein
• Folded to increase surface area
• Synthesis of ATP occurs here

Adenosine Phosphates (ATP,
ADP, and AMP)
• Adenosine phosphates of
interest:
• Adenosine monophosphate (AMP)
- One phosphate group
• Structural component of RNA
• Adenosine diphosphate (ADP) -
Two phosphate groups
• Key component of metabolic pathways
• Adenosine triphosphate (ATP) -
Three phosphate groups
• Key component of metabolic pathways
• A phosphoryl group is derived from a
phosphate ion when it becomes part of
another molecule
ATP + H 2 O ⎯⎯
→ ADP + Pi + H + + energy
ADP + H 2O ⎯⎯
→ AMP + Pi + H + + energy
ATP + 2H 2 O ⎯⎯
→ AMP + 2Pi + 2H + + energy
•The net energy produced in these reactions is

used for cellular reactions

•In cellular reactions, ATP functions as
both a source of a phosphate group and a
source of energy
• Example: Conversion of glucose to glucose-6-phosphate

Uridine triphosphate (UTP) -
Involved in carbohydrate
metabolism
Role of Other
Nucleotide Guanosine triphosphate (GTP)
- Involved in protein and
Triphosphates carbohydrate metabolism
in Metabolism
Cytidine triphosphate (CTP) -

Involved in lipid metabolism

Flavin Adenine Dinucleotide
(FAD)
• Coenzyme required in
numerous metabolic redox
reactions
• Flavin subunit is the active
form which gains H atoms
when FAD is converted to
FADH2
• Ribitol is a reduced form
of the sugar ribose
FIGURE 23.6 - STRUCTURAL
FORMULAS OF FAD AND NAD+

Flavin Adenine Dinucleotide
(FAD)
FAD is the oxidized form FADH2 is the reduced form
Typical cellular reaction in

In enzyme reactions, FAD goes
which FAD serves as oxidizing
back and forth from oxidized
agent involves conversion of
to reduced form
an alkane to an alkene

Nicotinamide Adenine
Dinucleotide (NAD)
• Has coenzyme functions in metabolic redox

pathways
• NAD+ is the oxidized form of NAD
• NADH is reduced form
• Typical cellular reaction in which NAD+ serves
as the oxidizing agent is the oxidation of a
secondary alcohol to give a ketone
Subunit Structures of NAD
• 3-subunit structure:

Coenzyme A
• Derivative of vitamin B pantothenic acid

• Active form of coenzyme A is the sulfhydryl
group (–SH group) in the ethanethiol subunit of
the coenzyme
• Acetyl-CoA - Formed when acetyl group bonds
to CoA–SH via a thioester bond

Coenzyme A Subunit Structures

CLASSIFICATION OF METABOLIC
INTERMEDIATE COMPOUNDS
Metabolic intermediate compounds can be classified into three groups
based on their functions
Carboxylate Ions (Metabolic Acids)
• Polyfunctional acids formed as intermediates of

metabolic reactions
• There are 5 such acids that serve as substrates
for enzymes in metabolic reactions:
− 3 succinic acid derivatives (C4 diacid)
− Fumarate, oxaloacetate, and malate
− 2 glutaric acid derivatives (C5 diacid)
− a-ketoglutarate and citrate
High-Energy Phosphate
Compounds
• Several phosphate-containing
compounds found in metabolic
pathways are known as high-energy
compounds
• High-energy compounds: Have
greater free energy of hydrolysis
than a typical compound
• They contain at least one reactive
bond called strained bond
• Energy to break these bonds is less
than a normal bond
• More negative the free energy of
hydrolysis, greater the bond strain

• Typically the free-energy release is
greater than 6.0 kcal/mole (indicative of
bond strain)
• Strained bonds are represented by the
sign ~ (squiggle)

