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Part two

9-principle of biochemistry
Metabolism and biological energy
1-Glycolysis

Course code: HFB 324


Credit hours: 3 hours
Dr. Siham Gritly

Dr. Siham M.O. Gritly 1


Glossary
• endergonic reaction; is a chemical reaction in which the
standard change in free energy is positive, and energy is
absorbed
• exergonic reaction; is a chemical reaction where the
change in the Gibbs free energy or ∆G is negative,
indicating a spontaneous reaction
• acetyl-CoA coenzyme A attached to an acetate group
(CH3COO−) via a high-energy bond; acetyl-CoA is the
central molecule of metabolism
• Aerobe oxygen-requiring organism; uses glycolysis, the
tricarboxylic acid cycle and the electron transport chain to
produce ATP
Dr. Siham M.O. Gritly 2
• spontaneous reaction-A process that will occur without any energy
input from the surroundings. A process that will occur on its own
• Non-spontaneous reaction- A reaction which cannot occur without
the input of work from an external source. ΔG > 0 for
nonspontaneous reactions at constant tem and pressure
• Anaerobe non-oxygen-requiring organism; uses only glycolysis to
produce ATP
• electrostatic repulsion the force repelling two ionic groups each
holding the same charge
• Entropy thermodynamic measure of disorder
• metabolism: reactions that living organism carry out consist of
catabolism: energy and simple molecules are formed from complex
molecules and anabolism: synthesis of complex biomolecules using
energy

Dr. Siham M.O. Gritly 3


• nonequilibrium thermodynamics the steady-state conditions that
exist in vivo
• physiological free energy the amount of energy available to a
organism based on both the standard free energy of a reaction and
also the actual concentrations of the reactants and products of this
reaction in vivo
• Resonance The property of a compound having simultaneously the
characteristics of two or more structural forms that differ only in the
distribution of electrons. Such compounds are highly stable and
cannot be properly represented by a single structural formula.
• standard free energy thermodynamic value; amount of energy
released when a mole of reactant is allowed to come to equilibrium
(with product) under fixed conditions of temperature, pressure, and
hydronium ion concentration

Dr. Siham M.O. Gritly 4


• elimination reaction in which a molecule decomposes
into two molecules of unequal size
• metabolic pathway tightly regulated, interconnected set
of biochemical reactions in which some chemicals
(nutrients) are used up and other chemicals are produced
(e.g., glucose → ATP)
• Oxidation loss of one or a pair of electrons
• redox reaction in which electrons are transferred from
one molecule to another
• Reduction gain of one or a pair of electrons
Dr. Siham M.O. Gritly 5
Lecture outlines
• Bioenergetics and the lows of thermodynamics
• the difference between standard free energies ( ΔG°′) and
physiological free energies (ΔG).
• ATP-Cycle
• Metabolism: Oxidation and Reduction Processes
• Glucose metabolism and the role of ATP
• glycolysis,
• Tri-carboxylic Acid cycle,
• Electron transport system
• Pentose Shunt Pathway,
• Glycogenolysis, Gloconegensis,

Dr. Siham M.O. Gritly 6


Bioenergetics and the lows of
thermodynamics
• Bioenergetics (thermodynamics), is the study
of energy changes accompanying by chemical
reactions.
• It is the quantitative study of energy
relationships and energy conversions in
biological systems.
• Biological energy transformations obey the
laws of thermodynamics

Dr. Siham M.O. Gritly 7


• The inter-conversions of energy led to two
fundamental lows of thermodynamics
• 1-transformation of energy from one state to
another is unchanged

• The entropy (is the energy in a state of


randomness) of the system must increase if the
process is to occur spontaneously

Dr. Siham M.O. Gritly 8


• All chemical reactions are influenced by two forces:
• 1- enthalpy, H the tendency to achieve the most stable
bonding state (enthalpy, H,)
• 2- entropy, S. is the energy in a state of randomness
or disorder of the system
• The net energetic force in a reaction is ΔG, the free-
energy change, free-energy change represents the net
effect of these two factors under condition of constant
tem. & pressure; ΔG = ΔH - TΔS.

Dr. Siham M.O. Gritly 9


• ΔG = ΔH - TΔS.
• ΔG =change of free energy in the reacting
system
• ΔH =change in its heat content or enthalpy
• T = temperature at which the reaction takes
place
• ΔS = change in entropy of the reaction

Dr. Siham M.O. Gritly 10


Concept of standard free energy ( ΔG°) and
physiological free energies (ΔG).

