Lipoprotein lipase acts in:

A) hydrolysis of triacylglycerols of plasma lipoproteins to supply fatty acids to various tissues.

B) intestinal uptake of dietary fat.
C) intracellular lipid breakdown of lipoproteins.
D) lipoprotein breakdown to supply needed amino acids.
) none of the above.
!ree fatty acids in the bloodstream are:
A) bound to hemoglobin.
B) carried by the protein serum albumin.
C) freely soluble in the a"ueous phase of the blood.
D) none#istent$ the blood does not contain free fatty acids.
) present at levels that are independent of epinephrine.
%he role of hormone&sensitive triacylglycerol lipase is to:
A) hydroly'e lipids stored in the liver.
B) hydroly'e membrane phospholipids in hormone&producing cells.
C) hydroly'e triacylglycerols stored in adipose tissue.
D) synthesi'e lipids in adipose tissue.
) synthesi'e triacylglycerols in the liver.
%he glycerol produced from the hydrolysis of triacylglycerides enters glycolysis as:
A) glucose.
B) glucose&(&phosphate.
C) dihydro#yacetone phosphate.
D) pyruvate.
) glyceryl CoA.
%ransport of fatty acids from the cytoplasm to the mitochondrial matri# re"uires:
A) A%)* carnitine* and coen'yme A.
B) A%)* carnitine* and pyruvate dehydrogenase.
C) A%)* coen'yme A* and he#okinase.
D) A%)* coen'yme A* and pyruvate dehydrogenase.
) carnitine* coen'yme A* and he#okinase.
!atty acids are activated to acyl&CoAs and the acyl group is further transferred to carnitine because:
A) acyl&carnitines readily cross the mitochondrial inner membrane* but acyl&CoAs do not.
B) acyl&CoAs easily cross the mitochondrial membrane* but the fatty acids themselves will not.
C) carnitine is re"uired to o#idi'e +AD
,
to +AD-.
D) fatty acids cannot be o#idi'ed by !AD unless they are in the acyl&carnitine form.

) +one of the above is true.
Carnitine is:
A) a ./&carbon fatty acid.
B) an essential cofactor for the citric acid cycle.
C) essential for intracellular transport of fatty acids.
D) one of the amino acids commonly found in protein.
) present only in carnivorous animals.
0hich of these is able to cross the inner mitochondrial membrane1
A) Acetyl2CoA
B) !atty acyl2carnitine
C) !atty acyl2CoA
D) 3alonyl2CoA
) +one of the above can cross.
0hat is the correct order of function of the following en'ymes of o#idation1
.. &-ydro#yacyl&CoA dehydrogenase
4. %hiolase
5. noyl&CoA hydratase
6. Acyl&CoA dehydrogenase
A) .* 4* 5* 6
B) 5* .* 6* 4
C) 6* 5* .* 4
D) .* 6* 5* 4
) 6* 4* 5* .
7f the .(&carbon saturated fatty acid palmitate is o#idi'ed completely to carbon dio#ide and water
8via the &o#idation pathway and the citric acid cycle)* and all of the energy&conserving products are
used to drive A%) synthesis in the mitochondrion* the net yield of A%) per molecule of palmitate is:
A) 5.
B) .9.
C) 4/.
D) .9:.
) .*999.
;aturated fatty acids are degraded by the stepwise reactions of o#idation* producing acetyl&CoA.
<nder aerobic conditions* how many A%) molecules would be produced as a conse"uence of
removal of each acetyl&CoA1
A) 4
B) 5
C) 6
D) /
) (