TABLE 23.1 - FREE ENERGIES OF
HYDROLYSIS OF COMMON PHOSPHATE-
CONTAINING METABOLIC COMPOUNDS
TABLE 23.1 - FREE ENERGIES OF
HYDROLYSIS OF COMMON PHOSPHATE-
CONTAINING METABOLIC COMPOUNDS
An Overview of Biochemical
Energy Production
• Energy needed to run the human body is obtained from food via a
multistep process involving several different catabolic pathways
• There are four general stages in the biochemical energy production
process:
• Stage 1: Digestion
• Stage 2: Acetyl group formation
• Stage 3: Citric acid cycle
• Stage 4: Electron transport chain and oxidative phosphorylation
• Each stage also involves numerous reactions

Stage 1: Digestion
• Begins in mouth (saliva contains starch-digesting enzymes),

continues in the stomach (gastric juices), and is completed in small
intestine
• Results in small molecules that can cross intestinal membrane into
the blood stream
• End products which are absorbed and transported to blood cells:
• Glucose and monosaccharides from carbohydrates
• Amino acids from proteins
• Fatty acids and glycerol from fats and oils

Stage 2: Acetyl Group
Formation
• The small molecules from Stage 1 are further

oxidized
• End product of these oxidations is acetyl CoA
and reduced coenzyme NADH
• This stage involves numerous reactions which
occur both in the cytosol as well as the
mitochondria of the cells
Stage 3: Citric Acid Cycle
• Takes place inside the mitochondria

• Acetyl group is oxidized to produce CO2 and energy
• Some energy produced in this stage is lost in the form
of heat
• Most energy is trapped in reduced coenzymes
NADH and FADH2
• The carbon dioxide we exhale comes primarily from
this stage

Stage 4: Electron Transport Chain
and Oxidative Phosphorylation
•Takes place in mitochondria

•NADH and FADH2 are oxidized to release
H ions and electrons
• Needed for the production of ATP, primary
energy carrier in metabolic pathways
•O2 inhaled is converted into H2O in this
stage
Citric Acid Cycle - An Introduction
• Is the series of biochemical reactions in which

the acetyl portion of acetyl CoA is oxidized to
carbon dioxide and the reduced coenzymes
FADH2 and NADH are produced
• Also know as:
• Tricarboxylic acid cycle (TCA) - Presence of three
carboxylate groups in citric acid
• Krebs cycle - Named after Hans Krebs who
elucidated this pathway
Citric Acid Cycle - An Introduction
• Important reactions in the citric acid
cycle include:
• Reduction of NAD+ and FAD to produce NADH and FADH2
• Decarboxylation of citric acid to produce carbon dioxide
• Summary of citric acid cycle reactions:

FIGURE
23.11 - THE
CITRIC
ACID
CYCLE

Step 1: Formation of Citrate
(1) the condensation of (2) hydrolysis of the

acetyl CoA and thioester bond in citryl CoA
oxaloacetate to form citryl to produce CoA-SH and
CoA, a process catalyzed by citrate, also catalyzed by
the enzyme citrate the enzyme citrate
synthase synthase

Step 2: Formation of Isocitrate
• Citrateis converted to its less
symmetrical isomer isocitrate
in an isomerization process that
involves a dehydration
followed by a hydration, both
catalyzed by the enzyme
aconitase

Step 3: Oxidation of Isocitrate
and Formation of CO2
•oxidation–reduction and
decarboxylation.
•isocitrate dehydrogenase

Step 4: Oxidation of a-Ketoglutarate
and Formation of CO2.
•a-ketoglutarate
dehydrogenase complex

Step 5: Thioester Bond Cleavage in Succinyl
CoA and Phosphorylation of GDP.
•succinyl-CoA synthetase

Steps 6 through 8 of the citric acid cycle involve a sequence of
functional group changes that have been encountered several
times in the organic sections of the text. The reaction sequence
is
•Step 6: Oxidation of Succinate.