• standard free energies ( ΔG°′) when reactant are present


in concentration of 1.0 mol/L under standard temperature
and pressure
• At pH 7 in a biological systems is designated ΔG°.
Standard free energy change can be calculated from the
equilibrium constant K’eq (-2.303 RT log K’eq)
• ΔG°′= -2.303 RT log K’eq
• Where R=gas constant
• T= absolute temperature
• K’eq = equilibrium constant
Dr. Siham M.O. Gritly 11
Difference between ΔG° & ΔG

• During spontaneous chemical or physical reaction the free


energy is decrease (negative ΔG becomes less negative as
the reaction continue till the reaction at equilibrium (zero)
• spontaneous reaction-is a process that will occur without
any energy input from the surroundings. A process that will
occur on its own

• Standard free energy ΔG° can be positive, negative or


zero depending on the equilibrium constant of the
reaction. ΔG° usually unchanging (constant tem. & pH)

Dr. Siham M.O. Gritly 12


Bioenergetics
Metabolism and biological energy
• Living cells continuously carry out work and thus
require energy. Energy is found in different forms;
Electrical, Chemical, Mechanical, Heat, Light energy
all are interconvert

• Energy is needed for; -muscular secretion, secretion


of the glands, synthesis of new substances, regulation
of body temperature, absorption of food from gastro-
intestinal tract, for the basal metabolism e.g. heart
beat, respiration ect
Dr. Siham M.O. Gritly 13
• There are two types of useful energy in living
organism
• 1-free energy ∆G (potential energy) available
energy; useful amount of available energy to perform
work (under a constant temperature, and pressure).
This form of energy trapped in the form of ATP
• 2-heat energy; can do work only through a change
in temperature, heat is not source of energy but is
used to facilitate the chemical energy

Dr. Siham M.O. Gritly 14


• The primary energy-releasing pathways of
metabolism involve the breakdown and
synthesis of; carbohydrates, lipids, amino acids
• There are two types of metabolic pathways:
• anabolic, involving the synthesis of
biochemical from simpler molecules where
energy is required (ΔG +ve value endergonic
reaction where energy is needed)

Dr. Siham M.O. Gritly 15


• catabolic, involving the breakdown of
biochemicals into simpler compounds with
releasing of energy (ΔG -ve in value
exergonic reaction)
• During catabolism of nutrients some of free
energy is trapped to form ATP from ADP
(adenosine diphosphate) and inorganic
phosphate Pi

Dr. Siham M.O. Gritly 16


Exergonic and endergonic reactions
Endergonic reactions are those that require an input of energy
Exergonic reactions are those that release energy

• In a biological reaction enthalpy ΔH is equal to ΔE, the


total change in internal energy of the reaction expressed
as;
ΔG = ΔE – TΔS
• If ΔG -ve in value =less energy reaction proceeds with
loss of energy this is known as exergonic (catabolism)
• If ΔG +ve value =more free energy than the reactant the
reaction proceeds only if free energy gained endergonic
(anabolism)
• If ΔG is zero this means that the reaction is at equilibrium

Dr. Siham M.O. Gritly 17


Coupling reactions
ATP-Cycle

• Most coupling reactions use the break down of


adenosine triphosphate (ATP) as the
intermediate process to drive chemical
synthesis
• ATP is the chemical link between catabolism
and anabolism. Its exergonic conversion to
ADP and Pi, or to AMP and PPi, is coupled to
a large number of endergonic reactions and
processes

Dr. Siham M.O. Gritly 18


• ATP undergo hydrolysis (breaking down) to
ADP and Pi by donate its chemical energy to
energy requiring reaction

• Each reaction is catalyzed by an enzyme and is


linked to other reactions through a pathway

Dr. Siham M.O. Gritly 19


ATP
adenosine tri-phosphate is intermediary compound that has the ability to enter in many
reactions
.