0hich of the following is 8are) true of the o#idation of . mol of palmitate 8a .(&carbon saturated
fatty acid$ .(:9) by the &o#idation pathway* beginning with the free fatty acid in the cytoplasm1
.. Activation of the free fatty acid re"uires the e"uivalent of two A%)s.
4. 7norganic pyrophosphate 8))
i
) is produced.
5. Carnitine functions as an electron acceptor.
6. : mol of !AD-4 are formed.
/. : mol of acetyl&CoA are formed.
(. %here is no direct involvement of +AD
,
.
A) . and / only
B) .* 4* and /
C) .* 4* and (
D) .* 5* and /
) / only
Complete o#idation of . mole of which fatty acid would yield the most A%)1
A) .(&carbon saturated fatty acid
B) .:&carbon mono&unsaturated fatty acid
C) .(&carbon mono&unsaturated fatty acid
D) .(&carbon poly&unsaturated fatty acid
) .6&carbon saturated fatty acid
0hich of the following statements apply 8applies) to the o#idation of fatty acids1
.. %he process takes place in the cytosol of mammalian cells.
4. Carbon atoms are removed from the acyl chain one at a time.
5. Before o#idation* fatty acids must be converted to their CoA derivatives.
6. +AD)
,
is the electron acceptor.
/. %he products of o#idation can directly enter the citric acid cycle for further o#idation.
A) . and 5 only
B) .* 4* and 5
C) .* 4* and /
D) 5 and / only
) 6 only
0hich of the following statements concerning the o#idation of fatty acids is true1
A) About .*499 A%) molecules are ultimately produced per 49&carbon fatty acid o#idi'ed.
B) =ne !AD-4 and two +AD- are produced for each acetyl&CoA.
C) %he free fatty acid must be carbo#ylated in the position by a biotin&dependent reaction before
the process of o#idation commences.
D) %he free fatty acid must be converted to a thioester before the process of o#idation
commences.
) %wo +AD- are produced for each acetyl&CoA.
%he balanced e"uation for the degradation of C-58C-4).9C==- via the &o#idation pathway is:

A) C-58C-4).9C==- , /!AD , /+AD
,
, (CoA>;- , /-4= , A%)
( Acetyl&CoA , /!AD-4 , /+AD- , /-
,
, A3) , ))i

B) C-58C-4).9C==- , /!AD , /+AD
,
, (CoA>;- , /-4=
( Acetyl&CoA , /!AD-4 , /+AD- , /-
,

C) C-58C-4).9C==- , (!AD , (+AD
,
, (CoA>;- , (-4= , A%)
( Acetyl&CoA , (!AD-4 , (+AD- , (-
,
, A3) , ))i

D) C-58C-4).9C==- , (!AD , (+AD
,
, (CoA>;- , (-4=
( Acetyl&CoA , (!AD-4 , (+AD- , (-
,

0hich compound is an intermediate of the o#idation of fatty acids1
A) C-5>8C-4)49>C=>C==-
B) C-5>C-4>C=>C-4>C=>=)=5
42

C) C-5>C-4>C=>C-4>=-
D) C-5>C-4>C=>C=>;>CoA
) C-5>C=>C-4>C=>;>CoA
%he conversion of palmitoyl&CoA 8.(:9) to myristoyl&CoA 8.6:9) and . mol of acetyl&CoA by the &
o#idation pathway results in the net formation of:
A) . !AD-4 and . +AD-.
B) . !AD-4 and . +AD)-.
C) . !AD-4* . +AD-* and . A%).
D) 4 !AD-4 and 4 +AD-.
) 4 !AD-4* 4 +AD-* and . A%).
0hich of the following is not true regarding the o#idation of . mol of palmitate 8.(:9) by the &
o#idation pathway1
A) . mol of A%) is needed.
B) : mol of acetyl&CoA are formed.
C) : mol of !AD-4 are formed.
D) A3) and ))
i
are formed.
) %he reactions occur in the mitochondria.
7f an aerobic organism 8e.g.* the bacterium E. coli) were fed each of the following four compounds as
a source of energy* the energy yield per mole from these molecules would be in the order:
A) alanine ? glucose ? palmitate 8.(:9)
B) glucose ? alanine ? palmitate
C) glucose ? palmitate ? alanine
D) palmitate ? alanine ? glucose
) palmitate ? glucose ? alanine
0hich of the following is 8are) true of the o#idation of long&chain fatty acids1
.. %he en'yme comple# that cataly'es the reaction contains biotin.
4. !AD-4 serves as an electron carrier.
5. +AD- serves as an electron carrier.

