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Fat Metabolism
Shahneaz Ali Khan
By complete oxidation of 1 gm of Fat produce 9.3 calories energy
whereas carbohydrate only 4 cal/gm
Digestion of dietary lipids in vertebrates
Oxidation of Fat
Fat
Hydrolysis
Fatty acid Glycerol
Glycerol Kinase
Glycerol-3-Phosphate
Dihydroxy Acetone Phosphate
Gluconeogenic pathway
Pyruvate
TCA
Oxidation of Fatty Acid
Requires different stges:
1. Activation of fatty acid in cytoplasm
2. Transfer of Acyl CoA from cytosol to
mitochondria (Carnitine Shuttle system)
3. Beta Oxidation in mitochondria
 Dehydrogenation
 Hydration
 2nd dehydrogenation
 Thyolytic cleavage
1. Activation of fatty acid
2. Transfer of Acyl CoA from
cytosol to mitochondria
(Carnitine Shuttle system)
3. Beta Oxidation in mitochondria
Dehydrogen
ation
Hydration
2nd
Dehydrogen
ation
Thyolytic
Cleavage/
Acylation
Fatty Acid oxidation
• Major Pathway
– β-oxidation
• Minor Pathway
– α-oxidation
(branch-chain FA,e.g. Phytanic acid)
– ω-oxidation
β-oxidation Pathway• Oxidation of fatty acids takes place in
mitochondria where the various enzymes for
fatty acid oxidation are present close to the
enzymes of the electron transport chain.
• Fatty acid oxidation is a major source of cell
ATP
• Oxidation of FAs occur at the β-carbon atom
resulting in the elimination of the two terminal
carbon atoms as acetyl CoA leaving fatty acyl
CoA which has two carbon atoms less than
the original fatty acid.
• β-oxidation has 4 steps:
1-Dehydrogenation (FAD-dependent)
2- Hydration
3-Dehydrogenation (NAD-dependent)
4-Cleavage (Remove 2C as acetyl CoA)
Calculations
Carbons in Fatty
Acid
Acetyl CoA
C/2
β-oxidation cycles
(C/2) -1
12 6 5
14 7 6
16 8 7
18 9 8
Note: In each round of β-oxidation one molecule of FADH2 and
one molecule of NADH+H+ are produced which generates 2 and
3 ATP molecules, respectively
Example: Energy of palmitoyl ~Co A
(16 C) oxidation
• Number of cycles= n/2 -1 = 7 cycles
• Number of acetyl ~Co A = n/2 =8
 So, 7 NADH, each provide 3 ATP when oxidized in the ETC
7X3=21 ATP
 7 FADH2 each provide 2 ATP when oxidized in the ETC
7x 2=14 ATP
 8 acetyl ~Co A , each provides 12 ATP when converted to
CO2& H2O by the TCA cycle 8x12= 96 ATP
So total energy yield of oxidation of palmitoyl ~Co A = 21 +
14 + 96 = 131 ATP
• As 2 molecules of ATP are used in the activation of a
molecule of fatty acid Therefore, there is a net yield of
129 molecules of ATP
Regulation of fatty acid β-oxidation
1- The level of ATP in the cell :If it is high in the cell, the rate of β-
oxidation will decrease (Feed back inhibition)
2- Malonyl-CoA
* (which is also a precursor for fatty acid synthesis) inhibits Carnitine
Palmitoyl Transferase I and thus, inhibits β-oxidation
* Malonyl-CoA is produced from acetyl-CoA by the enzyme Acetyl-CoA
Carboxylase
Oxidation of Unsaturated Fatty
Acid
• Slightly more complicated Requires additional enzymes
• Oxidation of unsaturated FAs produce less energy than that
of saturated FAs (because they are less highly reduced,
therefore, fewer reducing equivalents can be produced from
these structures)
Oxidation of Odd Numbered Fatty
Acid
• Requires three additional extra
reactions.
• Odd numbered lipids are present in
plants and marine organisms
• Fatty acids with odd number of
carbon atoms are also oxidized by
the same process β-oxidation as
even chain FAs, removing 2
carbons as acetyl CoA in each
round of the oxidative process BUT
the final round of β-oxidation of a
fatty acid with an odd number of C
atoms yields acetyl-CoA &
propionyl-CoA (3C).
