Biochemistry-8-1-lipids 1
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•Found mostly in fatty tissues, membranes, and other nonpolar biological structures
•Nonpolar: hydrophobic (“water hating”) or lipophilic (“fat loving”)
•Polar: hydrophilic (“water loving”) or lipophobic (“fat hating”)
Lipids
Definitions
Lipid (Greek: lipos, fat): organic molecule of biological origin that is insoluble in water
and soluble in nonpolar solvents (CH2Cl2, CH3CH2OCH2CH3, etc.)
Example: A phospholipid
O
O
O
O
O P
O
O
OCH2CH2NH3
•Lipid solubility properties due to large nonpolar regions
Biochemistry-8-1-lipids 2
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Lipids
Categories
General Categories of Lipids
Fatty acids
Waxes
Triacylglycerols
Phospholipids
Prostaglandins
Steroids
Lipophilic vitamins
Terpenes •Produced mostly by plants
Biochemistry-8-1-lipids 3
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Biological role
1. Fuel
-(adipocytes)
Oxidation in mitochondria
2. Nutrients
Amphipathic lipids- build cell membrane (ph.l.,
glycolipids, cholesterol)
3. Insulation
Mechanical and thermal insulation
4. special task
-signaling function, cofactors
Biochemistry-8-1-lipids 4
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Fatty Acids
Fatty acid: unbranched carboxylic acid
•Most have even number of carbons: two carbons added at a time during biosynthesis
•12-20 carbons most common
•Most biologically-important fatty acids have 18 carbons: stearic, oleic, and linoleic acids
•Main biological function: component of other lipids
•Categorized by C=C in chain: saturated (no C=C) or unsaturated (one or more C=C)
Saturated fatty acids
Lauric acid (12 C)
Myristic acid (14 C)
Palmitic acid (16 C)
Stearic acid (18 C)
Arachidic acid (20 C)
OH
O
OH
O
OH
O
OH
O
OH
O
Biochemistry-8-1-lipids 7
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Fatty Acids
•Monounsaturated: contains one C=C
•Polyunsaturated: contains more than one C=C
•Cis C=C much more common than trans C=C
OH
O
Oleic acid (C18)
OH
O
Linoleic acid (C18)
OH
O
Linolenic acid (C18)
Unsaturated fatty acids
Some important unsaturated fatty acids
Arachidonic acid (C20)
OH
O
C
C C
C
HH
C
C H
C
CH
Cis alkene
Common in fatty acids
Trans alkene
Rare in fatty acids
Biochemistry-8-1-lipids 8
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Triacylglycerols
Triacylglycerol (triacylglyceride): fatty acid triester of glycerol (glycerin)
•Triacylglycerol = fat if solid at room temperature; oil if liquid
•Most abundant natural lipids
•Main biological function: energy storage
•Hydrolysis (“water breaking”) of animal fats yields soap
- 3 H2O
OH
OH
OH + R'CO2
- Na+
+ RCO2
- Na+
+ R"CO2
- Na+
mixture = soap
OH
OH
+ RCOOH
+ R"COOH
HO + R'COOH
Glycerol Fatty acids
NaOH, H2O
heat
Triacylglycerol
O
O
O
R
O
O
R"
R'
O
Biochemistry-8-1-lipids 9
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Lower chain: eight carbons
Upper chain: seven carbons
Prostaglandins
Prostaglandin: molecule having the prostanoic acid skeleton
Prostanoic acid
OH
O
Prostaglandin F2a (PGF2a)
COOH
OH
H
H
HO
HO
•Nomenclature: based on stereochemistry, number of OH, C=C, C=O groups
•Biological functions: mostly as regulators and signal molecules
- cause constriction or dilatation in vascular and other smooth muscle cells
- regulate aggregation and disaggregation of platelets
- sensitize spinal neurons to pain
- regulate inflammatory mediation, calcium movement, hormones
- control cell growth
Biochemistry-8-1-lipids 10
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Phosphate esterFatty acid esters
Phospholipids
Phospholipid: Glycerol esterified with two fatty acids and one phosphate group
•Fatty acids are usually palmitic (C16), stearic (C18), and oleic (C18)
•Second most abundant group of natural lipids
•Main biological function: cell membranes (phospholipid bilayer)
•Hydrophobic effect: hydrophobic tails avoid water
Simplified cell membrane
cell exterior (aqueous)
cytoplasm (aqueous)
hydrophobic tails
hydrophilic heads
Generic phospholipid structure
R"
O
O
O
O
O
R'
P
O
O
OCH2CH2NR3
Biochemistry-8-1-lipids 11
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Arachidonic acid
O
O
Prostaglandin I2
(PGI2; a prostacyclin)
prostacyclin synthase
OH
PGH-PGE
isomerase
COOH
Prostaglandin E2 (PGE2)
PGH-PGF2a reductase
OHHO
Prostaglandin H2 (PGH2)
O
cyclooxygenase-2 (COX-2)
and other enyzmes
COOH
OH
COOH
HO
O
OH
COOH
HO
HO
Prostaglandin F2a (PGF2a)
COOH
Inhibited by aspirin, Vioxx, etc.
