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===Interaction of Tol with Colicins===
===Interaction of Tol with Colicins===
<structure load='2ivz' size='300' align='right' caption='Interaction of TolB and Colicin E9' (PDB entry [[2ivz]]  |  SCENE= Tol/Tolbcol/1 />
<structure load='2ivz' size='300' align='right' caption='Interaction of TolB and Colicin E9 [[2ivz]]' (PDB entry [[2ivz]]  |  SCENE= Tol/Tolbcol/1 />
It is interesting to note that the name 'Tol' derives from experiments which show that the ''tol'' mutant is tolerant to colicin action<ref name='Davies'>PMID: 1095546</ref>.
 
The Tol-Pal system is used by group A colicins in order to translocate across the outer membrane, targeting mainly the inner membrane component TolA as well as TolQ and TolR.  The colicins set up a translocon, constituting of the outer membrane receptor, translocator proteins and one or more periplasmic translocator proteins.  The colicins recruit the Tol proteins using a Tol binding antigen, or epitope, which is embedded in the IUTD (Intrinsically Unstructured [[Translocation domain]]) found on the N-terminal (T-) domain of the colicin<ref>PMID: 21060316</ref>.
The Tol-Pal system is used by group A colicins in order to translocate across the outer membrane, targeting mainly the inner membrane component TolA as well as TolQ and TolR.  The colicins set up a translocon, constituting of the outer membrane receptor, translocator proteins and one or more periplasmic translocator proteins.  The colicins recruit the Tol proteins using a Tol binding antigen, or epitope, which is embedded in the IUTD (Intrinsically Unstructured [[Translocation domain]]) found on the N-terminal (T-) domain of the colicin<ref>PMID: 21060316</ref>.


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The structure shows the interaction between TolB and the <scene name='Tol/Tolbcolclose/1'>translocation domain</scene> of [[Colicin E9]].
The structure shows the interaction between TolB and the <scene name='Tol/Tolbcolclose/1'>translocation domain</scene> of [[Colicin E9]].


The Tol system can be exploited by the following group A colicins<ref name="Cascales">PMID: 17347522</ref><ref>PMID: 1095546</ref>:
The Tol system can be exploited by the following group A colicins<ref name="Cascales">PMID: 17347522</ref><ref name='Davies'>PMID: 1095546</ref>:
*[[Colicin A]]
*[[Colicin A]]
*[[Colicin E1]]
*[[Colicin E1]]
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==Regulation==
==Regulation==
It has been shown that the regulation of Tol genes is linked to the cell envelope stability<ref name="Cascales">PMID: 17347522</ref>.  In studies with E. coli, the ''tol-pal'' genes have been induced by RcsC in response to cell envelope stress<ref>PMID: 8821933</ref>.  RcsC is a transmembrane sensor kinase, and along with the cytoplasmic response regulator RcsB, makes up the RcsBC regulatory system<ref>PMID: 16166540</ref>.  RcsBC is involved in the regulation of ''cps'' genes which code for the biosynthsis machinery of a major component for the capsula, cholanic acid.
It has been shown that the regulation of Tol genes is linked to the cell envelope stability<ref name="Cascales">PMID: 17347522</ref>.  In studies with E. coli, the ''tol-pal'' genes have been induced by RcsC in response to cell envelope stress<ref>PMID: 8821933</ref>.  RcsC is a transmembrane sensor kinase, and along with the cytoplasmic response regulator RcsB, makes up the RcsBC regulatory system<ref>PMID: 16166540</ref>.  RcsBC is involved in the regulation of ''cps'' genes which code for the biosynthsis machinery of a major component for the capsula, cholanic acid.
==3D structures of Tol proteins==
[[TolA]], [[TolB]], [[TolR]]


== References==
== References==
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