α-Oxidation Pathway
• α-oxidation occurs in brain tissue in
order to oxidize short chain FAs
• Inα-oxidation,there is one carbon
atom removed at time from α
position
• It does not require CoA and does
not generate high- energy
phosphates
• This type of oxidation is significant
in the metabolism of dietary FAs
that are methylated on β-carbon
e.g. phytanic acid (peroxisomes)
ω-Oxidation Pathway
• ω-oxidation is a minor
pathway and occurs in the
endoplasmic reticulum of
many tissues rather than
the mitochondria, the site
of β-oxidation.
• This process occurs
primarily with medium
chain FAs of adipose
tissue which are mobilized
to the liver under
conditions of ketosis
Energy Production
Acetyl CoA will enter into TCA cycle and will give energy
The entry of acetyl CoA into the citric acid cycle
depends on the availability of oxaloacetate.
The concentration of oxaloacetate is lowered if
carbohydrate is unavailable (starvation) or improperly
utilized (diabetes).
Oxaloacetate is
normally formed from
pyruvate by pyruvate
carboxylase
(anaplerotic reaction).
Fats burn in the flame
of carbohydrates.
KETONE BODIES
In fasting or diabetes the gluconeogenesis is activated
and oxaloacetate is consumed in this pathway.
Fatty acids are oxidized producing excess of acetyl CoA
which is converted to ketone bodies:
b-Hydroxybutyrate
Acetoacetate
Acetone
Ketone bodies are fuel
molecules (can fuel brain and
other cells during starvation)
Ketone bodies are synthesized
in liver mitochondria and
exported to different organs.
A. Synthesis of ketone bodies
Two molecules
of acetyl CoA
condense to
form
acetoacetyl CoA.
Enzyme –
thiolase.
Acetoacetyl
CoA reacts
with acetyl
CoA and water
to give 3-
hydroxy-3-
methylglutaryl
CoA (HMG-
CoA) and CoA.
Enzyme:
HMG-CoA
synthase
3-Hydroxy-3-
methylglutaryl
CoA is then
cleaved to
acetyl CoA and
acetoacetate.
Enzyme:
HMG-CoA lyase.
3-Hydroxybutyrate is
formed by the reduction of
acetoacetate by
3-hydroxybutyrate
dehydrogenase.
Acetoacetate also
undergoes a slow,
spontaneous
decarboxylation to
acetone.
The odor of acetone may
be detected in the breath
of a person who has a high
level of acetoacetate in
the blood.
B. Ketone bodies are a major fuel
in some tissues
Ketone bodies diffuse from the liver
mitochondria into the blood and are transported
to peripheral tissues.
Ketone bodies are important molecules in energy
metabolism.
Heart muscle and the renal cortex use
acetoacetate in preference to glucose in
physiological conditions.
The brain adapts to the utilization of
acetoacetate during starvation and diabetes.
3-Hydroxybutyrate is oxidized to produce
acetoacetate as well as NADH for use in
oxidative phosphorylation.
3-hydroxybutyrate
dehydrogenase
Acetoacetate is activated
by the transfer of CoA
from succinyl CoA in a
reaction catalyzed by a
specific CoA transferase.
Acetoacetyl CoA is cleaved
by thiolase to yield two
molecules of acetyl CoA
(enter the citric acid
cycle).
CoA transferase is present
in all tissues except liver.