Prostaglandins
•Biological origin: prostaglandin cascade
•May occur at wound site, leading to inflammation
•in vivo half-life typically 5 minutes or less
•Similar structures but wide range of functions
Biochemistry-8-1-lipids 12
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Steroids
Steroid: a molecule having the ring system shown below
Steroid skeleton
A B
C D
Steroid example: cholesterol
HO
H3C
H H
H
H3C
•Shape: fairly flat and fairly rigid
•Verify and explore with a model
Biochemistry-8-1-lipids 13
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Steroids
Steroid Biosynthesis
•More than sixty steps from acetyl CoA  cholesterol
H3C S-CoA
O
H3C
H3C
HO
CH3
H
acetylcoenzyme A
many enyzmatic steps
H3C
H3C
HO
H
squalene oxide
squalene
epoxidase
lanosterol
H
H
25 or more
enyzmatic
steps
epoxide
squalene
squalene
oxidocyclase
cholesterol
O
Cholesterol is biological precursor to all other steroids
Biochemistry-8-1-lipids 14
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Steroids
Steroid categories and examples
Sex hormones:
Corticoid hormones:
Testosterone (an androgen)
O
H3C
H3C
H
H H
OH
Estrone (an estrogen)
HO
H3C
H
H H
O
Cortisone
•A glucocorticoid hormone
•Regulates inflammation
•Regulates glucose metabolism
O
H3C
H3C
H
H H
O
CH2OH
OH
O
Aldosterone
•A mineralocorticoid hormone
•Regulates Na+/K+ balance
O
H3C H
H H
HO
O
CH2OH
O
H
•Synthesized in the adrenal complex
•Regulate metabolic processes
Biochemistry-8-1-lipids 15
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Steroids
Steroid categories and examples
Bile acids
•Aid in digestion by emulsifying fats in intestine
HO OH
H H
HO
H3C
H
H
CH3
H3C
CO2H
Steroids have similar structures but wide range of functions
Biochemistry-8-1-lipids 16
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Lipophilic Vitamins
Vitamin: an organic compound, other than fat, protein or carbohydrate, required for the
normal growth and maintenance of animals
•Very broad range of structures and functions
Vitamin E
•Mixture of isomers; a-tocopherol most important
•Protects against oxidative damage to cells from radicals
a-Tocopherol
Hydrophobic antioxidant vitamin
O
CH3
HO
CH3
CH3
CH3
CH3 CH3
Vitamin C (ascorbic acid)
Hydrophilic antioxidant vitamin
O
HO OH
O
OH
OH
H
Biochemistry-8-1-lipids 17
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Lipophilic Vitamins
Vitamin A (retinol)
•Essential to vision
•Incorporated into rhodopsin (photon-harvesting protein)
OH
Biochemie-8-1-lipidy 18
Lipid metabolism
metabolism of TG and FA
100 g/day
energy source
metabolism of
structural lipids
2 g/day
Compared to most of the carbohydrates and FA are lipids (mainly TG, FA,
esterified cholesterol) hydrophobic (non-polar). However, the environment in
which the metabolism of nutrients takes place is filled with water which is polar.
Therefore, in the body they are natural surfactants, able to receive, transport
and enable the metabolism of lipids.
