Team:Imperial College London/M2/genes

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Through the course of evolution, <i>E.coli</i> have equipped themselves with a multitude of defences to enable colonisation of the intestine. We are using two global transcription factors (RcsB & YgiV) to hijack this natural process in a way that maximises acid resitance. We have additionally upregulated a third enzyme (rfal) to enhance the encapsulation of single cells (over and above colony encapsulation). Finally, the two biosynthetic genes (OtsA & OtsB) code for enzymes responsible for the production of trehalose. Our manipulation of endogenous pathways reduces virulence while enchancing pill functionality.  
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===<b>R</b>egulation for <b>C</b>apsular <b>S</b>ynthesis (RCS) Phosphorelay System===
 +
 
 +
RcsB is a transcription factor that forms part of a phosphorelay system. As the name suggests, a phosphorelay system consists of a series of nodes down which a phosphate group is passed. The transfer of this phosphate group down along the Rcs relay system is initiated by membrane stress. Once the phosphate group has been passed to RcsB, it can bind to DNA and act as a transcription factor. Phosphorylated RcsB is thought to regulate ~5% of the E.coli genome <sup>4</sup>.
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[[Image:PhosRelay.png|left|400px]]
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The Rcs phosphorelay is composed of two membrane-anchored proteins, RcsC and RcsD, and a cytoplasmic protein, RcsB. In response to a signal, the sensor kinase RcsC autophosphorylates and the phosphoryl group is transferred to RcsB, via RcsD. RcsB can both homodimerise and heterodimerise with RcsA, these two complexes serve to regulate different operons.
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===RcsB===
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===<b>Signal Amplification:</b>===
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<b>Background:</b>
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RcsB is able to heterodimerise with the protein RcsA. The RcsB+RcsA complex acts as a transcription factor that induces the expression of a number of genes. One of these genes codes for RcsA, this initiates a positive feedback loop.
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RcsB is a transcription factor that forms part of the phosphorelay system. In response to membrane stress, RcsB is phosphorylated into its DNA binding form. In this state, it is able to both upregulate and downregulate a large number of genes.
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<html>
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<p class="ecoparagraph">
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<map name="MAP91791">
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<area coords="147,18,177,35" onmouseover="return overlib('H-NS transcriptional dual regulator represses transcription from associated promoter<br>The exact location of this binding site is unknown or not specified.<br><b>Evidence:</b> Gene expression analysis<br>The nature of binding of this protein is not well known.');" onmouseout="return nd();">
 +
<area coords="0,74,32,91" onmouseover="return overlib('RcsAB transcriptional dual regulator activates transcription from associated promoter<br><b>Distance to transcription start site:</b> -166.5 (ranging from -173 to -160)<br><b>Evidence:</b> Gene expression analysis [Ebel99], Automated inference based on similarity to consensus sequences [Ebel99]');" onmouseout="return nd();">
 +
<area coords="34,74,66,91" onmouseover="return overlib('RcsAB transcriptional dual regulator activates transcription from associated promoter<br><b>Distance to transcription start site:</b> -125.5 (ranging from -132 to -119)<br><b>Evidence:</b> Binding of cellular extracts, Site mutation [Wehland00], Binding of purified proteins [Wehland00]');" onmouseout="return nd();">
 +
<area coords="147,0,184,17" onmouseover="return overlib('GadE transcriptional activator activates transcription from associated promoter<br>The exact location of this binding site is unknown or not specified.<br><b>Evidence:</b> Binding of purified proteins [Hommais04], Gene expression analysis [Hommais04]');" onmouseout="return nd();">
 +
<area coords="104,60,165,91" onmouseover="return overlib(' <b>Gene:</b>  rcsA    EG10820    b1951    ECK1949<br><b>Synonyms:</b> cpsR<br><b>Location:</b> 2,021,992 -> 2,022,615  <br> <b>Product:</b> positive DNA-binding transcriptional regulator of capsular polysaccharide synthesis, activates its own expression, subunit of RcsAB transcriptional dual regulator');" onmouseout="return nd();">
 +
<area coords="24,0,184,108" onmouseover="return overlib('<b>Operon:</b> rcsAp (Experim. ev.)  <BR><b>Tr.Activators:</b> (GadE RcsAB) <BR><b>Tr.Inhibitors:</b> (Hns)');" onmouseout="return nd();">
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<area coords="0,0,184,108"  onmouseover="return overlib('<b>Operon</b>: rcsA');" onmouseout="return nd();">
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</map>
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<img src="https://static.igem.org/mediawiki/2009/8/89/Op8.gif" usemap="#MAP91791" border="0">
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</p>
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</html>
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{{Imperial/09/Division}}
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===<b>Colanic Acid Biosynthesis:</b>===
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The overexpression of RcsB is akin to fitting a stethoscope to the end of a megaphone. In essence, the sensitivity of the RCS Phosphorelay system becomes such that the smallest of membrane perturbations becomes sufficient to trigger colanic acid production. Thus, the background noise switches the RCS Phosphorelay system on permanently.
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There are a total of 19 genes involved in colonic acid biosynthesis, the best characterised of which are shown below.
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<html>
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</p><p class="ecoparagraph">
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<map name="MAP79413">
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<area coords="0,55,32,72" onmouseover="return overlib('RcsAB transcriptional dual regulator activates transcription from associated promoter<br><b>Distance to transcription start site:</b> -105.5 (ranging from -99 to -112)<br>
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 +
<b>Evidence:</b> Binding of purified proteins [Wehland00], Site mutation [Wehland00]');" onmouseout="return nd();">
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<area coords="32,55,64,72" onmouseover="return overlib('RcsAB transcriptional dual regulator activates transcription from associated promoter<br><b>Distance to transcription start site:</b> -78.5 (ranging from -72 to -85)<br>
 +
 