Ketone bodies are a water-
soluble, transportable
form of acetyl units
Impairment of the tissue function, most importantly in the central
nervous system
KETOSIS
The absence of insulin in diabetes mellitus
 liver cannot absorb glucose
 inhibition of glycolysis
 activation of gluconeogenesis
 deficit of oxaloacetate
 activation of fatty acid
mobilization by adipose tissue
 large amounts of acetyl CoA which can not be
utilized in Krebs cycle
 large amounts of ketone bodies (moderately strong acids)
 severe acidosis (ketosis)
Pathways for Pyruvate
• The pyruvate produced from glucose during
glycolysis can be further metabolized in three
possible ways
• For aerobic organisms, when oxygen is plentiful the
pyruvate is converted to acetyl coenzyme A (acetyl
CoA)
• For aerobic organisms, when oxygen is scarce, and
for some anaerobic organisms, the pyruvate is
reduced to lactate
• For some anaerobic organisms (like yeast), the
pyruvate is fermented to ethanol
Conversion of Pyruvate to Acetyl CoA
• Under aerobic conditions, pyruvate from glycolysis is
decarboxylated to produce acetyl CoA, which enters
the citric acid cycle as well as other metabolic
pathways
- the enzyme involved is pyruvate dehydrogenase
and the coenzyme NAD+ is also required
• This pathway provides the most energy from glucose
O
||
CH3—C—COO- + HS—CoA + NAD+ 
pyruvate
O
||
CH3—C—S—CoA + CO2 + NADH + H+
acetyl CoA
Conversion of Pyruvate to Lactate
• For aerobic organisms under anaerobic conditions,
pyruvate is reduced to lactate, which replenishes NAD+
to continue glycolysis
• During strenuous exercise, muscle cells quickly use up
their stored oxygen, creating anaerobic conditions
- lactate accumulates, leading to muscle fatigue and
soreness
• Anaerobic bacteria can also produce lactate, which is
how we make pickles and yogurt (among other things)
O lactate
|| dehydrogenase
CH3—C—COO- + NADH + H+ 
pyruvate
OH
|
CH3—CH—COO- + NAD+
lactate
Conversion of Pyruvate to Ethanol
• Anaerobic microorganisms such as yeast, convert
pyruvate to ethanol by fermentation
- pyruvate is decarboxylated to acetaldehyde, which is
reduced to ethanol
- NAD+ is regenerated to continue glycolysis
• The CO2 produced during fermentation make the
bubbles in beer and champagne, and also makes bread
rise
• Alcoholic beverages produced by fermentation can be
up to around 15% ethanol
- above that concentration the yeast die
O
O
O
pyruvate
decarboxylase
H+
CO2
H
O
OH
alcohol
dehydrogenase
NADH + H+
NAD+
Pyruvate Acetaldehyde Ethanol
Overview of Pyruvate Pathways
Cori Cycle
• When anaerobic conditions occur in active muscle,
glycolysis produces lactate
• The lactate moves through the blood stream to the
liver, where it is oxidized back to pyruvate.
• Gluconeogenesis converts pyruvate to glucose,
which is carried back to the muscles
• The Cori cycle is the flow of lactate and glucose
between the muscles and the liver
Pathways for Glucose
Michaelis-Menten equation
• Michaelis-Menten equation is the rate equation for an enzyme –
catalyzed reaction and is the mathematical description of the
hyperbolic curve we have discussed earlier. The formula is
][
][max
0
SK
SV
V
m 

Where,
V0 is the initial velocity
Vmax is the maximum velocity
[S] is the substrate concentration
Km (Michaelis-Menten constant) is the substrate concentration
at which the reaction velocity is the half of the maximum velocity.
Graphical determination of KM
Thank you!!!

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Fat metabolism

  • 2. By complete oxidation of 1 gm of Fat produce 9.3 calories energy whereas carbohydrate only 4 cal/gm
  • 3. Digestion of dietary lipids in vertebrates
  • 4.
  • 5.
  • 6. Oxidation of Fat Fat Hydrolysis Fatty acid Glycerol Glycerol Kinase Glycerol-3-Phosphate Dihydroxy Acetone Phosphate Gluconeogenic pathway Pyruvate TCA
  • 7. Oxidation of Fatty Acid Requires different stges: 1. Activation of fatty acid in cytoplasm 2. Transfer of Acyl CoA from cytosol to mitochondria (Carnitine Shuttle system) 3. Beta Oxidation in mitochondria  Dehydrogenation  Hydration  2nd dehydrogenation  Thyolytic cleavage
  • 8. 1. Activation of fatty acid
  • 9. 2. Transfer of Acyl CoA from cytosol to mitochondria (Carnitine Shuttle system)
  • 10. 3. Beta Oxidation in mitochondria Dehydrogen ation Hydration 2nd Dehydrogen ation Thyolytic Cleavage/ Acylation
  • 11. Fatty Acid oxidation • Major Pathway – β-oxidation • Minor Pathway – α-oxidation (branch-chain FA,e.g. Phytanic acid) – ω-oxidation
