a .

I r r e v e r s i b i l i t y,

b. Presence of an exergonic, first

committed step,

c. Strict regulations at the first

committed step, and,

d. Occurrence in specific sub cellular

compartments in eukaryotic cells.
The reaction pathways involved in
biosyntheses are called anabolic
pathways.

The process also takes place in

three stages.
A D P + H P O 42-
NADP+

ATP
NADPH
 An adipocyte (fat cell)

Structure of a TAG
Triacylglycerol Diacylglycerol Monoacylglycerol
Triacyl glycerol

3 H2 O
even number

odd number
 On the outer mitochondrial membrane.

 Catalysed by a family of at least three

acyl-CoA synthetases (fatty acid thiokinases).

RCOOH + CoA-SH + ATP RCo-SCoA + AMP + PPi

fatty acid fatty-acyl CoA
(ΔG°ʹ= -15 kJ/mol)
 The free energy change associated with the
overall reaction is close to zero.

 Such a reaction (ΔG°ʹ= -15 kJ/mol) is driven

to completion (ΔG°ʹ= -34kJ/mol) in the cell
by the highly exergonic hydrolysis
(ΔG°ʹ= -19 kJ/mol) of the pyrophosphate
(PPi) to inorganic orthophosphate (Pi) in
a reaction catalysed by inorganic pyrophosphatase.
RCOO - + CoA-SH + ATP + H2O RCO-SCoA + AMP + Pi + 2H+
fatty acid fatty-acyl CoA

(ΔG°ʹ= -34kJ/mol)
 Fatty acids are activated in the
outer mitochondrial membrane, but are
oxidised in the mitochondrial matrix.

 A long chain fatty acyl-CoA cannot directly

cross the inner mitochondrial membrane as
CoA-thioesters.
4 reactions are involved:

a . F o r m a t i o n o f t r a n s - Δ 2- e n o y l C o A .

Acyl CoA +E-FAD t r a n s - Δ 2- e n o y l C o A + E - F A D H 2

 b) Hydration to form a L-ß-hydroxy-acyl
CoA or L-3- hydroxy-acyl CoA.

t r a n s - Δ 2- e n o y l C o A + H 2O L-ß-hydroxy-acyl CoA

c) NAD+-requiring dehydrogenation to form

the corresponding ß-keto-acyl CoA.

L-ß-hydroxy-acyl CoA + NAD+ ß-keto-acyl CoA

+ NADH + H+
d) Acetyl CoA (derived from C-1 and C-2 of the original fatty acid
chain) and a new, long-chain, saturated fatty acyl-CoA, shortened
by 2 C-atoms with respect to the original one are now formed.

(n carbons) (n-2 carbons)

 oxidation

hydration

oxidation
thiolysis
 Overall equation for one turn of the cycle:

Cn-acyl CoA + CoA + FAD + NAD+ + H2O Cn-2-acyl CoA + acetyl-CoA +

FADH2 + NADH + H+

For example, starting from palmitoyl-CoA:

Palmitoyl-CoA + CoA + FAD + NAD+ + H2O myristoyl-CoA + acetyl-CoA

+ FADH2 + NADH + H+
Each round of ß-oxidation produces one NADH,
one FADH2 and one acetyl CoA. Oxidation of
acetyl CoA via the TCA cycle generates
additional FADH2 and NADH, which are
reoxidised through oxidative phosphorylation
to form ATP.