20
Triglycerides (TG)-90%
phospholipids (PL)
Cholesteryl ester (CHE)
glycolipids (GL)
lipophilic vitamins (LV)
primary products :
free FA
2-monoacylglycerols
lysophospholipids
cholesterol
lipophilic vitamins
pancreatic lipase
+ colipaseBile acids
Lingual and
Gastric Lipase
Resorption to the
enterocytes
in the form of mixed
micelles (particles <20 nm)
Digestion of lipids
21
CH2
CH
O C
O
CH2
OC
O
O
C
O
Cleavage of lipids in the intestine by
pancreatic enzymes
• pancreatic lipase
triacylglycerol  2-monoacylglycerol + 2 FA
< 1/4 TG
triacylglycerol  glycerol + FA
Orlistat – Anti-obesity agent inhibits lipase digestive tract, reduces absorption of about 30% of dietary fat
Biochemie-8-1-lipidy
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Lipids are hydrophobic
triacylglycerols
free fatty acids
esterified cholesterol
Their transport and metabolism takes place
through various natural surfactants.
The first problem with the fact that they are non-polar lipids and internal
environment of our bodies is polar occurs in the small intestine.
Solution to the problem is the formation of mixed micelles, which is provided by
surfactants of the small intestine as bile acids, phospholipids, salts of free FA
(soaps) and 2-glycerides. Nonpolar lipids "hide" between polar surfactants and in
this polar packaging can be transported into the cells of the intestinal mucosa.
Biochemie-8-1-lipidy 23
Natural surfactants in the
absorption of fat
Form a micelle, which enters into the
enterocytes
Surfactant Type Origin
Bile acids
2-Acylglycerol
Anionts of FA
Phospholipids
anionic
nonionic
anionic
amphoteric
from cholesterol in the liver
TAG hydrolysis in the intestine
TAG hydrolysis in the intestine
food
24
Resorption of lipid cells of the intestinal
mucosa (Mostly in the jejunum, bile acids to the ileum)
mixed micelles
brush border epithelial cells
The diameter of
<20 nm Passive diffusion of monoglycerides and FA
25
TG
chylomicron
PL
AA    apoprotein B-48
(apoprotein A-I)
The portal vein
lymph
vessels
FA short
chain
glycerol
CHE
Transport of lipids from cells of the
intestinal mucosa
lipophilic vitamins
Biochemie-8-1-lipidy 26
Blood plasma
transport of triacylglycerols in the form of
lipoproteins
fatty acids bound to albumin.
Biochemie-8-1-lipidy 27
Lipoproteins are the transport form of
non-polar lipids in blood
More specifically, we will focus
on lipoproteins, which we can
say that they are a "transport
form" otherwise non-polar lipids
in the blood.
Lipoprotein is composed
of a core and cover. At
the core we find
transmitted lipids (TAG,
cholesterol esters), the
packaging is made of
phospholipids,
cholesterol, and various
proteins (integral and
peripheral).
Lipoprotein size for most types
does not exceed the size of the
colloidal particles (ie the 500
mm), only one type
(chylomicrons) has a diameter
greater than 500 microns. http://1.bp.blogspot.com/_fS2Wvz7uG
Bo/SxyYjpj0ApI/AAAAAAAAALI/fy
of6TlCCnY/s400/lipoprotein+structure
Biochemie-8-1-lipidy 28
NOVÁK, Jan. Biochemie I. Brno: Muni,
2009, Metabolismus lipidů s. 3
Biochemie-8-1-lipidy 29
Types of lipoproteins
VLDL
LDL
Chylomikron
CM
HDL
TG
Proteiny
CH
PL
Density of lipoprotein is determined by its composition. If we focus on the 4 basic structural
components of lipoprotein (TAG - triacylgylcetroly CH - cholesterol, PL - phospholipids, P - proteins),
the percentage of these components can be expressed by the following graphs.
NOVÁK, Jan. Biochemie
I. Brno: Muni, 2009,
Metabolismus lipidů s. 4
Biochemie-8-1-lipidy 30
Metabolism of triglycerides
CH2 O
CH2 O
CHOC
C
C R
O
O
O R
R
1. Hydrolytic cleavage of fatty acids
2. Metabolism of fatty acids and glycerol
The most commonly accepted dietary triglycerides, which are esters of glycerol and fatty acids. Their
metabolism begins by hydrolysis to glycerol and FA. FA and glycerol then go through a completely
different metabolic pathways.
Decomposition of TAG into glycerol and FA by enzymes called lipases (enzymes from the group of
hydrolases) which cleave the ester bond between the glycerol and the chain of FA.