6. =#idation of an .:&carbon fatty acid produces si# molecules of propionyl&CoA.
/. =#idation of a ./&carbon fatty acid produces at least one propionyl&CoA.
A) .* 4* and 5
B) .* 4* and /
C) 4* 5* and 6
D) 4* 5* and /
) 5 and / only
%he following fatty acid* in which the indicated carbon is labeled with
.6
C* is fed to an animal:
.6
C-58C-4)@C==-
After allowing 59 minutes for fatty acid o#idation* the label would most likely be recovered in:
A) acetyl&CoA.
B) beta&hydro#y butyryl&CoA.
C) both acetyl&CoA and propionyl&CoA.
D) palmitoyl&CoA.
) propionyl&CoA.
%he carbon atoms from a fatty acid with an odd number of carbons will enter the citric acid cycle as
acetyl&CoA and:
A) butyrate.
B) citrate.
C) malate.
D) succinyl&CoA.
) &ketoglutarate.
7n the disease sprue* vitamin B.4 8cobalamin) is poorly absorbed in the intestine* resulting in B.4
deficiency. 7f each of the following fatty acids were in the diet* for which one would the process of
fatty acid o#idation be most affected in a patient with sprue1
A) C-58C-4).9C==-
B) C-58C-4)..C==-
C) C-58C-4).4C==-
D) C-58C-4).6C==-
) C-58C-4).:C==-
0hich en'yme is the maAor regulatory control point for &o#idation1
A) pyruvate carbo#ylase
B) carnitine acyl transferase 7
C) acetyl CoA dehydrogenase
D) enoyl CoA isomerase
) methylmalonyl CoA mutase
During o#idation of fatty acids* BBBBBBBBBBB is produced in pero#isomes but not in mitochondria.
A) acetyl&CoA
B) !AD-4


C) -4=
D) -4=4

) +AD-
0hen comparing the &o#idation and &o#idation pathways* which one of the following statements
is correct1
A) &o#idation and &o#idation occur in the cytoplasm.
B) o#idation occurs at the carbo#yl end of the fatty acid whereas o#idation occurs at the methyl
end.
C) o#idation occurs at the methyl end of the fatty acid whereas o#idation occurs at the carbo#yl
end.
D) o#idation occurs mainly in the cytoplasm whereas o#idation occurs mainly in the
mitochondria.
) o#idation occurs mainly in the mitochondria whereas o#idation occurs mainly in the
cytoplasm.
Cetone bodies are formed in the liver and transported to the e#trahepatic tissues mainly as:
A) acetoacetyl&CoA.
B) acetone.
C) beta&hydro#ybutyric acid.
D) beta&hydro#ybutyryl&CoA.
) lactic acid.
%he maAor site of formation of acetoacetate from fatty acids is the:
A) adipose tissue.
B) intestinal mucosa.
C) kidney.
D) liver.
) muscle.
0hy is it more efficient to store energy as lipid rather than as glycogen1
!irst* the energy yield per gram of lipid 8about 5: kDEg) is more than twice that for carbohydrate
8about .F kDEg). ;econd* lipid is stored as anhydrous lipid droplets* but carbohydrates such as
glycogen and starch are stored hydrated* and the water of hydration roughly triples the effective
weight of the carbohydrate* reducing the energy yield to about ( kDEg.
7n the first step of fatty acid o#idation* the fatty acid 8G>C==-) is converted to its coen'yme A
derivative in the following reaction:
G2C==- , A%) , CoA2;- G2C=2;2CoA , A3) , ))i
%he standard free&energy change 8G') for this reaction is 2./ kDEmol