  • 12. β-oxidation Pathway• Oxidation of fatty acids takes place in mitochondria where the various enzymes for fatty acid oxidation are present close to the enzymes of the electron transport chain. • Fatty acid oxidation is a major source of cell ATP • Oxidation of FAs occur at the β-carbon atom resulting in the elimination of the two terminal carbon atoms as acetyl CoA leaving fatty acyl CoA which has two carbon atoms less than the original fatty acid. • β-oxidation has 4 steps: 1-Dehydrogenation (FAD-dependent) 2- Hydration 3-Dehydrogenation (NAD-dependent) 4-Cleavage (Remove 2C as acetyl CoA)
  • 13. Calculations Carbons in Fatty Acid Acetyl CoA C/2 β-oxidation cycles (C/2) -1 12 6 5 14 7 6 16 8 7 18 9 8 Note: In each round of β-oxidation one molecule of FADH2 and one molecule of NADH+H+ are produced which generates 2 and 3 ATP molecules, respectively
  • 14. Example: Energy of palmitoyl ~Co A (16 C) oxidation • Number of cycles= n/2 -1 = 7 cycles • Number of acetyl ~Co A = n/2 =8  So, 7 NADH, each provide 3 ATP when oxidized in the ETC 7X3=21 ATP  7 FADH2 each provide 2 ATP when oxidized in the ETC 7x 2=14 ATP  8 acetyl ~Co A , each provides 12 ATP when converted to CO2& H2O by the TCA cycle 8x12= 96 ATP So total energy yield of oxidation of palmitoyl ~Co A = 21 + 14 + 96 = 131 ATP • As 2 molecules of ATP are used in the activation of a molecule of fatty acid Therefore, there is a net yield of 129 molecules of ATP
  • 15. Regulation of fatty acid β-oxidation 1- The level of ATP in the cell :If it is high in the cell, the rate of β- oxidation will decrease (Feed back inhibition) 2- Malonyl-CoA * (which is also a precursor for fatty acid synthesis) inhibits Carnitine Palmitoyl Transferase I and thus, inhibits β-oxidation * Malonyl-CoA is produced from acetyl-CoA by the enzyme Acetyl-CoA Carboxylase
  • 16. Oxidation of Unsaturated Fatty Acid • Slightly more complicated Requires additional enzymes • Oxidation of unsaturated FAs produce less energy than that of saturated FAs (because they are less highly reduced, therefore, fewer reducing equivalents can be produced from these structures)
  • 17. Oxidation of Odd Numbered Fatty Acid • Requires three additional extra reactions. • Odd numbered lipids are present in plants and marine organisms • Fatty acids with odd number of carbon atoms are also oxidized by the same process β-oxidation as even chain FAs, removing 2 carbons as acetyl CoA in each round of the oxidative process BUT the final round of β-oxidation of a fatty acid with an odd number of C atoms yields acetyl-CoA & propionyl-CoA (3C).
  • 18. α-Oxidation Pathway • α-oxidation occurs in brain tissue in order to oxidize short chain FAs • Inα-oxidation,there is one carbon atom removed at time from α position • It does not require CoA and does not generate high- energy phosphates • This type of oxidation is significant in the metabolism of dietary FAs that are methylated on β-carbon e.g. phytanic acid (peroxisomes)
  • 19. ω-Oxidation Pathway • ω-oxidation is a minor pathway and occurs in the endoplasmic reticulum of many tissues rather than the mitochondria, the site of β-oxidation. • This process occurs primarily with medium chain FAs of adipose tissue which are mobilized to the liver under conditions of ketosis
  • 20. Energy Production Acetyl CoA will enter into TCA cycle and will give energy
  • 21. The entry of acetyl CoA into the citric acid cycle depends on the availability of oxaloacetate. The concentration of oxaloacetate is lowered if carbohydrate is unavailable (starvation) or improperly utilized (diabetes). Oxaloacetate is normally formed from pyruvate by pyruvate carboxylase (anaplerotic reaction). Fats burn in the flame of carbohydrates. KETONE BODIES
  • 22. In fasting or diabetes the gluconeogenesis is activated and oxaloacetate is consumed in this pathway. Fatty acids are oxidized producing excess of acetyl CoA which is converted to ketone bodies: b-Hydroxybutyrate Acetoacetate Acetone Ketone bodies are fuel molecules (can fuel brain and other cells during starvation) Ketone bodies are synthesized in liver mitochondria and exported to different organs.
  • 23. A. Synthesis of ketone bodies Two molecules of acetyl CoA condense to form acetoacetyl CoA. Enzyme – thiolase.
  • 24. Acetoacetyl CoA reacts with acetyl CoA and water to give 3- hydroxy-3- methylglutaryl CoA (HMG- CoA) and CoA. Enzyme: HMG-CoA synthase
  • 25. 3-Hydroxy-3- methylglutaryl CoA is then cleaved to acetyl CoA and acetoacetate. Enzyme: HMG-CoA lyase.
  • 26. 3-Hydroxybutyrate is formed by the reduction of acetoacetate by 3-hydroxybutyrate dehydrogenase. Acetoacetate also undergoes a slow, spontaneous decarboxylation to acetone. The odor of acetone may be detected in the breath of a person who has a high level of acetoacetate in the blood.