Biochemie-8-1-lipidy 31
CH2 O
CH2 O
CHOC
C
C R
O
O
O R
R
Lipases catalyze the hydrolysis of
triacylglycerols
Cleave the ester bond between the glycerol and FA
Biochemie-8-1-lipidy 32
Lipases
Extracellular Intracellular
pancreatic lipase
(small intestine)
• lipoprotein lipase
(blood)
• hepatic lipase
(surface of
sinusoid)
• hormone-sensitive
(adipocytes - fat cells)
• acidic lipase (lysosomes)
Biochemie-8-1-lipidy 33
(+ colipase)
• operates in the small intestine, splits fats
ingested
• triacylglycerol  2-monoacylglycerol + 2 FA
CH2O-CO-R
R-CO-OCH
CH2O-CO-R
Pancreatic lipase
Biochemie-8-1-lipidy 34
Effect of pancreatic lipase
CH2
CH
O C
O
CH2
OC
O
O
C
O
2 H2O
CH2
CH
OH
CH2 OH
OC
O
HOOC2
Biochemie-8-1-lipidy 35
Lipoprotein lipase
• acts on chylomicrons and VLDL in
blood
cleaves triglycerides contained therein
triacylglycerol  glycerol + 3 FA
CH2O-CO-R
R-CO-OCH
CH2O-CO-R
Biochemie-8-1-lipidy 36
Adipocytal lipase (hormone
sensitive)
active in adipocytes
• depends on hormone action
(glucagon - starvation, adrenaline,
noradrenaline - stress)
• releases fatty acids into the blood
triglyceride  glycerol + 3 FA
Biochemie-8-1-lipidy 37
Transport of fatty acids in ECT
FA in blood - albumin binding
(1 mmol/l, half-life 2 min)
TAG release of the
ECT (CM, VLDL)
Release of TG in adipocytes,
hormonsensitive action of lipase
(Hormonal regulation)
NOVÁK, Jan. Biochemie I. Brno: Muni,
2009, Metabolismus lipidů s. 6
Biochemie-8-1-lipidy 38
• specific membrane proteins facilitate transportation of FA
in cells
transport in cells using FABP (fatty acid binding protein)
• across the mitochondrial membrane by carnitine
Transport of fatty acids in the cells
Biochemie-8-1-lipidy 39
-Oxidation of fatty acids
• Meaning: energy source
In virtually all cells
• Location: mitochondrial matrix
Progress: stepwise removal of acetyl-CoA
Biochemie-8-1-lipidy 40
For transport of FA into mitochondria
carnitine is needed
short chain FA do not require carnitine
(2-hydroxy-3-carboxypropyl)trimethylammonium
N
CH3
CH3
CH2H3C CH CH2
OH
COOH
*
Biochemie-8-1-lipidy 41
FA is transmitted in the form of acylcarnitine
ester bond
N
CH3
CH3
CH2H3C CH CH2
O
COOH
O
C
Biochemie-8-1-lipidy 42
Sources of carnitine
Protein-CH2CH2CH2CH2NH3 protein-CH2CH2CH2CH2N(CH3)3
Side lysine residue
proteolysis
trimethyllysine
carnitine
SAM
Dietary intake: about 100 mg / day (Animal sources:
meat, milk. It is also found in plant sources.)
Liver, brain, kidneys
transport by blood
Synthesis in an organism
+ +
Biochemie-8-1-lipidy 43
carnitine deficiency
• congenital disorder of carnitine transport
• in certain diseases (especially organs that can
synthesize it)
• large losses (diarrhea, hemodialysis, burns ...)
• inhibition of transport into the cell by some drugs
(doxorubicin, cisplatin, lidocaine)
decreased biosynthesis (malnutrition)
Carnitine supplementation in these disorders is
required.
Biochemie-8-1-lipidy 44
Consequences of lack of carnitine
• Lack of carnitine in liver : hypoketotic hypoglycemia during
starvation
-oxidation is required during fasting for the production of acetyl-CoA for
ketogenesis and ATP production for gluconeogesis
Lack in muscle - muscle weakness and cramps
Biochemie-8-1-lipidy 45
The importance of increased intake of carnitine
especially for athletes leads to numerous disputes.
Although many findings about the function and
dynamics of carnitine in the body suggests beneficial
effects of increased intake of dietary supplement in
particular, excessive physical exertion, no convincing
and reliable evidence for this assumption so far been
filed.
Carnitine from food is poorly absorbed, intestinal
bacteria can metabolize to form trimethylamine.
Carnitine as a dietary supplement?