 +
<b>Evidence:</b> Gene expression analysis [Ebel99], Automated inference based on similarity to consensus sequences [Ebel99]');" onmouseout="return nd();">
 +
<area coords="274,41,338,72" onmouseover="return overlib(' <b>Gene:</b>  wzb    G7106    b2061    ECK2055<br><b>Location:</b> 2,133,679 <- 2,134,122  <br> <b>Product:</b> tyrosine phosphatase');" onmouseout="return nd();">
 +
<area coords="745,41,816,72" onmouseover="return overlib(' <b>Gene:</b>  wcaB    G7103    b2058    ECK2052<br><b>Location:</b> 2,130,091 <- 2,130,579  <br> <b>Product:</b> predicted colanic acid biosynthesis acyl transferase');" onmouseout="return nd();">
 +
<area coords="631,41,745,72"  onmouseover="return overlib(' <b>Gene:</b>  wcaA    G7104    b2059    ECK2053<br><b>Location:</b> 2,130,582 <- 2,131,421  <br> <b>Product:</b> predicted colanic acid biosynthesis glycosyl transferase');" onmouseout="return nd();">
 +
<area coords="122,41,274,72" onmouseover="return overlib(' <b>Gene:</b>  wza    G7107    b2062    ECK2056<br><b>Location:</b> 2,134,128 <- 2,135,267  <br> <b>Product:</b> lipoprotein required for capsular polysaccharide translocation through the outer membrane, subunit of capsular polysaccharide export apparatus');" onmouseout="return nd();">
 +
<area coords="338,41,619,72"  onmouseover="return overlib(' <b>Gene:</b>  wzc    G7105    b2060    ECK2054<br><b>Location:</b> 2,131,514 <- 2,133,676  <br> <b>Product:</b> tyrosine kinase involved in colanic acid biosynthesis, subunit of capsular polysaccharide export apparatus');" onmouseout="return nd();">
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<area coords="16,0,135,89"  onmouseover="return overlib('<b>Operon:</b> wzap (Experim. ev.)  <BR><b>Tr.Activators:</b> (RcsAB)');" onmouseout="return nd();">
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<area coords="0,0,818,89" onmouseover="return overlib('<b>Operon</b>: wza-wzb-wzc-wcaAB<br><b>Synonyms:</b> OP00290, cps, cpsB, cpsGB');" onmouseout="return nd();">
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</map>
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<img src="https://static.igem.org/mediawiki/2009/8/89/Op1.gif" usemap="#MAP79413" border="0">
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</html>
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<html>
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</p><p class="ecoparagraph">
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<map name="MAP91795">
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<area coords="41,55,72,72" onmouseover="return overlib('IHF transcriptional dual regulator activates transcription from associated promoter<br><b>Distance to transcription start site:</b> -71 (ranging from -77 to -65)<br><b>Evidence:</b> Binding of purified proteins [Murtin98], Human inference based on similarity to consensus sequences [Murtin98]');" onmouseout="return nd();">
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<area coords="0,55,32,72" onmouseover="return overlib('RcsAB transcriptional dual regulator activates transcription from associated promoter<br><b>Distance to transcription start site:</b> -147.5 (ranging from -154 to -141)<br><b>Evidence:</b> Gene expression analysis [Ferrieres07], Human inference based on similarity to consensus sequences [Ferrieres07]');" onmouseout="return nd();">
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<area coords="98,41,137,72" onmouseover="return overlib(' <b>Gene:</b>  yjbE    EG11923    b4026    ECK4018<br><b>Location:</b> 4,233,929 -> 4,234,171  <br> <b>Product:</b> predicted protein');" onmouseout="return nd();">
 +
<area coords="154,41,247,72" onmouseover="return overlib(' <b>Gene:</b>  yjbF    EG11924    b4027    ECK4019<br><b>Location:</b> 4,234,285 -> 4,234,923  <br> <b>Product:</b> predicted lipoprotein');" onmouseout="return nd();">
 +
<area coords="246,41,353,72" onmouseover="return overlib(' <b>Gene:</b>  yjbG    EG11925    b4028    ECK4020<br><b>Location:</b> 4,234,920 -> 4,235,657  <br> <b>Product:</b> conserved protein');" onmouseout="return nd();">
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<area coords="352,41,640,72"  onmouseover="return overlib(' <b>Gene:</b>  yjbH    EG11926    b4029    ECK4021<br><b>Location:</b> 4,235,657 -> 4,237,753  <br> <b>Product:</b> predicted porin');" onmouseout="return nd();">
 +
<area coords="22,0,143,89" onmouseover="return overlib('<b>Operon:</b> yjbEp (No experim. ev.)  <BR><b>Tr.Activators:</b> (RcsAB IHF)');" onmouseout="return nd();">
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<area coords="0,0,642,89" onmouseover="return overlib('<b>Operon</b>: yjbEFGH');" onmouseout="return nd();">
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</map>
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<img src="https://static.igem.org/mediawiki/2009/2/28/Op11.gif" usemap="#MAP91795" border="0">
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</html>
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==Other genes involved in colanic acid biosynthesis are listed below.==
 +
 
 +
=====<b><i>wcaF</i> </b>=====
 +
 
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*Putative colanic acid biosynthesis acetyltransferase <sup>2</sup>
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=====<b><i>gmd</i> </b>=====
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Codes for GDP-mannose 4,6-dehydratase which is a subunit of GDP-mannose 4,6-dehydratase. This enzyme is involved in colanic acid biosynthesis.
 +
 
 +
=====<b><i>wcaD</i> </b>=====
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* Predicted colanic acid polymerase
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=====<b><i>fcl</i> </b>=====
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* Codes for a subunit of GDP-fucose synthase. This enzyme is involved in colanic acid biosynthesis.
 +
 
 +
 
 +
{{Imperial/09/Division}}
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===<b>General Protective:</b>===
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The following proteins are thought play a role in membrane fortification to confer protection against the rigours of dessication, acid exposure and osmotic stress.
 +
 
 +
<html>
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 +
<p class="ecoparagraph">
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<map name="MAP91783">
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<area coords="17,55,48,72" onmouseover="return overlib('RcsB-Pasp trancriptional activator activates transcription from associated promoter<br><b>Distance to transcription start site:</b> -41.5 (ranging from -35 to -48)<br><b>Evidence:</b> Gene expression analysis [Boulanger05], Human inference based on similarity to consensus sequences [Boulanger05]');" onmouseout="return nd();">
 +
<area coords="101,41,162,72" onmouseover="return overlib(' <b>Gene:</b>  osmB    EG10679    b1283    ECK1278<br><b>Location:</b> 1,341,134 <- 1,341,352  <br> <b>Product:</b> OsmB osmotically inducible lipoprotein');" onmouseout="return nd();">
 +
<area coords="0,0,115,89" onmouseover="return overlib('<b>Operon:</b> osmBp1 (Experim. ev.)  <BR><b>Tr.Activators:</b> (phospho-RcsB)');" onmouseout="return nd();">
 +
<area coords="0,0,164,89" onmouseover="return overlib('<b>Operon</b>: osmB');" onmouseout="return nd();">
 +
</map>
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<img src="https://static.igem.org/mediawiki/2009/9/9f/Op5.gif" usemap="#MAP91783" border="0">
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</p>
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</html>
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<html>
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<p class="ecoparagraph">
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<map name="MAP91779">
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<area coords="142,0,172,17" onmouseover="return overlib('H-NS transcriptional dual regulator represses transcription from associated promoter<br>The exact location of this binding site is unknown or not specified.<br><b>Evidence:</b> Gene expression analysis [Bouvier98]<br>The nature of binding of this protein is not well known.');" onmouseout="return nd();">
 +
<area coords="1,56,32,73" onmouseover="return overlib('Lrp transcriptional dual regulator represses transcription from associated promoter<br><b>Distance to transcription start site:</b> -52 (ranging from -58 to -46)<br><b>Evidence:</b> Gene expression analysis, Binding of purified proteins, Human inference based on similarity to consensus sequences, Site mutation');" onmouseout="return nd();">
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<area coords="25,20,61,37"  onmouseover="return overlib('RcsB-Pasp trancriptional activator activates transcription from associated promoter<br><b>Distance to transcription start site:</b> -41.5 (ranging from -48 to -35)<br><b>Evidence:</b> Site mutation [DavalosGar01], Gene expression analysis [DavalosGar01], Binding of purified proteins [FrancezCha05]');" onmouseout="return nd();">
 +
<area coords="19,38,70,55"  onmouseover="return overlib('NhaR-Na+ transcriptional activator activates transcription from associated promoter<br><b>Distance to transcription start site:</b> -41 (ranging from -50 to -32)<br><b>Evidence:</b> Gene expression analysis [Toesca01], Binding of cellular extracts [Sturny03], Site mutation [Toesca01]');" onmouseout="return nd();">
 +
<area coords="87,42,177,73" onmouseover="return overlib(' <b>Gene:</b>  osmC    EG10680    b1482    ECK1476<br><b>Location:</b> 1,554,649 -> 1,555,080  <br> <b>Product:</b> osmotically inducible peroxidase OsmC');" onmouseout="return nd();">
 +
<area coords="19,0,172,90" onmouseover="return overlib('<b>Operon:</b> osmCp1 (Experim. ev.)  <BR><b>Tr.Activators:</b> (NhaR-Na<SUP>+</SUP> phospho-RcsB) <BR><b>Tr.Inhibitors:</b> (Lrp Hns)');" onmouseout="return nd();">
 +
<area coords="0,0,179,90" onmouseover="return overlib('<b>Operon</b>: osmC');" onmouseout="return nd();">
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</map>
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<img src="https://static.igem.org/mediawiki/2009/1/10/Op3.gif" usemap="#MAP91779" border="0">
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</p>
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 +
</html>
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 +
==Other RcsB induced genes coding for protective proteins are listed below.==
 +
 
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=====<b><i>KatE</i> </b>=====
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* Codes for a subunit of heme d synthase / hydroperoxidase. This bacterial catalase protects against osmotic stress.
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 +
{{Imperial/09/Division}}
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===<b>Host Defence:</b>===
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 +
Humans synthesise a number of enzymes that are able to burst bacterial cells. These enzymes are called lysosymes. RcsB is able to induce a number of proteins that act as lysosyme inhibitors that will prevent the host-mediated lysis of our chassis.
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====RcsB upregulates the following genes:====
 