  • 27. B. Ketone bodies are a major fuel in some tissues Ketone bodies diffuse from the liver mitochondria into the blood and are transported to peripheral tissues. Ketone bodies are important molecules in energy metabolism. Heart muscle and the renal cortex use acetoacetate in preference to glucose in physiological conditions. The brain adapts to the utilization of acetoacetate during starvation and diabetes.
  • 28. 3-Hydroxybutyrate is oxidized to produce acetoacetate as well as NADH for use in oxidative phosphorylation. 3-hydroxybutyrate dehydrogenase
  • 29. Acetoacetate is activated by the transfer of CoA from succinyl CoA in a reaction catalyzed by a specific CoA transferase. Acetoacetyl CoA is cleaved by thiolase to yield two molecules of acetyl CoA (enter the citric acid cycle). CoA transferase is present in all tissues except liver. Ketone bodies are a water- soluble, transportable form of acetyl units
  • 30. Impairment of the tissue function, most importantly in the central nervous system KETOSIS The absence of insulin in diabetes mellitus  liver cannot absorb glucose  inhibition of glycolysis  activation of gluconeogenesis  deficit of oxaloacetate  activation of fatty acid mobilization by adipose tissue  large amounts of acetyl CoA which can not be utilized in Krebs cycle  large amounts of ketone bodies (moderately strong acids)  severe acidosis (ketosis)
  • 31. Pathways for Pyruvate • The pyruvate produced from glucose during glycolysis can be further metabolized in three possible ways • For aerobic organisms, when oxygen is plentiful the pyruvate is converted to acetyl coenzyme A (acetyl CoA) • For aerobic organisms, when oxygen is scarce, and for some anaerobic organisms, the pyruvate is reduced to lactate • For some anaerobic organisms (like yeast), the pyruvate is fermented to ethanol
  • 32. Conversion of Pyruvate to Acetyl CoA • Under aerobic conditions, pyruvate from glycolysis is decarboxylated to produce acetyl CoA, which enters the citric acid cycle as well as other metabolic pathways - the enzyme involved is pyruvate dehydrogenase and the coenzyme NAD+ is also required • This pathway provides the most energy from glucose O || CH3—C—COO- + HS—CoA + NAD+  pyruvate O || CH3—C—S—CoA + CO2 + NADH + H+ acetyl CoA
  • 33. Conversion of Pyruvate to Lactate • For aerobic organisms under anaerobic conditions, pyruvate is reduced to lactate, which replenishes NAD+ to continue glycolysis • During strenuous exercise, muscle cells quickly use up their stored oxygen, creating anaerobic conditions - lactate accumulates, leading to muscle fatigue and soreness • Anaerobic bacteria can also produce lactate, which is how we make pickles and yogurt (among other things) O lactate || dehydrogenase CH3—C—COO- + NADH + H+  pyruvate OH | CH3—CH—COO- + NAD+ lactate
  • 34. Conversion of Pyruvate to Ethanol • Anaerobic microorganisms such as yeast, convert pyruvate to ethanol by fermentation - pyruvate is decarboxylated to acetaldehyde, which is reduced to ethanol - NAD+ is regenerated to continue glycolysis • The CO2 produced during fermentation make the bubbles in beer and champagne, and also makes bread rise • Alcoholic beverages produced by fermentation can be up to around 15% ethanol - above that concentration the yeast die O O O pyruvate decarboxylase H+ CO2 H O OH alcohol dehydrogenase NADH + H+ NAD+ Pyruvate Acetaldehyde Ethanol
  • 36. Cori Cycle • When anaerobic conditions occur in active muscle, glycolysis produces lactate • The lactate moves through the blood stream to the liver, where it is oxidized back to pyruvate. • Gluconeogenesis converts pyruvate to glucose, which is carried back to the muscles • The Cori cycle is the flow of lactate and glucose between the muscles and the liver
  • 38. Michaelis-Menten equation • Michaelis-Menten equation is the rate equation for an enzyme – catalyzed reaction and is the mathematical description of the hyperbolic curve we have discussed earlier. The formula is ][ ][max 0 SK SV V m   Where, V0 is the initial velocity Vmax is the maximum velocity [S] is the substrate concentration Km (Michaelis-Menten constant) is the substrate concentration at which the reaction velocity is the half of the maximum velocity.
  • 40.
  • 41.