Two Component Signaling in bacteria

In this system two components are involved. The first

component is a sensory kinase which has a kinase
activity. It senses the change in the surrounding and
phosphorylates the second component. The second
component is a response regulator which acts according
to the signal received from sensory kinase. This two
component signaling is found in bacteria and eukaryotes.
It has been extensively studied in bacterial systems.
Bacteria use it in transformation competence,
pathogenicity, flagellar motility, cell division, membrane
transport, antibiotic resistance and other metabolisms.
The reactions which occur in the system are as follows:
ATP HK ADP HK P
HK P RR RR P HK
RR P RR Pi
Sensory kinase which is normally a histidine kinase
(HK) upon change in the surrounding
autophosphorylates itself on histidine moiety. This
phosphate is passed on to aspartate of response regulator
(RR). Response regulator loses its phosphate with or
without enzymatic intervention.

Systems in bacteria
PHOSPHORUS SYSTEM
Bacteria starved for Phosphorus




Bacteria having abundant Phosphorus

In this system PhoR and PhoB are two components.
PhoR is a sensory kinase and PhoB is a response
regulator. When cell is starved ion Phosphorus PhoR
phosphorylates PhoB showing its kinase activity. The
activates PhoB* combines with RNA polymerase and
tanscribes the gene for Phosphorus synthesis (PhoA).
PhoR acts as a Phosphatase when Phosphorus is
abundant. It dephophorylates PhoB*, which later does
not combine with RNA polymerase so no transcription
of PhoA gene for Phosphorus synthesis occurs.
PORINS

In this system the sensory kinase is EnvZ an osmolarity
protein sensor) and response regulator is OmpR
(regulator for EnvZ). OmpC and OmpF are porins.


When the solute concentration is low, Omp F is
predominant and has a larger pore size than that of Omp
C. When the solute concentration is high, Omp C is
predominant.
As osmotic pressure increases, Env-Z gets
phosphorylated. The phosphorylated Env-Z (Env-Z P)
will pass on its phosphate to Omp R. Higher the
regulation, higher the concentration oI Omp R P.
Under low osmotic pressure, the concentration oI
OmpR P is not very high. The OmpR P would bind to
the high affinity site upstream of Omp F.
This high aIIinity site when bound to it OmpR P
enhance the transcription of Omp F. so omp F porins
will be predominant at low osmotic pressure.
As the osmolarity goes on increasing, Omp R P
increases, it will bind upstream of Omp F which will
downregulate transcription of Omp F.
Additionally, it will also bind at a site upstream of Omp
C, so it will upregulate transcription of Omp C.

CARBON / NITROGEN METABOLISM




P
11
is a phosphatase action of NR
11
which is a sensory
kinase. NR
1
is a response regulator. NR
11

autophosphorylates NR
1
.
NR
1
~ P will bind upstream to those genes that are
responsible for the assimilation of ammonia. The first
gene is Glutamine synthase [gln A]. the state of UMP
with P
11
is sensed. P
11
can be found in two forms: one is
unmodified P
11
and other is modified with UMP. If P
11
is
unmodified then NR
11
has to phosphatase activity and
NR
1
~ P becomes NR
1
.
If the amount of Glutamine in the body is sufficiently
high then P
11
~UMP will become P
11
. UMP Transferase
(UT) and UMP Remover (UR) plays a catalytic role in
this conversion. UR acts when concentration of Gln is
high |N| whereas UT will act when concentration oI
Gln is low |N|. When Gln is high NR
11
cannot
phosphorylate NR
1
which will then bind upstream and
start transcription and assimilation.


CHEMOTAXIS


In this system Che R is a methyl transferase , Che B is a
methyl esterase, Che A is a sensory kinase and Che Y is
a response regulator. Che Y is superimposed on the other
two regulators i.e Che B and Che R.
This is an example of regain of chemotaxis with two
component signaling. Various receptors related to this
system are Tar for arginine, Tsr for serine and Tap for
small peptides. In the absence of nutrients or in the
presence of toxins Che A will get autophosphorylated to
Che A~P. when it gets autophosphorylated it passes its
phosphate to Che Y. Che Y~P will go and bind to the
flagella and cause tumbling.
CheB also gets phosphorylated by Che A. CheB is a
methyl esterase and CheB~P is its active form. It will
remove methyl moiety from CheA which is glutamate
moiety. When methyl group is removed from CheA, it
attenuates i.e. reduces the activity of , Che A.
Che R adds methyl groups to these glutamate moieties. It
sensitizes CheA which is a histidine kinase. It again
phosphorylates and starts phosphorylating Che Y which
will lead to tumbling and also demethylation.