=====<b><i>ivy</i> (Inhibitor of Vertebrate lysozyme) </b>=====
=====<b><i>ivy</i> (Inhibitor of Vertebrate lysozyme) </b>=====
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*Discovered in 2001 as the first bacterial lysozyme inhibitor. This Type-C lysozyme inhibitor resides in the periplasm.<cite>1</cite>
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*Discovered in 2001 as the first bacterial lysozyme inhibitor. This Type-C lysozyme inhibitor resides in the periplasm.<sup>1</sup>
=====<b><i>MilC</i> (Membrane-bound lysozyme inhibitor of Type C lysozyme) </b>=====
=====<b><i>MilC</i> (Membrane-bound lysozyme inhibitor of Type C lysozyme) </b>=====
Line 21: Line 165:
*This is a lipoprotein that resides in the membrane. <sup>1</sup>  
*This is a lipoprotein that resides in the membrane. <sup>1</sup>  
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<b>References</b>
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*[http://www.ncbi.nlm.nih.gov/pubmed/19136591 <b>The Rcs two-component system regulates expression of lysozyme inhibitors and is induced by exposure to lysozyme</b>]
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{{Imperial/09/Division}}
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===<b>Global Regulators</b>:===
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<html>
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 +
<script type="text/javascript" src="http://biocyc.org/overLIB/Mini/overlib_mini.js"><!-- overLIB (c) Erik Bosrup --></script>
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<script type="text/javascript" src="http://biocyc.org/overLIB/Mini/overlib_exclusive_mini.js"></script>
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<script type="text/javascript" src="http://biocyc.org/overLIB/Mini/overlib_positioncap_mini.js"></script>
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<p class="ecoparagraph">
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<map name="MAP80844">
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<area coords="18,55,49,72" onmouseover="return overlib('RcsB-phosphate activates transcription from associated promoter<br><b>Distance to transcription start site:</b> -42.5 (ranging from -47 to -38)<br><b>Evidence:</b> Gene expression analysis [Majdalani02], Site mutation [Majdalani02], Human inference based on similarity to consensus sequences [Majdalani02]');" onmouseout="return nd();">
 +
<area coords="59,41,91,72" onmouseover="return overlib(' <b>Gene:</b>  rprA    G0-8863    b4431    ECK1686<br><b>Synonyms:</b> psrA5, IS083<br><b>Location:</b> 1,768,396 -> 1,768,500  <br> <b>Product:</b> RprA small RNA, regulator of RpoS');" onmouseout="return nd();">
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<area coords="0,0,105,89" onmouseover="return overlib('<b>Operon</b>: rprA');" onmouseout="return nd();">
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<area coords="0,0,105,89" onmouseover="return overlib('<b>Operon:</b> rprAp (Experim. ev.)  <BR><b>Tr.Activators:</b> (RcsB-phosphate)');" onmouseout="return nd();">
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</map>
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<img src="https://static.igem.org/mediawiki/2009/1/1c/Op2.gif" usemap="#MAP80844" border="0">
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</p>
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</html>
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<html>
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<p class="ecoparagraph">
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<map name="MAP91781">
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<area coords="17,55,48,72" onmouseover="return overlib('RcsB-Pasp trancriptional activator activates transcription from associated promoter<br><b>Distance to transcription start site:</b> -41.5 (ranging from -35 to -48)<br><b>Evidence:</b> Binding of purified proteins [FrancezCha05], Gene expression analysis [FrancezCha05]');" onmouseout="return nd();">
 +
<area coords="165,41,207,72"  onmouseover="return overlib(' <b>Gene:</b>  sra    EG11508    b1480    ECK1474<br><b>Synonyms:</b> rpsV<br><b>Location:</b> 1,553,850 <- 1,553,987  <br> <b>Product:</b> 30S ribosomal subunit protein S22, subunit of ribosome. Sra is more abundant at stationary phase than during log phase growth.  ');" onmouseout="return nd();">
 +
<area coords="75,41,136,72" onmouseover="return overlib(' <b>Gene:</b>  bdm    G6776    b1481    ECK1475<br><b>Synonyms:</b> yddX<br><b>Location:</b> 1,554,089 <- 1,554,304  <br> <b>Product:</b> biofilm-dependent modulation protein');" onmouseout="return nd();">
 +
<area coords="0,0,103,89" onmouseover="return overlib('<b>Operon:</b> bdmp (Experim. ev.)  <BR><b>Tr.Activators:</b> (phospho-RcsB)');" onmouseout="return nd();">
 +
<area coords="0,0,209,89" onmouseover="return overlib('<b>Operon</b>: bdm-sra');" onmouseout="return nd();">
 +
</map>
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<img src="https://static.igem.org/mediawiki/2009/a/ae/Op4.gif" usemap="#MAP91781" border="0">
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</p>
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</html>
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<html>
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<p class="ecoparagraph">
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<map name="MAP91785">
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<area coords="12,55,43,72" onmouseover="return overlib('RcsB-Pasp trancriptional activator activates transcription from associated promoter<br><b>Distance to transcription start site:</b> -45 (ranging from -51 to -39)<br><b>Evidence:</b> Gene expression analysis, Human inference based on similarity to consensus sequences, Site mutation');" onmouseout="return nd();">
 +
<area coords="0,0,99,89" onmouseover="return overlib('<b>Operon:</b> ftsAp1 (Experim. ev.)  <BR><b>Tr.Activators:</b> (phospho-RcsB)');" onmouseout="return nd();">
 +
<area coords="521,41,829,72" onmouseover="return overlib(' <b>Gene:</b>  ftsZ    EG10347    b0095    ECK0096<br><b>Synonyms:</b> sfiB, sulB<br><b>Location:</b> 105,305 -> 106,456  <br> <b>Product:</b> essential cell division protein FtsZ');" onmouseout="return nd();">
 +
<area coords="167,41,505,72" onmouseover="return overlib(' <b>Gene:</b>  ftsA    EG10339    b0094    ECK0095<br><b>Synonyms:</b> divA<br><b>Location:</b> 103,982 -> 105,244  <br> <b>Product:</b> essential cell division protein FtsA');" onmouseout="return nd();">
 +
<area coords="0,0,831,89" onmouseover="return overlib('<b>Operon</b>: ftsAZ');" onmouseout="return nd();">
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</map>
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<img src="https://static.igem.org/mediawiki/2009/e/e8/FtsA.gif" usemap="#MAP91785" border="0">
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</p>
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</html>
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{{Imperial/09/Division}}
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===YgiV===
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==Repression:==
 +
 