• *ATP is a source of high-energy phosphate .


present everywhere in the cytoplasm and
nucleoplasm of all cells and essentially for all
the physiological mechanisms that required
energy.
• it is nucleotide compound composed of
adenine (nitrogen base), ribose (pentose sugar)
and three phosphate radicals.
• *ATP is known as energy currency of the cells
Dr. Siham M.O. Gritly 20
• *within the cells the nutrients amino acids, fatty acids
and glucose react with oxygen (oxidation) to form
carbon dioxide, water and energy.
• Energy produced used to form adenosine tri-phosphate
• *creatine phosphate (CP) is another high energy rich
compound and is considered the reservoir of high
energy phosphate. this compound is stored in the body
in large quantities than ATP.
• *the rapid chemical changes involved in ATP
production and breakdown are brought about by the
action of enzymes, co-enzymes and hormones.

Dr. Siham M.O. Gritly 21


ATP (adenosine triphophate) structure
is nucleotide phosphate ester to a ribose and adenosine nucleoside)
(with Mg++ element attached active region)

ATP or adenosine tri-phosphate has the ability


to enter all cells. it is known as energy currency
of the cells Dr. Siham M.O. Gritly 22
Carbohydrates metabolism

• Carbohydrates utilized by the body in


different ways;
*for the immediate energy needs through oxidation to
carbon dioxide and water via the processes of
glycolysis and tri-carboxylic acid cycle

• *stored as glycogen in the liver and muscles tissue


through the process glycogenesis.

• *converted to fatty acids and stored as triglyceride.


Dr. Siham M.O. Gritly 23
• *convert to other necessary CHO such as
ribose, fructose and deoxyribose which is
necessary for the formation of genetic material
DNA.
• *some become the carbon skeletons for
production of essential amino acids

Dr. Siham M.O. Gritly 24


• The first step in the metabolism of digestible
carbohydrate is the conversion of the higher
polymers to simpler, soluble forms that can be
transported across the intestinal wall and
delivered to the tissues.
• The breakdown of polymeric sugars begins in
the mouth by the Saliva α amylase that begins
the digestion of carbohydrates.
• Passes through the esophagus to stomach

Dr. Siham M.O. Gritly 25


• in the stomach, acid hydrolysis contributes to its
degradation;

• The mixture of gastric secretions, saliva, and food, (chyme),


moves to the small intestine and absorbed by blood as
glucose

• the portal blood carry glucose to the liver. in the liver


galactose and fructose are converted to glucose by
specific enzymes.

• Glucose then enter into; metabolic pass ways (glycolysis)


Dr. Siham M.O. Gritly 26
Metabolic pathways
Glucose metabolism
• Glucose metabolism is a process that make
energy locked in glucose molecules is
available for all physiological functions of the
cells and tissues.

• Glucose is the main source of energy. it is the


final common pathway of carbohydrates, fats
and proteins additional to alcohols

Dr. Siham M.O. Gritly 27


Glucose metabolism

Dr. Siham M.O. Gritly 28


Glycolysis pathway
Oxidation of glucose
• Glycolysis is process of splitting of glucose
molecule (6C) to form 2 molecules of pyruvic
acid (3C) with the release of energy
• Free energy released in this process is stored as
2 molecules of ATP, and 2 molecules of NADH.

• Glucose + 2 ADP + 2 NAD+ + 2 Pi ——> 2


Pyruvate + 2 ATP + 2 NADH + 2 H+

Dr. Siham M.O. Gritly 29


• The 2 NADH molecules are oxidized in
mitochondria under aerobic condition and the free
energy released to synthesize ATP by oxidative
phosphorylation reaction.
• 2 molecules of pyruvic acid (3C) Under the aerobic
condition is catabolized further in mitochondria
through pyruvate dehydrogenase in citric acid cycle
where all the carbon atoms are oxidized to CO2.
• The free energy released is used in the synthesis of
ATP, NADH and FADH2 again .

Dr. Siham M.O. Gritly 30


Anaerobic Glycolysis

• Under anaerobic conditions pyruvate is


converted to lactate by the enzyme lactate
dehydrogenase (LDH),
• the lactate is transported out of the cell into the
blood circulatory system.
• 2 lactic acid is the end product of glycolysis
under anaerobic condition)

Dr. Siham M.O. Gritly 31


• Aerobic Glycolysis occurs by 10 steps of
chemical reactions, each reaction is catalyzed by
one specific protein enzyme.