•succinatedehydrogenase
•Step 7: Hydration of Fumarate.
• Fumarase
•Step 8: Oxidation
of L-Malate to
Regenerate Oxaloacetate
• malate dehydrogenase.
FIGURE
23.11 -
THE
CITRIC
ACID
CYCLE

• The rate at which the citric acid
cycle operates is controlled by the
body’s need for ATP
• When ATP supply is high, ATP
inhibits citrate synthase (Step 1
Regulation of of the cycle)
the Citric Acid • When ATP levels are low, ADP
Cycle activates citrate synthase
• Similarly, ADP and NADH control
isocitrate dehydrogenase
• NADH acts as an inhibitor
• ADP acts as an activator

The Electron Transport Chain - An
Introduction
Series of biochemical reactions in which

intermediate carriers aid the transfer of Ultimately react with molecular oxygen to
electrons and hydrogen ions from NADH give H2O
and FADH2
Lost energy is used to synthesize ATP in oxidative phosphorylation

Enzyme and Electron Carriers for
ETC
Located along inner mitochondrial membrane
Organized into four distinct protein complexes and two mobile

carriers
• Protein complexes tightly bound to membrane:
• Complex I: NADH–coenzyme Q reductase
• Complex II: Succinate–coenzyme Q reductase
• Complex III: Coenzyme Q–cytochrome c reductase
• Complex IV: Cytochrome c oxidase
• Two mobile electron carriers:
• Coenzyme Q and cytochrome c

• NADH from citric acid cycle is the source
of electrons processed via this complex
Complex I: • It contains > 40 subunits including flavin
NADH– mononucleotide (FMN) and several iron–
sulfur protein clusters (FeSP)
Coenzyme Q • Net result is the transfer of electrons
Reductase from NADH to coenzyme Q (CoQ)
• Several intermediate carriers are
involved in this electron transfer

Smaller than complex I
Complex II: Contains only four subunits including

Succinate– two FeSPs
Coenzyme Succinate is converted to fumarate

via this complex
Q
Reductase This complex processes FADH2
• CoQ is the final recipient of the electrons from

FADH2

• Contains 11 different subunits
• Several FeSP proteins and
cytochromes are electron carriers
in this complex
• Cytochrome: Heme iron protein
Complex III: in which reversible oxidation of
Coenzyme Q– an iron atom occurs
Cytochrome c • Various cytochromes, cyt a, cyt b,
Reductase cyt c, and so on, differ from each
other in:
• Their protein constituents
• The manner in which the heme
is bonded to the protein
• Attachments to the heme ring

Complex IV: Cytochrome c Oxidase
• Contains 13 subunits including two cytochromes
• The electrons flow from cyt c to cyt a to cyt a3
• In the final stage of electron transfer, the electrons
from cyt a3 and hydrogen ions combine with oxygen
(O2) to form water
O2 + 4H + 4e ⎯⎯
+
→ 2H2O -
• It is estimated that 95% of the oxygen used by cells

serves as the final electron acceptor for the ETC

SUMMARY OF THE FLOW OF ELECTRONS
THROUGH THE FOUR COMPLEXES OF
THE ELECTRON TRANSPORT CHAIN

Oxidative Phosphorylation - An
Introduction
• Oxidative phosphorylation: Process by which
ATP is synthesized from ADP using the energy
released in the electron transport chain
• Can be coupled reactions
• Coupled reactions: Pairs of biochemical

reactions that occur concurrently in which
energy released by one reaction is used in the
other reaction
• Examples: Oxidative phosphorylation and the
oxidation reactions of the electron transport chain
Oxidative Phosphorylation - An
Introduction
• Coupling of ATP synthesis with the reactions of

the ETC is related to the movement of protons
(H+ ions) across the inner mitochondrial
membrane
• Complexes I, III, and IV of ETC chain have a
second function
• Serve as “proton pumps” transferring protons from the
matrix side of the inner mitochondrial membrane to
the intermembrane space
• For every two electrons passed
through ETC, four protons cross the
inner mitochondrial membrane
through complex I, four through
complex III, and two more though
complex IV
• This proton flow causes a buildup of
H+ in the intermembrane space
• This high concentration of
protons passing through ATP
synthase becomes the basis for
the ATP synthesis