 +
 
 +
 
 +
RcsB is also is able to act as a repressor, preventing the expression of both curli and fimbriae. This reduces cell motility and plays a role in biofilm formation.
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<html>
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<p class="ecoparagraph">
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<map name="MAP91787">
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<area coords="1,128,32,145" onmouseover="return overlib('CRP-cAMP transcriptional dual regulator activates transcription from associated promoter<br><b>Distance to transcription start site:</b> -170.5 (ranging from -160 to -181)<br><b>Evidence:</b> Gene expression analysis [Zheng04], Automated inference based on similarity to consensus sequences [Zheng04]');" onmouseout="return nd();">
 +
<area coords="39,128,70,145"  onmouseover="return overlib('CpxR-Phosphorylated represses transcription from associated promoter<br><b>Distance to transcription start site:</b> -100.5 (ranging from -93 to -107)<br><b>Evidence:</b> Binding of purified proteins [Jubelin05], Gene expression analysis [Jubelin05], Human inference based on similarity to consensus sequences [Jubelin05]');" onmouseout="return nd();">
 +
<area coords="135,128,166,145"  onmouseover="return overlib('CpxR-Phosphorylated represses transcription from associated promoter<br><b>Distance to transcription start site:</b> -20.5 (ranging from -13 to -27)<br><b>Evidence:</b> Binding of purified proteins [Jubelin05], Gene expression analysis [Jubelin05], Human inference based on similarity to consensus sequences [Jubelin05]');" onmouseout="return nd();">
 +
<area coords="166,128,197,145"  onmouseover="return overlib('CpxR-Phosphorylated represses transcription from associated promoter<br><b>Distance to transcription start site:</b> 2.5 (ranging from 10 to -5)<br><b>Evidence:</b> Binding of purified proteins [Jubelin05], Gene expression analysis [Jubelin05], Human inference based on similarity to consensus sequences [Jubelin05]');" onmouseout="return nd();">
 +
<area coords="79,74,110,91"  onmouseover="return overlib('RstA-Phosphorylated transcriptional regulator represses transcription from associated promoter<br><b>Distance to transcription start site:</b> -50.5 (ranging from -44 to -57)<br><b>Evidence:</b> Binding of purified proteins [Ogasawara07a], Gene expression analysis [Ogasawara07a], Human inference based on similarity to consensus sequences [Ogasawara07a]');" onmouseout="return nd();">
 +
<area coords="74,128,109,145"  onmouseover="return overlib('CpxR-Phosphorylated represses transcription from associated promoter<br><b>Distance to transcription start site:</b> -52 (ranging from -45 to -59)<br><b>Evidence:</b> Binding of purified proteins [PrigentCom01], Gene expression analysis [PrigentCom01], Human inference based on similarity to consensus sequences [PrigentCom01]');" onmouseout="return nd();">
 +
<area coords="76,110,111,127" onmouseover="return overlib('CpxR-Phosphorylated represses transcription from associated promoter<br><b>Distance to transcription start site:</b> -51 (ranging from -44 to -58)<br><b>Evidence:</b> Binding of purified proteins [Jubelin05], Gene expression analysis [Jubelin05], Human inference based on similarity to consensus sequences [Jubelin05]');" onmouseout="return nd();">
 +
<area coords="81,56,116,73"  onmouseover="return overlib('CpxR-Phosphorylated represses transcription from associated promoter<br><b>Distance to transcription start site:</b> -48.5 (ranging from -41 to -55)<br><b>Evidence:</b> Binding of purified proteins [Jubelin05], Gene expression analysis [Jubelin05], Human inference based on similarity to consensus sequences [Jubelin05]');" onmouseout="return nd();">
 +
<area coords="238,0,276,17" onmouseover="return overlib('CsgD transcriptional dual regulator activates transcription from associated promoter<br>The exact location of this binding site is unknown or not specified.');" onmouseout="return nd();">
 +
<area coords="238,18,280,35"  onmouseover="return overlib('RcsAB transcriptional dual regulator represses transcription from associated promoter<br>The exact location of this binding site is unknown or not specified.<br><b>Evidence:</b> Gene expression analysis [Vianney05]');" onmouseout="return nd();">
 +
<area coords="79,92,123,109" onmouseover="return overlib('OmpR-Phosphorylated transcriptional dual regulator activates transcription from associated promoter<br><b>Distance to transcription start site:</b> -47.5 (ranging from -38 to -57)<br><b>Evidence:</b> Binding of purified proteins [Jubelin05], Gene expression analysis [Jubelin05], Human inference based on similarity to consensus sequences [Jubelin05]');" onmouseout="return nd();">
 +
<area coords="87,38,135,55" onmouseover="return overlib('CRP-cAMP transcriptional dual regulator activates transcription from associated promoter<br><b>Distance to transcription start site:</b> -42.5 (ranging from -32 to -53)<br><b>Evidence:</b> Gene expression analysis [Zheng04], Automated inference based on similarity to consensus sequences [Zheng04]');" onmouseout="return nd();">
 +
<area coords="218,114,279,145" onmouseover="return overlib(' <b>Gene:</b>  csgD    G6546    b1040    ECK1026<br><b>Location:</b> 1,101,769 <- 1,102,419  <br> <b>Product:</b> CsgD transcriptional dual regulator');" onmouseout="return nd();">
 +
<area coords="279,114,340,145" onmouseover="return overlib(' <b>Gene:</b>  csgE    G6545    b1039    ECK1025<br><b>Location:</b> 1,101,375 <- 1,101,764  <br> <b>Product:</b> curli production assembly/transport component, subunit of Curli Secretion and Assembly Complex');" onmouseout="return nd();">
 +
<area coords="343,114,404,145" onmouseover="return overlib(' <b>Gene:</b>  csgF    G6544    b1038    ECK1024<br><b>Location:</b> 1,100,934 <- 1,101,350  <br> <b>Product:</b> curli assembly component, subunit of Curli Secretion and Assembly Complex');" onmouseout="return nd();">
 +
 
 +
<area coords="407,114,476,145" onmouseover="return overlib(' <b>Gene:</b>  csgG    G6543    b1037    ECK1023<br><b>Location:</b> 1,100,074 <- 1,100,907  <br> <b>Product:</b> curli production component, subunit of Curli Secretion and Assembly Complex');" onmouseout="return nd();">
 +
<area coords="121,0,280,162" onmouseover="return overlib('<b>Operon:</b> csgDp1 (Experim. ev.)  <BR><b>Tr.Activators:</b> (OmpR-P<SUP>asp</SUP> CsgD CRP-cAMP) <BR><b>Tr.Inhibitors:</b> (RstA-P<SUP>asp</SUP> RcsAB CpxR-P<SUP>asp</SUP>)');" onmouseout="return nd();">
 +
<area coords="0,0,478,162" onmouseover="return overlib('<b>Operon</b>: csgDEFG');" onmouseout="return nd();">
 +
</map>
 +
<img src="https://static.igem.org/mediawiki/2009/a/aa/Op6.gif" usemap="#MAP91787" border="0">
 +
</p>
 +
 
 +
</html>
 +
 
 +
 
 +
 
 +
<html>
 +
<p class="ecoparagraph">
 +
<map name="MAP91789">
 +
<area coords="198,18,228,35" onmouseover="return overlib('H-NS transcriptional dual regulator activates transcription from associated promoter<br>The exact location of this binding site is unknown or not specified.<br><b>Evidence:</b> Binding of purified proteins [Soutourina99], Gene expression analysis [Soutourina99]<br>The nature of binding of this protein is not well known.');" onmouseout="return nd();">
 +
<area coords="1,92,32,109" onmouseover="return overlib('OmpR-Phosphorylated transcriptional dual regulator represses transcription from associated promoter<br><b>Distance to transcription start site:</b> -145.5 (ranging from -136 to -155)<br><b>Evidence:</b> Human inference based on similarity to consensus sequences [Shin95]');" onmouseout="return nd();">
 +
<area coords="33,92,64,109" onmouseover="return overlib('LrhA transcriptional dual regulator represses transcription from associated promoter<br><b>Distance to transcription start site:</b> -112 (ranging from -105 to -119)<br><b>Evidence:</b> Gene expression analysis [Lehnen02], Binding of purified proteins [Lehnen02]');" onmouseout="return nd();">
 +
<area coords="64,92,95,109" onmouseover="return overlib('IHF transcriptional dual regulator represses transcription from associated promoter<br><b>Distance to transcription start site:</b> -92 (ranging from -86 to -98)<br><b>Evidence:</b> Human inference based on similarity to consensus sequences [Shin95]');" onmouseout="return nd();">
 +
<area coords="95,92,126,109"  onmouseover="return overlib('CRP-cAMP transcriptional dual regulator activates transcription from associated promoter<br><b>Distance to transcription start site:</b> -71.5 (ranging from -61 to -82)<br><b>Evidence:</b> Gene expression analysis, Binding of purified proteins, Human inference based on similarity to consensus sequences');" onmouseout="return nd();">
 +
<area coords="199,92,230,109" onmouseover="return overlib('Fur-Fe+2 transcriptional dual regulator represses transcription from associated promoter<br><b>Distance to transcription start site:</b> 115 (ranging from 124 to 106)<br><b>Evidence:</b> Automated inference based on similarity to consensus sequences [Stojiljkov94]');" onmouseout="return nd();">
 +
<area coords="134,92,166,109" onmouseover="return overlib('RcsAB transcriptional dual regulator represses transcription from associated promoter<br><b>Distance to transcription start site:</b> 11.5 (ranging from 18 to 5)<br><b>Evidence:</b> Binding of purified proteins [FrancezCha03], Gene expression analysis [FrancezCha03], Site mutation [FrancezCha03]');" onmouseout="return nd();">
 +
<area coords="198,0,234,17" onmouseover="return overlib('QseB-Phosphorylated transcriptional activator activates transcription from associated promoter<br>The exact location of this binding site is unknown or not specified.<br><b>Evidence:</b> Gene expression analysis [Sperandio02]');" onmouseout="return nd();">
 +
<area coords="198,36,234,53" onmouseover="return overlib('HdfR transcriptional repressor represses transcription from associated promoter<br>The exact location of this binding site is unknown or not specified.<br><b>Evidence:</b> Binding of purified proteins [Ko00], Gene expression analysis [Ko00]');" onmouseout="return nd();">
 +
<area coords="144,74,188,91" onmouseover="return overlib('OmpR-Phosphorylated transcriptional dual regulator represses transcription from associated promoter<br><b>Distance to transcription start site:</b> 18.5 (ranging from 28 to 9)<br><b>Evidence:</b> Human inference based on similarity to consensus sequences [Shin95]');" onmouseout="return nd();">
 +
<area coords="241,78,296,109" onmouseover="return overlib(' <b>Gene:</b>  flhD    EG10320    b1892    ECK1893<br><b>Synonyms:</b> flbB<br><b>Location:</b> 1,975,871 <- 1,976,221  <br> <b>Product:</b> FlhD, subunit of FlhDC transcriptional dual regulator');" onmouseout="return nd();">
 +
<area coords="296,78,357,109" onmouseover="return overlib(' <b>Gene:</b>  flhC    EG10319    b1891    ECK1892<br><b>Synonyms:</b> flaI<br><b>Location:</b> 1,975,290 <- 1,975,868  <br> <b>Product:</b> FlhC, subunit of FlhDC transcriptional dual regulator');" onmouseout="return nd();">
 +
<area coords="77,0,234,126"  onmouseover="return overlib('<b>Operon:</b> flhDp (Experim. ev.)  <BR><b>Tr.Activators:</b> (Hns CRP-cAMP QseB-Phosphorylated transcriptional activator) <BR><b>Tr.Inhibitors:</b> (OmpR-P<SUP>asp</SUP> LrhA HdfR RcsAB Fur IHF)');" onmouseout="return nd();">
 +
<area coords="0,0,359,126" onmouseover="return overlib('<b>Operon</b>: flhDC<br><b>Synonyms:</b> OP00183');" onmouseout="return nd();">
 +
</map>
 +
 