• it has 2 phases,
• Phase 1-energy requiring reactions (ATP)
Preparatory phase
• Phase 2-energy releasing reactions the pay-off
phase
Dr. Siham M.O. Gritly 32
Glycolysis Phase one –energy requiring
Ref. Michael W King, PhD | © 1996–2012 themedicalbiochemistrypage.org, LLC | info @ themedicalbiochemistrypage.org

In the preparatory
phase of glycolysis two
molecules of ATP have
been consumed. The
hexose chain has been
cleaved into two triose
phosphates

Cleavage of 6-
carbon sugar
phosphate to
two 3-carbon
sugar
phosphate

Dr. Siham M.O. Gritly 33


Glycolysis Phase two–energy yeilding
Ref. Michael W King, PhD | © 1996–2012 themedicalbiochemistrypage.org, LLC | info @ themedicalbiochemistrypage.org

The two high energy


First ATP forming
intermediates whose
reaction-
oxidations are
substrate level
coupled to ATP
phosphorylation
synthesis are 1,3-
bisphosphoglycerate
& phospho-
enolpyruvate.
second ATP
forming reaction-
substrate level
phosphorylation

Dr. Siham M.O. Gritly 34


Step 1 phosphorylation of glucose
Hexokinase reaction
One molecule of ATP is required during this reaction
• The ATP-dependent phosphorylation of glucose to form
glucose 6-phosphate (C 6)
• This reaction is catalyzed by tissue-specific isoenzymes
known as hexokinases

Glucose (C6H12O6) + hexokinase + ATP →


ADP + Glucose 6-phosphate (C6H 11O6P1)
Dr. Siham M.O. Gritly 35
Step 2 Phosphohexose Isomerase
Phosphoglucoisomerase
isomerization, in which G6P is converted to its isomer fructose 6-phosphate (F6P). The
reaction is freely reversible at normal cellular concentrations of the two hexose phosphates

Glucose 6-phosphate (C6H11O6P1) + Phosphoglucoisomerase


→ Fructose 6-phosphate (C 6H11O6P1)
Dr. Siham M.O. Gritly 36
• The second reaction of glycolysis is an isomerization, in
which G6P is converted to fructose 6-phosphate (F6P).
• The enzyme catalyzing this reaction is phosphohexose
isomerase (also known as phosphoglucoisomerase).

• This enzyme catalyzes the reversible isomerization of


G6P (an aldohexose) to F6P (a ketohexose).This
specific enzyme requires Mg ++ for its activity.

Dr. Siham M.O. Gritly 37


step 3- 6-Phosphofructo-1-Kinase (Phosphofructokinase-1, PFK-1):

• The next reaction of glycolysis involves the


utilization of a second ATP to convert F6P to
fructose 1,6-bisphosphate (F1,6BP).

• This reaction is catalyzed by 6-


phosphofructo-1-kinase, or known as
phosphofructokinase-1 or PFK-1 (Class2
Transferases).

Dr. Siham M.O. Gritly 38


• Transfer of phosphoryl group from ATP to C-1
of F6P to produce Fructose 1,6 biphosphate.

Fructose 6-phosphate (C6H11O6P1) +


phosphofructokinase + ATP → ADP +
Fructose 1, 6-bisphosphate (C6H10 O6P2)M.O. Gritly
Dr. Siham 39
Step 4 Aldolase
Class 4. Lyases-
cleavage step-Cleavage of Fructose 1,6 biphosphate

• Aldolase catalyses the hydrolysis of F1,6BP


into two 3-carbon products (trioses):
• *dihydroxyacetone phosphate (DHAP) (a
ketose).

• *glyceraldehyde 3-phosphate (G3P) (an


aldose) .
Dr. Siham M.O. Gritly 40
• the standard free energy change is positive in the
forward direction, meaning it requires energy.
• Fructose 1, 6-bisphosphate (C6H10O6P2) + aldolase → Dihydroxyacetone phosphate
(C3H5O3P1) + Glyceraldehyde phosphate (C3H5O3P1)

Dr. Siham M.O. Gritly 41


Step 5 Triose Phosphate Isomerase:
• Triose phosphate isomerases reaction:
Conversion of Dihydroxyacetone phosphate to
glyceraldehyde 3 Phosphate are both 3 carbon
molecules .