0hat will tend to make the reaction more favorable when it takes place within a cell1
%he hydrolysis of ))
i
by inorganic pyrophosphatase* for which G' is 2.@ kDEmol* makes the
overall G' more negative.
%he o#idation of acetyl&CoA added to isolated* intact mitochondria is stimulated strongly by
carnitine. 0hy1
Carnitine is essential in the transport of fatty acyl groups into the mitochondrial matri#* where fatty
acid o#idation occurs.
%he o#idation of fatty acids begins with this activation reaction:
G2C-42C-42C-42C==- , A%) , CoA2;-
G2C-42C-42C-42C=2;2CoA , A3) , ))i
0hat are the ne#t two steps 8after transport into the mitochondria)1 ;how structures and indicate
where any cofactors participate.
%he reactions are those cataly'ed by fatty acyl2CoA dehydrogenase and enoyl hydratase. ;ee !ig.
.F&:a* p. (/5.
Draw the four basic steps in the o#idation of a saturated fatty acid 8the &o#idation pathway). ;how
structures* name en'ymes* and indicate where any cofactors participate.
;ee !ig. .F&:a* p. (/5.
;how the last step in the se"uence of the four reactions in the &o#idation pathway for fatty acid
degradation. 7nclude the structures of reactant and product* the en'yme name* and indicate where any
cofactors participate.
;ee the thiolase reaction* !ig. .F&:a* p. (/5.
=ne of the steps in fatty acid o#idation in mitochondria involves the addition of water across a
double bond. 0hat is the ne#t step in the process1 ;how structures and indicate where any
cofactor8s) participate8s).
%he reaction is that cataly'ed by &hydro#yacyl&CoA dehydrogenase* for which +AD
,
is cofactor.
;ee !ig. .F&:a* p. (/5.
7n the citric acid cycle* a double bond is introduced into a four&carbon compound containing the >
C-
4
>C-
4
> group* producing fumarate. ;how a similar reaction that occurs in the &o#idation
pathway.
;ee !ig. .F&:a* p. (/5.
0rite a balanced e"uation for the o#idation of palmitoyl&CoA* a .(&carbon* fully saturated fatty
acid* and indicate how much of each product is formed.

%he overall reaction is:
)almitoyl&CoA , FCoA&;- , F!AD , F+AD
,
, F-
4
=
: acetyl&CoA , F!AD-
4
, F+AD- , F-
,
!or each two&carbon increase in the length of a saturated fatty acid chain* how many additional
moles of A%) can be formed upon complete o#idation of one mole of the fatty acid to C=4 and -4=1
ach >C-4>C-4> unit yields .6 e#tra A%) molecules. %he two o#idations of the &o#idation
pathway produce . !AD-4 and . +AD-* which yield ../ and 4./ A%)* respectively* by o#idative
phosphorylation. %he e#tra acetyl&CoA* when o#idi'ed via the citric acid cycle* yields another .9
A%) e"uivalents: 5 +AD-* . !AD-4* and . A%) or H%).
0rite a balanced e"uation for the complete o#idation 8to acetyl&CoA and any other products that
might be formed) of pelargonic acid* C-58C-4)FC==-.
%he odd&chain fatty acid is first activated to the CoA derivative* then o#idi'ed to 5 acetyl&CoA and .
propionyl&CoA by o#idation. %he propionyl&CoA is converted to succinyl&CoA through the
se"uence of reactions shown on p. ((9* !ig. .F&... %he overall reaction is therefore:
)elargonic acid , -C=5
2
, A%) , 6CoA;- , 5!AD , 5+AD
,