 +
<img src="https://static.igem.org/mediawiki/2009/3/3c/Op7.gif" usemap="#MAP91789" border="0">
 +
</p>
 +
 
 +
</html>
 +
 
 +
<html><center><a href="https://2009.igem.org/Team:Imperial_College_London/M2"><img width=150px src="https://static.igem.org/mediawiki/2009/1/10/II09_TourArrow.png"></a>
 +
</html>
 +
 
-
<b>Background:</b>
 
-
In nature, the colanic acid synthesis phase occurs prior to biofilm formation. The latter process of biofilm formation is associated with the upregulation of a number of virulence factors. The transcription factor YgiV blocks the progression into biofilm formation by maintaining colanic acid production. Thus YgiV serves to increase acid resistance and decrease virulence.
 
{{Imperial/09/Division}}
{{Imperial/09/Division}}
-
===Rfal===
+
<b>References</b>
 +
 
 +
1) [http://www.ncbi.nlm.nih.gov/pubmed/19136591 <b>The Rcs two-component system regulates expression of lysozyme inhibitors and is induced by exposure to lysozyme</b>]
 +
 
 +
2) [http://www.ncbi.nlm.nih.gov/pubmed/19346340 <b>Gene Expression Induced in Escherichia coli O157:H7 upon Exposure to Model Apple Juice</b>]
 +
 
 +
3) [http://www.ncbi.nlm.nih.gov/pubmed/14627744 <b>Structural and functional features of the Escherichia coli hydroperoxide resistance protein OsmC.</b>]
 +
 
 +
4) [http://www.ncbi.nlm.nih.gov/pubmed/16153174?ordinalpos=5&itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DefaultReportPanel.Pubmed_RVDocSum <b>The Rcs phosphorelay: a complex signal transduction system.</b>]
 +
 
 +
 
 +
<hr>  <h3> <b>Interactive Image References</b>
 +
 
 +
Note, interactive operons are credited to EcoCyc: Nucleic Acids Research 37:D464-70 2009
 +
 
 +
<html>
 +
</h3>  <p class="ecoparagraph"> <a name="16606699"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=16606699&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('16606699');" onmouseout="return nd();">  <b>Arifuzzama06</b>: "Large-scale identification of protein-protein interaction of Escherichia coli K-12." Genome Res 16(5);686-91. PMID: 16606699
 +
</a>  </p><p class="ecoparagraph"> <a name="15838058"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=15838058&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('15838058');" onmouseout="return nd();">  <b>Boulanger05</b>:  "Multistress regulation in Escherichia coli: expression of osmB involves two independent promoters responding either to sigmaS or to the RcsCDB His-Asp phosphorelay." J Bacteriol 187(9);3282-6. PMID: 15838058
 +
</a>  </p><p class="ecoparagraph"> <a name="2836365"></a>
 +
 
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=2836365&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('2836365');" onmouseout="return nd();">  <b>Brill88</b>:  "Fine-structure mapping and identification of two regulators of capsule synthesis in Escherichia coli K-12." J Bacteriol 1988;170(6);2599-611. PMID: 2836365
 +
</a>  </p><p class="ecoparagraph"> <a name="10564486"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=10564486&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('10564486');" onmouseout="return nd();">  <b>Carballes99</b>:  "Regulation of Escherichia coli cell division genes ftsA and ftsZ by the two-component system rcsC-rcsB." Mol Microbiol 1999;34(3);442-50. PMID: 10564486
 +
</a>  </p><p class="ecoparagraph"> <a name="16740933"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=16740933&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('16740933');" onmouseout="return nd();">  <b>CastanieCo06</b>:  "RcsF is an outer membrane lipoprotein involved in the RcsCDB phosphorelay signaling pathway in Escherichia coli." J Bacteriol 188(12);4264-70. PMID: 16740933
 +
 
 +
</a>  </p><p class="ecoparagraph"> <a name="17185552"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=17185552&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('17185552');" onmouseout="return nd();">  <b>CastanieCo07</b>:  "The glutamate-dependent acid resistance system in Escherichia coli: essential and dual role of the His-Asp phosphorelay RcsCDB/AF." Microbiology 153(Pt 1);238-46. PMID: 17185552
 +
</a>  </p><p class="ecoparagraph"> <a name="11758943"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=11758943&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('11758943');" onmouseout="return nd();">  <b>Chen01c</b>:  "Characterization of the RcsC--&gt;YojN--&gt;RcsB phosphorelay signaling pathway involved in capsular synthesis in Escherichia coli." Biosci Biotechnol Biochem 65(10);2364-7. PMID: 11758943
 +
</a>  </p><p class="ecoparagraph"> <a name="19026860"></a>
 +
 
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=19026860&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('19026860');" onmouseout="return nd();">  <b>Corbett08</b>:  "Capsular polysaccharides in Escherichia coli." Adv Appl Microbiol 65;1-26. PMID: 19026860
 +
</a>  </p><p class="ecoparagraph"> <a name="11566985"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=11566985&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('11566985');" onmouseout="return nd();">  <b>DavalosGar01</b>:  "Regulation of osmC gene expression by the two-component system rcsB-rcsC in Escherichia coli." J Bacteriol 183(20);5870-6. PMID: 11566985
 +
</a>  </p><p class="ecoparagraph"> <a name="8759878"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=8759878&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('8759878');" onmouseout="return nd();">  <b>Dierksen96</b>:  "Identification of a second RcsA protein, a positive regulator of colanic acid capsular polysaccharide genes, in Escherichia coli." J Bacteriol 178(16);5053-6. PMID: 8759878
 +
 
 +
</a>  </p><p class="ecoparagraph"> <a name="11309136"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=11309136&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('11309136');" onmouseout="return nd();">  <b>Dominguez01</b>:  "Leishmania immune adherence reaction in vertebrates." Parasite Immunol 23(5);259-65. PMID: 11309136
 +
</a>  </p><p class="ecoparagraph"> <a name="9882673"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=9882673&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('9882673');" onmouseout="return nd();">  <b>Ebel99</b>:  "Escherichia coli RcsA, a positive activator of colanic acid capsular polysaccharide synthesis, functions to activate its own expression." J Bacteriol 1999;181(2);577-84. PMID: 9882673
 +
</a>  </p><p class="ecoparagraph"> <a name="17379715"></a>
 +
 