Class 5. Isomerases- enzyme that catalyzes the


structural rearrangement of isomers
Dr. Siham M.O. Gritly 42
• Net result for steps 4 and 5:
• Fructose 1, 6-bisphosphate (C6H10O6P2) ↔ 2
molecules of Glyceraldehyde phosphate
(C3H5O3P1)

Dr. Siham M.O. Gritly 43


Second phase of glycolysis (the payoff phase)

• Step 6 The second phase of glucose catabolism where


the energy-yielding glycolytic reactions.
• In this reaction two high energy phosphate bonds are
produced. These 2 high energy phosphoryl groups are
transferred to ADP to generate 2 molecules of ATP
and NADH.
• NAD+ is the cofactor in this reaction which acts as an
oxidizing agent. The free energy released in the
oxidation reaction is used in the formation of
acylphosphate.
Dr. Siham M.O. Gritly 44
Step 6 Glyceraldehyde-3-Phosphate Dehydrogenase (Class 1
Oxidoreductase).
The first step of the second phase of glycolysis is the oxidation and
phosphorylation of glyceraldehydes 3-phosphate to form 1, 3-
bisphosphoglycerate
.

Dr. Siham M.O. Gritly 45


• First the enzyme transfers a hydrogen (H-) from
glyceraldehyde 3 phosphate to the oxidizing agent
nicotinamide adenine dinucleotide (NAD+) to form
NADH

• triose phosphate dehydrogenase adds a phosphate (P)


from the cytosol to the oxidized glyceraldehyde
phosphate to form 1, 3-bisphosphoglycerate

• The oxidation of an aldehyde to the carboxylic acid is


an exergonic process.
Dr. Siham M.O. Gritly 46
Step 7
Phosphoglycerate Kinase
The high-energy phosphate of 1,3-BPG is used to form ATP and 3-phosphoglycerate
(3PG) by the enzyme phosphoglycerate kinase.

(substrate level phosphorylation)


Dr. Siham M.O. Gritly 47
• In Phosphoglycerate kinase Reaction the
phosphoryl group Transfer from 1,3
bisphosphoglycerate to ADP to form ATP
(substrate level phosphorylation)

Dr. Siham M.O. Gritly 48


Step 8 Phosphoglycerate Mutase
In the eighth reaction of glycolysis, 3-phosphoglycerate is converted to 2-
phosphoglycerate by the enzyme phosphoglycerate mutase. This reaction is an
isomerization

the phosphoryl group from the C-3 of


the intermediate is transferred to the
enzyme and 2-PG is released.
Dr. Siham M.O. Gritly 49
• Step 8 mutase enzyme (group transfer) convert
3-phosphoglycerate into 2-phosphateglycerate.
The enzyme catalyzes the reversible move of
the phosphate ester between C2 and C3 of
glycerate.
• Mg2+ is required for enzymatic activity.

Dr. Siham M.O. Gritly 50


Step 9 Enolase
In the ninth reaction of glycolysis, enolase catalyzes a
dehydration reaction of 2-phosphoglycerate in which water is
eliminated, to yield phosphoenolpyruvate.

Dehydration of 2-phosphoglycerate to
phosphoenolpyruvate

Dr. Siham M.O. Gritly 51


• Enolase is the enzyme that catalyzes the
reversible elimination reaction of water to
convert 2-phosphoglycerate into phosphoenol
pyruvate.
• Mg2+ required for enzymatic activity.
• Enolase dehydrates the substrate to form a
high energy phosphoenol

Dr. Siham M.O. Gritly 52


Step 10-Pyruvate Kinase
The tenth and final reaction of glycolysis is catalyzed by pyruvate
kinase, which converts phosphoenolpyruvate to pyruvate as it transfers
the phosphoryl group to ADP to yield ATP

This is the second substrate


level phosphorylation reaction
of glycolysis Dr. Siham M.O. Gritly exergonic reaction 53
• 2 molecules of phosphoenolpyruvate (PEP)
(C3H3O3P1) + pyruvate kinase + 2 ADP → 2
molecules of pyruvic acid (C3H4O3) + 2 ATP
keto form of pyruvate

The loss of phosphate by PEP leads to the


production of pyruvate in an unstable enol
form, which spontaneously tautomerizes to the
more stable, keto form of pyruvate.
Dr. Siham M.O. Gritly 54
Pyruvate oxidation
Pyruvate dehydrogenase complex

The overall reaction for pyruvate oxidation is: 2-carbon molecule


Pyruvate + CoA + NAD+ → Acetyl-CoA + NADH
Dr. Siham M.O. Gritly 55
+ H+ + CO2
The last step of glycolysis produces 2 pyruvate molecules in
the cytosol, In mitochondrial matrix and by the action of
pyruvate dehydrogenase complex Pyruvate oxididised to
acetyle co A
• In this reaction (pyruvate oxidation);
• two electrons and two protons are released;
• the electrons and one proton are accepted by NAD+,
which make it into NADH,
• the other proton is simply released as an H+ ion.
• The acetyl group is attached to coenzyme A, a carrier
formed from nucleotides,
• formed molecule, Acetyl-CoA, is transported to the citric
acid cycle,
• the citric acid cycle occurs in the mitochondrial matrix.
Dr. Siham M.O. Gritly 56
Regulation of Glycolysis

• There are three steps in glycolysis that have


enzymes which regulate the flux of glycolysis.