FIGURE 23.18 - A SECOND FUNCTION FOR
PROTEIN COMPLEXES I, III, AND IV

ATP Formation
• For each mole of NADH oxidized in the
ETC, 2.5 moles of ATP are formed
• For each mole of FADH2 oxidized in the
ETC, only 1.5 moles of ATP are formed
• For each mole of GTP hydrolyzed, one
mole of ATP is formed
• Ten molecules of ATP are produced for
each acetyl CoA catabolized

>90% of inhaled oxygen via
respiration is consumed during
oxidative phosphorylation
Reactive
Oxygen Remaining O2 is converted to several
highly reactive oxygen species (ROS)
Species within the body, which include:
(ROS) • Hydrogen peroxide (H2O2)

• Superoxide ion (O2-)
• Hydroxyl radical (OH)
• Superoxide ion and hydroxyl radicals have
an unpaired electron

Reactive Oxygen Species (ROS)
• Can also be formed due to external influences such
as polluted air, cigarette smoke, and radiation
exposure
• Are both beneficial as well as problematic within the
body
• Example: White blood cells produce a significant
amount of superoxide free radicals via the following
reaction to destroy the invading bacteria and viruses
2O2 + NADPH ⎯⎯
→ 2O2- + NADP+ + H+
Reactive Oxygen Species (ROS)
• ROS formed are quickly converted to non-toxic
species
superoxide
2O2- + 2H + ⎯⎯⎯⎯
→ H 2O2 + O2
dismutase
2H 2O2 ⎯⎯⎯
catalase
→ 2H 2O + O2
• About 5% of ROS escape destruction by superoxide

dismutase and catalase enzymes

• Antioxidant molecules
present in the body help trap
ROS species
• Vitamin E
Reactive Oxygen • Vitamin C
Species (ROS) • Glutathione (GSH)
• Beta-carotene
• Major family of antioxidant
phytochemicals are
flavonoids

• Structurally modified B vitamins
function as coenzymes in metabolic
pathways
• Four B vitamins participate in various
B Vitamins and reactions:
• Niacin—as NAD+ and NADH
the Common • Riboflavin—as FAD, FADH2, and FMN
Metabolic • Thiamin—as TPP
Pathway • Pantothenic acid—as CoA
• In the absence of these B vitamins, the

body would be unable to utilize
carbohydrates, proteins, and fats as
energy sources

Biochemical Energy Production

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BIOCHEMICAL

Copyright ©2016 Cengage Learning. All Rights Reserved. 1

Source of energy for the functioning of the

Also needed for many of the cellular

Copyright ©2016 Cengage Learning. All Rights Reserved. 2

Metabolic Anabolism: All metabolic reactions in

Copyright ©2016 Cengage Learning. All Rights Reserved. 3

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Cytoplasm: Water-based material of a eukaryotic

Cell Mitochondrion: Generates most of the energy

Function Ribosome - Site for protein synthesis

Nucleus - Site where DNA is found

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•The net energy produced in these reactions is

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Cytidine triphosphate (CTP) -

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FAD is the oxidized form FADH2 is the reduced form

Typical cellular reaction in

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• Has coenzyme functions in metabolic redox

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• Derivative of vitamin B pantothenic acid

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• Polyfunctional acids formed as intermediates of

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• Begins in mouth (saliva contains starch-digesting enzymes),

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• The small molecules from Stage 1 are further

• Takes place inside the mitochondria

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•Takes place in mitochondria

• Is the series of biochemical reactions in which

• Summary of citric acid cycle reactions:

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(1) the condensation of (2) hydrolysis of the

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•Step 6: Oxidation of Succinate.

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Series of biochemical reactions in which

Lost energy is used to synthesize ATP in oxidative phosphorylation

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Organized into four distinct protein complexes and two mobile

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Complex II: Contains only four subunits including

Coenzyme Succinate is converted to fumarate

• CoQ is the final recipient of the electrons from