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=17379715&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('17379715');" onmouseout="return nd();">  <b>Ferrieres07</b>:  "The yjbEFGH locus in Escherichia coli K-12 is an operon encoding proteins involved in exopolysaccharide production." Microbiology 153(Pt 4);1070-80. PMID: 17379715
 +
</a>  </p><p class="ecoparagraph"> <a name="12864862"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=12864862&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('12864862');" onmouseout="return nd();">  <b>FrancezCha03</b>:  Francez-Charlot A, Laugel B, Van Gemert A, Dubarry N, Wiorowski F, Castani?-Cornet MP, Gutierrez C, Cam K (2003). "RcsCDB His-Asp phosphorelay system negatively regulates the flhDC operon in Escherichia coli." Mol Microbiol 49(3);823-32. PMID: 12864862
 +
</a>  </p><p class="ecoparagraph"> <a name="15901715"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=15901715&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('15901715');" onmouseout="return nd();">  <b>FrancezCha05</b>:  Francez-Charlot A, Castanie-Cornet MP, Gutierrez C, Cam K (2005). "Osmotic Regulation of the Escherichia coli bdm (Biofilm-Dependent Modulation) Gene by the RcsCDB His-Asp Phosphorelay." J Bacteriol 187(11);3873-7. PMID: 15901715
 +
 
 +
</a>  </p><p class="ecoparagraph"> <a name="16776655"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=16776655&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('16776655');" onmouseout="return nd();">  <b>Fredericks06</b>:  Fredericks CE, Shibata S, Aizawa S, Reimann SA, Wolfe AJ (2006). "Acetyl phosphate-sensitive regulation of flagellar biogenesis and capsular biosynthesis depends on the Rcs phosphorelay." Mol Microbiol 61(3);734-47. PMID: 16776655
 +
</a>  </p><p class="ecoparagraph"> <a name="1597415"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=1597415&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('1597415');" onmouseout="return nd();">  <b>Gervais92</b>:  Gervais FG, Phoenix P, Drapeau GR (1992). "The rcsB gene, a positive regulator of colanic acid biosynthesis in Escherichia coli, is also an activator of ftsZ expression." J Bacteriol 1992;174(12);3964-71. PMID: 1597415
 +
</a>  </p><p class="ecoparagraph"> <a name="GO_REF-0000004"></a>
 +
 
 +
<a href="http://www.geneontology.org/doc/GO.references" class="CITATION-FRAME" onmouseover="return overlib('GO_REF-0000004');" onmouseout="return nd();">  <b>GOA00</b>:  GOA (2000). "Gene Ontology annotation based on Swiss-Prot keyword mapping."
 +
</a>  </p><p class="ecoparagraph"> <a name="GO_REF-0000002"></a>
 +
<a href="http://www.geneontology.org/doc/GO.references" class="CITATION-FRAME" onmouseover="return overlib('GO_REF-0000002');" onmouseout="return nd();">  <b>GOA01</b>:  GOA, DDB, FB, MGI, ZFIN (2001). "Gene Ontology annotation through association of InterPro records with GO terms."
 +
</a>  </p><p class="ecoparagraph"> <a name="3888955"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=3888955&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('3888955');" onmouseout="return nd();">  <b>Gottesman85</b>:  Gottesman S, Trisler P, Torres-Cabassa A (1985). "Regulation of capsular polysaccharide synthesis in Escherichia coli K-12: characterization of three regulatory genes." J Bacteriol 162(3);1111-9. PMID: 3888955
 +
 
 +
</a>  </p><p class="ecoparagraph"> <a name="1943696"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=1943696&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('1943696');" onmouseout="return nd();">  <b>Gottesman91</b>:  Gottesman S, Stout V (1991). "Regulation of capsular polysaccharide synthesis in Escherichia coli K12." Mol Microbiol 1991;5(7);1599-606. PMID: 1943696
 +
</a>  </p><p class="ecoparagraph"> <a name="9209051"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=9209051&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('9209051');" onmouseout="return nd();">  <b>Gupte97</b>:  Gupte G, Woodward C, Stout V (1997). "Isolation and characterization of rcsB mutations that affect colanic acid capsule synthesis in Escherichia coli K-12." J Bacteriol 1997;179(13);4328-35. PMID: 9209051
 +
</a>  </p><p class="ecoparagraph"> <a name="12618449"></a>
 +
 
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=12618449&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('12618449');" onmouseout="return nd();">  <b>Hirakawa03</b>:  Hirakawa H, Nishino K, Hirata T, Yamaguchi A (2003). "Comprehensive studies of drug resistance mediated by overexpression of response regulators of two-component signal transduction systems in Escherichia coli." J Bacteriol 185(6);1851-6. PMID: 12618449
 +
</a>  </p><p class="ecoparagraph"> <a name="16427772"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=16427772&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('16427772');" onmouseout="return nd();">  <b>Huang06a</b>:  Huang YH, Ferrieres L, Clarke DJ (2006). "The role of the Rcs phosphorelay in Enterobacteriaceae." Res Microbiol 157(3);206-12. PMID: 16427772
 +
</a>  </p><p class="ecoparagraph"> <a name="8366025"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=8366025&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('8366025');" onmouseout="return nd();">  <b>Jayaratne93</b>:  Jayaratne P, Keenleyside WJ, MacLachlan PR, Dodgson C, Whitfield C (1993). "Characterization of rcsB and rcsC from Escherichia coli O9:K30:H12 and examination of the role of the rcs regulatory system in expression of group I capsular polysaccharides." J Bacteriol 1993;175(17);5384-94. PMID: 8366025
 +
 
 +
</a>  </p><p class="ecoparagraph"> <a name="14766922"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=14766922&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('14766922');" onmouseout="return nd();">  <b>Kuo04</b>:  Kuo MS, Chen KP, Wu WF (2004). "Regulation of RcsA by the ClpYQ (HslUV) protease in Escherichia coli." Microbiology 150(Pt 2);437-46. PMID: 14766922
 +
</a>  </p><p class="ecoparagraph"> <a name="12670962"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=12670962&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('12670962');" onmouseout="return nd();">  <b>Lee03b</b>:  Lee YY, Chang CF, Kuo CL, Chen MC, Yu CH, Lin PI, Wu WF (2003). "Subunit oligomerization and substrate recognition of the Escherichia coli ClpYQ (HslUV) protease implicated by in vivo protein-protein interactions in the yeast two-hybrid system." J Bacteriol 185(8);2393-401. PMID: 12670962
 +
</a>  </p><p class="ecoparagraph"> <a name="12410838"></a>
 +
 
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=12410838&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('12410838');" onmouseout="return nd();">  <b>Majdalani02</b>:  Majdalani N, Hernandez D, Gottesman S (2002). "Regulation and mode of action of the second small RNA activator of RpoS translation, RprA." Mol Microbiol 46(3);813-26. PMID: 12410838
 +
</a>  </p><p class="ecoparagraph"> <a name="16153174"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=16153174&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('16153174');" onmouseout="return nd();">  <b>Majdalani05</b>:  Majdalani N, Gottesman S (2005). "The Rcs phosphorelay: a complex signal transduction system." Annu Rev Microbiol 59;379-405. PMID: 16153174
 +
</a>  </p><p class="ecoparagraph"> <a name="16166540"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=16166540&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('16166540');" onmouseout="return nd();">  <b>Majdalani05b</b>:  Majdalani N, Heck M, Stout V, Gottesman S (2005). "Role of RcsF in signaling to the Rcs phosphorelay pathway in Escherichia coli." J Bacteriol 187(19);6770-8. PMID: 16166540
 +
 
 +
</a>  </p><p class="ecoparagraph"> <a name="14514676"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=14514676&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('14514676');" onmouseout="return nd();">  <b>Mouslim03</b>:  Mouslim C, Latifi T, Groisman EA (2003). "Signal-dependent requirement for the co-activator protein RcsA in transcription of the RcsB-regulated ugd gene." J Biol Chem 278(50);50588-95. PMID: 14514676
 +
</a>  </p><p class="ecoparagraph"> <a name="17468243"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=17468243&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('17468243');" onmouseout="return nd();">  <b>Ogasawara07a</b>:  Ogasawara H, Hasegawa A, Kanda E, Miki T, Yamamoto K, Ishihama A (2007). "Genomic SELEX search for target promoters under the control of the PhoQP-RstBA signal relay cascade." J Bacteriol 189(13);4791-9. PMID: 17468243
 +
</a>  </p><p class="ecoparagraph"> <a name="7699720"></a>
 +
 
