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In-Depth Information
Network
Shorthand
y 1
y 2
y 1
y 2
Genes:
f 2
f 1
Promoters
x 1
x 2
x 1
x 2
Products:
Fig. 7.2: Example 1, simple network with two genes. With feedforward and feed-
back connections included ( left ). Equivalent configuration redrawn with bidirec-
tional connections ( right ).
simple example, Example 1 (Fig. 7.2), if genes y 1 and y 2 lead to product x 1 , then by
definition the x 1 promoter f 1 affects genes y 1 and y 2 . Similarly if gene y 2 leads to
product x 2 , gene y 2 is additionally regulated by product x 2 . Thus gene y 1 is regulated
by product x 1 and gene y 2 is regulated by products x 1 and x 2 . Due to the feedback
model, the activation of gene y 1 can depend on the level of product x 2 , because
if products x 1 and x 2 are consumed equally, then gene y 2 will be promoted at the
expense of gene y 1 .
The network is evaluated until it settles onto a steady state. The solutions are pre-
sented as ( products consumed )
( genes expressed ). Since there are two products
and two genes in Example 1, the solution is written in the form
(
x 1
,
x 2
) (
y 1
,
y 2
)
.
The steady-state solution for Example 1 is
(
x 1
,
x 2
) (
y 1
=
x 1
x 2
,
y 2
=
x 2
)
.The
mathematical equations and their derivation follow:
y 1 (
t
)
x 1
y 2 (
t
)
x 1
x 2
y 2 (
y 1 (
t
+
dt
)=
) ,
y 2 (
t
+
dt
)=
) +
.
(7.5)
y 1 (
t
)+
y 2 (
t
2
y 1 (
t
)+
y 2 (
t
t
)
The network solution at steady state is derived by setting y 1 (
t
+
dt
)=
y 1 (
t
)
and
y 2 (
t
+
dt
)=
y 2 (
t
)
and solving these equations. The solutions are y 1 =
x 1
x 2 and
x 1 + x 2 .
This solution demonstrates efficient outcomes where minimal products are wasted.
Neither x 1 nor x 2 is produced if they are not needed. For example, when products
x 1 and x 2 are equally consumed
y 2 =
x 2 .If x 1
x 2 then y 1 =
0 and the equation for y 2 becomes y 2 =
(
=
)
then gene y 2 is expressed and gene y 1 is
silenced. This occurs because x 1 and x 2 equally usurp promoter f 1 . From the per-
spective of the genes, gene y 1 has all of its promoters reduced when f 1 is usurped,
while gene y 2 still has an independent promoter f 2 . Gene y 2 expression becomes
preferred and in the process inhibits gene y 1 . The final result is that if product x 2 is
not consumed, gene y 2 is not expressed.
If only product x 1 is consumed
x 1
x 2
then only gene y 1 is expressed avoiding
extraneous products. There are consumption patterns where this configuration is
not efficient. For example, if only product x 2 is consumed
(
x 2 =
0
)
(
x 1 =
0
)
then only y 2 is
expressed but extraneous product x 1 is produced.
 
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