The hexokinase (HK); inhibit by its production of


glucose-6-phosphate
The phoshofructokinase (PFK); increase in response
to insulin and decrease in response of gucagon
The pyruvate kinase; increase in concentration with
high insulin and decrease with glucagon
Dr. Siham M.O. Gritly 57
The importance of glycolysis pathway

• Glycolysis is important part of glucose metabolism


because;
• -It produces large amount of energy,
• -occur in all cells,
• Ability to provide ATP in the absence of oxygen
• Generates intermidiates for synthesis pathway;- pyruvate tp
alanine (transaminated), acetyyle-C A for fatty acid
synthesis, glycerpl-3-phosphate which form the backbone
of triacylglycerol
• -common pathway in breaking down all macro-nutrients
(proteins, fats and carbohydrates)
Dr. Siham M.O. Gritly 58
Some of Genetic diseases associated with glycolysis pathway

• Genetic deficiency of pyruvate kinase in


erythrocyte leads to chronic hemolytic anemia.
• Cancer; Malignant rapidly growing tumor cells
typically have glycolytic rates that are up to 200
times higher than those of their normal tissues of
origin
• Alzheimer's disease; Dis-functioning glycolysis
or glucose metabolism has been associated with
Alzheimer's disease
Dr. Siham M.O. Gritly 59
Glycolysis

Dr. Siham M.O. Gritly 60


Glucose metabolism---continue next lecture

• Tri-carboxylic Acid cycle,


• Electron transport system
• Pentose Shunt Pathway,
• Glycogenolysis, Gloconegensis

Dr. Siham M.O. Gritly 61


References
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themedicalbiochemistrypage.org
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and Company
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• Emsley, John (2011). Nature's Building Blocks: An A-Z Guide to the Elements (New
ed.). New York, NY: Oxford University Press. ISBN 978-0-19-960563-7.

Dr. Siham M.O. Gritly 62


• Koppenol, W. H. (2002). "Naming of New Elements (IUPAC Recommendations 2002)" (PDF). Pure and
Applied Chemistry 74 (5): 787–791. doi:10.1351/pac200274050787.
http://media.iupac.org/publications/pac/2002/pdf/7405x0787.pdf.
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• Murry K. Robert, Granner K. daryl, Mayes A. peter, Rodwell W. Victor (1999). Harpers Biochemistry.
Appleton and Lange , twent fifth edition
• Cooper GM 2000. The Central Role of Enzymes as Biological CatalystsThe Cell: A Molecular Approach. 2nd
edition. Sunderland (MA): Sinauer Associates; 2000
• Campbell, Neil A.; Brad Williamson; Robin J. Heyden (2006). Biology: Exploring Life. Boston, Massachusetts:
Pearson Prentice Hall
• A. Burtis, Edward R. Ashwood, Norbert W. Tietz (2000), Tietz fundamentals of clinical chemistry
• Maton, Anthea; Jean Hopkins, Charles William McLaughlin, Susan Johnson, Maryanna Quon Warner, David
LaHart, Jill D. Wright (1993). Human Biology and Health. Englewood Cliffs, New Jersey, USA: Prentice Hall.
pp. 52–59
• Maitland, Jr Jones (1998). Organic Chemistry. W W Norton & Co Inc (Np). p. 139. ISBN 0-393-97378-6.
• Nelson DL, Cox MM (2005). Lehninger's Principles of Biochemistry (4th ed.). New York, New York: W. H.
Freeman and Company.
• Matthews, C. E.; K. E. Van Holde; K. G. Ahern (1999) Biochemistry. 3rd edition. Benjamin Cummings.
• http://wiki.answers.com/Q/What_is_dehydration_synthesis#ixzz2BuiK645

Dr. Siham M.O. Gritly 63

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