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=7699720&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('7699720');" onmouseout="return nd();">  <b>Pao95</b>:  Pao GM, Saier MH (1995). "Response regulators of bacterial signal transduction systems: selective domain shuffling during evolution." J Mol Evol 1995;40(2);136-54. PMID: 7699720
 +
</a>  </p><p class="ecoparagraph"> <a name="1482126"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=1482126&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('1482126');" onmouseout="return nd();">  <b>Parkinson92</b>:  Parkinson JS, Kofoid EC (1992). "Communication modules in bacterial signaling proteins." Annu Rev Genet 1992;26;71-112. PMID: 1482126
 +
</a>  </p><p class="ecoparagraph"> <a name="8098993"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=8098993&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('8098993');" onmouseout="return nd();">  <b>Parkinson93</b>:  Parkinson JS (1993). "Signal transduction schemes of bacteria." Cell 1993;73(5);857-71. PMID: 8098993
 +
 
 +
</a>  </p><p class="ecoparagraph"> <a name="7534408"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=7534408&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('7534408');" onmouseout="return nd();">  <b>Sledjeski95</b>:  Sledjeski D, Gottesman S (1995). "A small RNA acts as an antisilencer of the H-NS-silenced rcsA gene of Escherichia coli." Proc Natl Acad Sci U S A 1995;92(6);2003-7. PMID: 7534408
 +
</a>  </p><p class="ecoparagraph"> <a name="8576059"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=8576059&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('8576059');" onmouseout="return nd();">  <b>Sledjeski96</b>:  Sledjeski DD, Gottesman S (1996). "Osmotic shock induction of capsule synthesis in Escherichia coli K-12." J Bacteriol 1996;178(4);1204-6. PMID: 8576059
 +
</a>  </p><p class="ecoparagraph"> <a name="2157156"></a>
 +
 
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=2157156&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('2157156');" onmouseout="return nd();">  <b>Stock90</b>:  Stock JB, Stock AM, Mottonen JM (1990). "Signal transduction in bacteria." Nature 1990;344(6265);395-400. PMID: 2157156
 +
</a>  </p><p class="ecoparagraph"> <a name="2404948"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=2404948&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('2404948');" onmouseout="return nd();">  <b>Stout90</b>:  Stout V, Gottesman S (1990). "RcsB and RcsC: a two-component regulator of capsule synthesis in Escherichia coli." J Bacteriol 1990;172(2);659-69. PMID: 2404948
 +
</a>  </p><p class="ecoparagraph"> <a name="1999391"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=1999391&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('1999391');" onmouseout="return nd();">  <b>Stout91</b>:  Stout V, Torres-Cabassa A, Maurizi MR, Gutnick D, Gottesman S (1991). "RcsA, an unstable positive regulator of capsular polysaccharide synthesis." J Bacteriol 173(5);1738-47. PMID: 1999391
 +
 
 +
</a>  </p><p class="ecoparagraph"> <a name="7855424"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=7855424&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('7855424');" onmouseout="return nd();">  <b>Stout94</b>:  Stout V "Regulation of capsule synthesis includes interactions of the RcsC/RcsB regulatory pair." Res Microbiol 1994;145(5-6);389-92. PMID: 7855424
 +
</a>  </p><p class="ecoparagraph"> <a name="8763957"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=8763957&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('8763957');" onmouseout="return nd();">  <b>Stout96</b>:  Stout V (1996). "Identification of the promoter region for the colanic acid polysaccharide biosynthetic genes in Escherichia coli K-12." J Bacteriol 178(14);4273-80. PMID: 8763957
 +
</a>  </p><p class="ecoparagraph"> <a name="3029041"></a>
 +
 
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=3029041&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('3029041');" onmouseout="return nd();">  <b>TorresCaba87</b>:  Torres-Cabassa AS, Gottesman S (1987). "Capsule synthesis in Escherichia coli K-12 is regulated by proteolysis." J Bacteriol 169(3);981-9. PMID: 3029041
 +
</a>  </p><p class="ecoparagraph"> <a name="UNIPROT-14-6"></a>
 +
<a href="http://BioCyc.org/ECOLI/reference.html?type=CITATION-FRAME&amp;object=UNIPROT-14-6" class="CITATION-FRAME" onmouseover="return overlib('UNIPROT-14-6');" onmouseout="return nd();">  <b>UniProt08</b>:  UniProt Consortium (2008). "UniProt version 14.6 released on 2008-12-16." Database.
 +
</a>  </p><p class="ecoparagraph"> <a name="16000739"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=16000739&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('16000739');" onmouseout="return nd();">  <b>Vianney05</b>:  Vianney A, Jubelin G, Renault S, Dorel C, Lejeune P, Lazzaroni JC (2005). "Escherichia coli tol and rcs genes participate in the complex network affecting curli synthesis." Microbiology 151(Pt 7);2487-97. PMID: 16000739
 +
 
 +
</a>  </p><p class="ecoparagraph"> <a name="10702265"></a>
 +
<a href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;list_uids=10702265&amp;dopt=Abstract" class="CITATION-FRAME" onmouseover="return overlib('10702265');" onmouseout="return nd();">  <b>Wehland00</b>:  Wehland M, Bernhard F (2000). "The RcsAB box. Characterization of a new operator essential for the regulation of exopolysaccharide biosynthesis in enteric bacteria." J Biol Chem 2000;275(10);7013-20. PMID: 10702265
-
<b>Background:</b>
+
</html>
-
In the majority of E.coli, the enzyme Rfal joins the O-antigen to the membrane-bound lipid core molecule. Since the K-12 strain has an insertion mutation in the gene coding for O-antigen, the enzyme Rfal is free to join colanic acid to the lipid core.
 
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Latest revision as of 21:47, 21 October 2009

Contents

Regulation for Capsular Synthesis (RCS) Phosphorelay System

RcsB is a transcription factor that forms part of a phosphorelay system. As the name suggests, a phosphorelay system consists of a series of nodes down which a phosphate group is passed. The transfer of this phosphate group down along the Rcs relay system is initiated by membrane stress. Once the phosphate group has been passed to RcsB, it can bind to DNA and act as a transcription factor. Phosphorylated RcsB is thought to regulate ~5% of the E.coli genome 4.

PhosRelay.png

The Rcs phosphorelay is composed of two membrane-anchored proteins, RcsC and RcsD, and a cytoplasmic protein, RcsB. In response to a signal, the sensor kinase RcsC autophosphorylates and the phosphoryl group is transferred to RcsB, via RcsD. RcsB can both homodimerise and heterodimerise with RcsA, these two complexes serve to regulate different operons.


Signal Amplification:

RcsB is able to heterodimerise with the protein RcsA. The RcsB+RcsA complex acts as a transcription factor that induces the expression of a number of genes. One of these genes codes for RcsA, this initiates a positive feedback loop.



Colanic Acid Biosynthesis:

The overexpression of RcsB is akin to fitting a stethoscope to the end of a megaphone. In essence, the sensitivity of the RCS Phosphorelay system becomes such that the smallest of membrane perturbations becomes sufficient to trigger colanic acid production. Thus, the background noise switches the RCS Phosphorelay system on permanently.

There are a total of 19 genes involved in colonic acid biosynthesis, the best characterised of which are shown below.


Other genes involved in colanic acid biosynthesis are listed below.

wcaF
  • Putative colanic acid biosynthesis acetyltransferase 2
gmd

Codes for GDP-mannose 4,6-dehydratase which is a subunit of GDP-mannose 4,6-dehydratase. This enzyme is involved in colanic acid biosynthesis.

wcaD
  • Predicted colanic acid polymerase
fcl
  • Codes for a subunit of GDP-fucose synthase. This enzyme is involved in colanic acid biosynthesis.



General Protective:

The following proteins are thought play a role in membrane fortification to confer protection against the rigours of dessication, acid exposure and osmotic stress.



Other RcsB induced genes coding for protective proteins are listed below.

KatE
  • Codes for a subunit of heme d synthase / hydroperoxidase. This bacterial catalase protects against osmotic stress.