The administration can be only L-carnitine. Dcarnitine
and racemate are officially banned.
Biochemie-8-1-lipidy 46
Cytoplasm
Activation of FA before binding to carnitine
R C
O
OH
+ HS CoA C
O
S CoA
R
ATP AMP + 2Pi
Loss of energy
equivalent to 2
ATP
Biochemie-8-1-lipidy 47
Formation of acyl-
carnitine
carnitinacyltransferase
CH2
CHCH2
N
OH
COOH(CH3)3
+
+
C
O
S CoA
H3C
CH2
CH
O
C
CH2
N COOH(CH3)3
+
O
CH3
+ CoASH
Takes place in the mitochondria
transmembrane space
Biochemie-8-1-lipidy 48
Transport of fatty acids into mitochondria
Two forms of carnitin-acyltransferase
RCO-S-CoA
CoA-SH
RCO-S-CoA
transmembrane space inner mitochondrial membrane matrix
Karnitin/acylkarnitin
výměník
Cn-OH
RCO-OCn
Cn-OH
RCO-OCn CoA-SH
Biochemie-8-1-lipidy 49
NOVÁK, Jan. Biochemie I. Brno:
Muni, 2009, Metabolismus lipidů s. 7
Biochemie-8-1-lipidy 50
-Oxidation of fatty acids
 the main FA degradation pathway
 acyl-CoA enters the reaction
 carbon (C-3) is oxidized
 general mechanism - the repetition of four
steps:
dehydrogenation  hydratation 
dehydrogenation  cleavage of acetyl-CoA
Biochemie-8-1-lipidy 51
(1) Dehydrogenation of acyl-CoA
trans configuration
Biochemie-8-1-lipidy 52
(2) Hydration of the double bond
-II
R CH2 CH CH C
S
O
CoA
-I -I
a,-nenasycený acyl-CoA
H2O
R CH2 CH CH2 C
S
O
CoAOH
0
-hydroxyacyl-CoA
Hydration is not a redox reaction, one C was reduced,
the second C oxidized, but the sum of carbon oxidation
numbers is the same
Biochemie-8-1-lipidy 53
(3) Dehydrogenation of hydroxyacyl
-II
-II
R CH2 CH CH2 C
S
O
CoAOH
0
-hydroxyacyl-CoA
NAD
+
NADH H
+
R CH2 C CH2 C
S
O
CoAO
-oxoacyl-CoA
II
+
Biochemie-8-1-lipidy 54
(4) Thiolysis of oxoacyl and cleavage of
acetyl-CoA
R CH2 C
O
CH2 C
S
O
CoA
CoAS
H
C
S
O
CoA
H3CCH2 C
S
O
CoA
R
acyl-CoA
(2 C shorter)
acetyl-CoA
CC
thiolase
Biochemie-8-1-lipidy 55
The overall progress
of -oxidation
• 1.dehydrogenation
(FAD)
• 2. hydratation
• 3. dehydrogenation
(NAD+)
• 4. transfer of acyl
to CoASH
acyl-CoA
dehydrogenase
2-enoyl-CoA
hydratase
3-hydroxyacyl-CoA
dehydrogenase
thiolase
http://classes.kumc.edu/som/cellbiology
/processes/atp/images/nr39.jpg
Biochemie-8-1-lipidy 56
Acyl-CoA dehydrogenases
3 main types
for FA with short
medium
long chain
Dehydrogenase deficiency for FA with medium chain
congenital disorder - intolerance to prolonged starvation
associated with hypoglycaemic coma (extended in
northwest Europe - up to 90 % of population)
Biochemie-8-1-lipidy 57
The energy yield of the oxidation
of palmitoyl-CoA (16 C)
Palmitoyl-CoA + 7 FAD + 7 NAD+ + 7 H2O + 7 CoA
8 acetyl-CoA + 7 FADH2 + 7 NADH + 7 H+
8 x 12 ATP = 96 ATP 14 ATP 21 ATP
Overall 131 – 2 = 129 ATP / palmitate
Equivalent
to 2 ATP is
consumed
during
formation of
acyl-CoA
Biochemie-8-1-lipidy 58
Comparison of energy-yield of -oxidation
and glycolysis:
Gain of ATP from glucose (6C)
38 ATP on 1 C of glucose 38/6 = 6,3 ATP
from FA (16 C) 129 ATP
on 1 C of FA 129/16 = 8,1 ATP
From 1 C of FA the average yield is
1.3 times more ATP Why?