5 acetyl&CoA , succinyl&CoA , 5!AD-4 , 5+AD- , A3) , ))
i
8a) Describe the steps in the metabolic pathway in which cells o#idi'e a four&carbon* straight&chain*
saturated fatty acid 8butyrate$ 6:9) to the fragments that enter the citric acid cycle. ;how the
structures of intermediates and products* and indicate where any cofactor8s) participate8s). 8b) 7n
what way would you change or add to your answer if the starting fatty acid had been five carbons
long 8also straight&chain and saturated)1
8a) Butyrate is first activated:
Butyrate , A%) , CoA>;- butyryl&CoA , A3) , ))
i
%hen* the butyryl group is transferred to carnitine and transported into the mitochondrial matri#*
where it is reconverted to the butyryl&CoA derivative. %his passes through the four steps of
o#idation. 8;ee !ig. .F&:a* p. (/5.) 8b) A five&carbon chain would undergo activation and one cycle
of o#idation* producing acetyl&CoA and propionyl&CoA. )ropionyl&CoA would be converted to
succinyl&CoA by the reaction se"uence in !ig. .F&..* p. ((9.
An e#perimenter studying the o#idation of fatty acids in e#tracts of liver found that when palmitate
8.(:9) was provided as substrate* it was completely o#idi'ed to C=4. -owever* when undecanoic
acid 8..:9) was added as substrate* incomplete o#idation occurred unless he bubbled C=4 through the
reaction mi#ture. %he addition of the protein avidin* which binds tightly to biotin* prevented the
complete o#idation of undecanoic acid even in the presence of C=4* although it had no effect on
palmitate o#idation. #plain these observations in light of what you know of fatty acid o#idation
reactions.
=#idation of odd&chain fatty acid yields acetyl&CoA , propionyl&CoA. %he reaction C=4 ,
propionyl&CoA methylmalonyl&CoA is cataly'ed by propionyl&CoA carbo#ylase* a biotin&
containing en'yme* which is therefore inhibited by avidin.

%wo vitamins* biotin and vitamin B.4* play crucial roles in the metabolism of propionic acid
8propionate). #plain this by showing the steps in which each is essential in propionate metabolism.
Biotin and vitamin B.4 act as cofactors for propionyl&CoA carbo#ylase and methylmalonyl&CoA
mutase* respectively$ see !ig. .F&..* p. ((9* for the complete se"uence of reactions.
%he total degradation of a fatty acid with an odd number of carbons yields acetyl&CoA and another
compound* I. ;how the structure of I* and describe the pathway by which it is converted into a
citric acid cycle intermediate* including where any cofactors participate.
I is propionyl&CoA* and its conversion into succinyl&CoA is accomplished by the reactions in !ig.
.F&..* p. ((9.
;how the shortest pathway by which propionyl&CoA can be converted into a citric acid cycle
intermediate. 7ndicate where any cofactors participate.
;ee !ig. .F&..* p. ((9.
7f you received a laboratory report showing the presence of a high concentration of ketone bodies in
the urine of a patient* what disease would you suspect1 0hy do ketone bodies accumulate in such
patients1
%he patient is probably an untreated diabetic* but the condition might also result from fasting. 7n
either case* the unavailability of glucose from the blood stimulates gluconeogenesis in the liver. As
the substrate for glucose formation* o#aloacetate is withdrawn from the citric acid cycle* bringing that
cycle to a near halt. %he fatty acids being o#idi'ed in the liver yield acetyl&CoA* which now cannot
be o#idi'ed via the citric acid cycle. Geversal of the thiolase reaction produces acetoacetyl&CoA*
which is then converted into ketone bodies and e#ported from the liver. ;ee !ig. .F&.:* p. (((.

Draw the structure of one ketone body* and describe circumstances under which you would e#pect to
find high concentrations of this compound in the urine of a human.
%he ketone bodies* acetoacetate* &hydro#ybutyrate* and acetone 8p. ((()* are overproduced in
untreated diabetes mellitus and during prolonged fasting* when fatty acids become the principle
energy source.
0hat are ketone bodies and why do they form during fasting1
%he ketone bodies* acetoacetate* &hydro#ybutyrate* and acetone* are overproduced during fasting*
when fatty acids from stored triacylglycerols become the principle o#idi'able fuel. Accumulation of
acetyl&CoA and its precursor acetoacetyl&CoA favors ketone body formation. Because o#aloacetate is
used for gluconeogenesis* it is withdrawn from the citric acid cycle* bringing that cycle to a near halt.
%he acetyl&CoA that is produced by o#idation can no longer be o#idi'ed via the citric acid cycle so
it accumulates.