Biochemie-8-1-lipidy 59
Oxidation of unsaturated fatty acids
oleic acid: cis 9-C18
cis 7-C16
cis 5-C14
cis 3-C12
trans 2-C12
isomerase
loss of FADH2
the same course with
-oxidation
Biochemie-8-1-lipidy 60
Polyunsaturated FA
linoleic acid: 9,12-C18
2-C8 cis
Other enzymes allow complete oxidation
Biochemie-8-1-lipidy 61
FA with an odd number C provide
propionyl
propionyl-CoA
CO2 + H2O
racemase
D-methylmalonyl-CoA
L-methylmalonyl-CoAsuccinyl-CoA
CH3CH2CO -S-CoA
ATP ADP
biotin
CH
COO-
CO-S-CoA
CH3
C H
COO-
CO-S-CoA
CH3CH2-CH2
COO-
CO-S-CoA
B12
produced also
during
metabolism of
some amino acids
Biochemie-8-1-lipidy 62
 -oxidation of FA is an important source of energy
When does it take place?
If a cell needs energy and
does not have enough glucose
-oxidation takes place in postresorption
phase and in starvation especially in the
muscles, myocardium and liver
Biochemie-8-1-lipidy 63
Lipids in postresorption phase
(glucagon)
liver
Acetyl-CoA
muscle
FA
adipose tissue
FA + glycerol-P
TAG
FA-albumin
Acetyl-CoA
the effect of glucagon
Biochemie-8-1-lipidy 64
NOVÁK, Jan. Biochemie I. Brno: Muni,
2009, Metabolismus lipidů s. 13
Biochemie-8-1-lipidy 65
Lipids in postresorption phase
• In adipose tissue lipolysis occurs
(hormone sensitive lipase)
• FA are transported in the ECF bound to
albumin
• FA are the source of energy for muscles
and myocardium
Biochemie-8-1-lipidy 66
Ketone bodies
Acetocetate, 3-hydroxybutyrate - metabolically
usable
acetone - waste product
produced by the liver
• pass into the blood
• they are processed by extrahepatic tissues
• level increased during fasting, diabetes
• the ratio of glucagon / insulin >>> 1
Biochemie-8-1-lipidy 67
Extrahepatically they are metabolized for energy gains 
Increased FA mobilization from adipose tissue 
transport to the liver
 Increased production of acetyl-CoA by -oxidation
capacity exceeded in the citric acid cycle (lack of
oxalacetate)
 synthesis of ketone bodies
Causes of ketone bodies
Increased production is associated with ketoacidosis 
Biochemie-8-1-lipidy 68
Interrelationship of ketone bodies
H3C C CH3
O
H3C CH CH2 C
O
OH
OH
H3C C CH2
O
C
O
O H
- CO2
- 2H
+ 2H
-hydroxymáselná kyselina acetoctová kyselina aceton
Acid pKA
Acetoacetic
-hydroxybutyric
3,52
4,70
Biochemie-8-1-lipidy 69
Synthesis of ketone bodies
matrix of mitochondria
of liver cells
transport via blood to
the extrahepatic tissues
http://dl1.cuni.cz/pluginfile.php/241144
/mod_page/content/4/ketola%CC%81tk
y-schema.jpg
Biochemie-8-1-lipidy 70
• In blood there is always a trace concentration of
ketone bodies
• their levels rise during fasting or
uncompensated diabetes
• unused acetyl-CoA from degradation of fatty
acids in the liver is used to gain energy in
extrahepatic tissues
Formation of ketone bodies
Biochemie-8-1-lipidy 71
Ketone bodies as an energy source in
extrahepatic tissues
H3C C CH2
O
COOH
sukcinyl-CoA sukcinát
H3C C CH2
O
C
SCoA
O
S
H
CoA
C
SCoA
O
H3C2
CC
Energie
acetoacetát
acetoacetyl-CoA
Biochemie-8-1-lipidy 72
NOVÁK, Jan. Biochemie
I. Brno: Muni, 2009,
Metabolismus lipidů s. 14
Biochemie-8-1-lipidy 73
Causes and utilization of ketone bodies
liver
Acetyl-CoA
ketone body ketone bodies in blood
CNS
CO2
muscle
FA
adipose tissue
FA + glycerol-P
TAG
FA-albumin
Acetyl-CoA
lack of oxaloacetate