Host Defence:

Humans synthesise a number of enzymes that are able to burst bacterial cells. These enzymes are called lysosymes. RcsB is able to induce a number of proteins that act as lysosyme inhibitors that will prevent the host-mediated lysis of our chassis.


ivy (Inhibitor of Vertebrate lysozyme)
  • Discovered in 2001 as the first bacterial lysozyme inhibitor. This Type-C lysozyme inhibitor resides in the periplasm.1
MilC (Membrane-bound lysozyme inhibitor of Type C lysozyme)
  • This is a lipoprotein that resides in the membrane. 1



Global Regulators:






Repression:

RcsB is also is able to act as a repressor, preventing the expression of both curli and fimbriae. This reduces cell motility and plays a role in biofilm formation.







References

1) [http://www.ncbi.nlm.nih.gov/pubmed/19136591 The Rcs two-component system regulates expression of lysozyme inhibitors and is induced by exposure to lysozyme]

2) [http://www.ncbi.nlm.nih.gov/pubmed/19346340 Gene Expression Induced in Escherichia coli O157:H7 upon Exposure to Model Apple Juice]

3) [http://www.ncbi.nlm.nih.gov/pubmed/14627744 Structural and functional features of the Escherichia coli hydroperoxide resistance protein OsmC.]

4) [http://www.ncbi.nlm.nih.gov/pubmed/16153174?ordinalpos=5&itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DefaultReportPanel.Pubmed_RVDocSum The Rcs phosphorelay: a complex signal transduction system.]


Interactive Image References

Note, interactive operons are credited to EcoCyc: Nucleic Acids Research 37:D464-70 2009

Arifuzzama06: "Large-scale identification of protein-protein interaction of Escherichia coli K-12." Genome Res 16(5);686-91. PMID: 16606699

Boulanger05: "Multistress regulation in Escherichia coli: expression of osmB involves two independent promoters responding either to sigmaS or to the RcsCDB His-Asp phosphorelay." J Bacteriol 187(9);3282-6. PMID: 15838058

Brill88: "Fine-structure mapping and identification of two regulators of capsule synthesis in Escherichia coli K-12." J Bacteriol 1988;170(6);2599-611. PMID: 2836365

Carballes99: "Regulation of Escherichia coli cell division genes ftsA and ftsZ by the two-component system rcsC-rcsB." Mol Microbiol 1999;34(3);442-50. PMID: 10564486

CastanieCo06: "RcsF is an outer membrane lipoprotein involved in the RcsCDB phosphorelay signaling pathway in Escherichia coli." J Bacteriol 188(12);4264-70. PMID: 16740933

CastanieCo07: "The glutamate-dependent acid resistance system in Escherichia coli: essential and dual role of the His-Asp phosphorelay RcsCDB/AF." Microbiology 153(Pt 1);238-46. PMID: 17185552

Chen01c: "Characterization of the RcsC-->YojN-->RcsB phosphorelay signaling pathway involved in capsular synthesis in Escherichia coli." Biosci Biotechnol Biochem 65(10);2364-7. PMID: 11758943

Corbett08: "Capsular polysaccharides in Escherichia coli." Adv Appl Microbiol 65;1-26. PMID: 19026860

DavalosGar01: "Regulation of osmC gene expression by the two-component system rcsB-rcsC in Escherichia coli." J Bacteriol 183(20);5870-6. PMID: 11566985

Dierksen96: "Identification of a second RcsA protein, a positive regulator of colanic acid capsular polysaccharide genes, in Escherichia coli." J Bacteriol 178(16);5053-6. PMID: 8759878

Dominguez01: "Leishmania immune adherence reaction in vertebrates." Parasite Immunol 23(5);259-65. PMID: 11309136

Ebel99: "Escherichia coli RcsA, a positive activator of colanic acid capsular polysaccharide synthesis, functions to activate its own expression." J Bacteriol 1999;181(2);577-84. PMID: 9882673

Ferrieres07: "The yjbEFGH locus in Escherichia coli K-12 is an operon encoding proteins involved in exopolysaccharide production." Microbiology 153(Pt 4);1070-80. PMID: 17379715

FrancezCha03: Francez-Charlot A, Laugel B, Van Gemert A, Dubarry N, Wiorowski F, Castani?-Cornet MP, Gutierrez C, Cam K (2003). "RcsCDB His-Asp phosphorelay system negatively regulates the flhDC operon in Escherichia coli." Mol Microbiol 49(3);823-32. PMID: 12864862

FrancezCha05: Francez-Charlot A, Castanie-Cornet MP, Gutierrez C, Cam K (2005). "Osmotic Regulation of the Escherichia coli bdm (Biofilm-Dependent Modulation) Gene by the RcsCDB His-Asp Phosphorelay." J Bacteriol 187(11);3873-7. PMID: 15901715

Fredericks06: Fredericks CE, Shibata S, Aizawa S, Reimann SA, Wolfe AJ (2006). "Acetyl phosphate-sensitive regulation of flagellar biogenesis and capsular biosynthesis depends on the Rcs phosphorelay." Mol Microbiol 61(3);734-47. PMID: 16776655

Gervais92: Gervais FG, Phoenix P, Drapeau GR (1992). "The rcsB gene, a positive regulator of colanic acid biosynthesis in Escherichia coli, is also an activator of ftsZ expression." J Bacteriol 1992;174(12);3964-71. PMID: 1597415

GOA00: GOA (2000). "Gene Ontology annotation based on Swiss-Prot keyword mapping."

GOA01: GOA, DDB, FB, MGI, ZFIN (2001). "Gene Ontology annotation through association of InterPro records with GO terms."

Gottesman85: Gottesman S, Trisler P, Torres-Cabassa A (1985). "Regulation of capsular polysaccharide synthesis in Escherichia coli K-12: characterization of three regulatory genes." J Bacteriol 162(3);1111-9. PMID: 3888955

Gottesman91: Gottesman S, Stout V (1991). "Regulation of capsular polysaccharide synthesis in Escherichia coli K12." Mol Microbiol 1991;5(7);1599-606. PMID: 1943696

Gupte97: Gupte G, Woodward C, Stout V (1997). "Isolation and characterization of rcsB mutations that affect colanic acid capsule synthesis in Escherichia coli K-12." J Bacteriol 1997;179(13);4328-35. PMID: 9209051

Hirakawa03: Hirakawa H, Nishino K, Hirata T, Yamaguchi A (2003). "Comprehensive studies of drug resistance mediated by overexpression of response regulators of two-component signal transduction systems in Escherichia coli." J Bacteriol 185(6);1851-6. PMID: 12618449

Huang06a: Huang YH, Ferrieres L, Clarke DJ (2006). "The role of the Rcs phosphorelay in Enterobacteriaceae." Res Microbiol 157(3);206-12. PMID: 16427772

Jayaratne93: Jayaratne P, Keenleyside WJ, MacLachlan PR, Dodgson C, Whitfield C (1993). "Characterization of rcsB and rcsC from Escherichia coli O9:K30:H12 and examination of the role of the rcs regulatory system in expression of group I capsular polysaccharides." J Bacteriol 1993;175(17);5384-94. PMID: 8366025

Kuo04: Kuo MS, Chen KP, Wu WF (2004). "Regulation of RcsA by the ClpYQ (HslUV) protease in Escherichia coli." Microbiology 150(Pt 2);437-46. PMID: 14766922

Lee03b: Lee YY, Chang CF, Kuo CL, Chen MC, Yu CH, Lin PI, Wu WF (2003). "Subunit oligomerization and substrate recognition of the Escherichia coli ClpYQ (HslUV) protease implicated by in vivo protein-protein interactions in the yeast two-hybrid system." J Bacteriol 185(8);2393-401. PMID: 12670962

Majdalani02: Majdalani N, Hernandez D, Gottesman S (2002). "Regulation and mode of action of the second small RNA activator of RpoS translation, RprA." Mol Microbiol 46(3);813-26. PMID: 12410838

Majdalani05: Majdalani N, Gottesman S (2005). "The Rcs phosphorelay: a complex signal transduction system." Annu Rev Microbiol 59;379-405. PMID: 16153174

Majdalani05b: Majdalani N, Heck M, Stout V, Gottesman S (2005). "Role of RcsF in signaling to the Rcs phosphorelay pathway in Escherichia coli." J Bacteriol 187(19);6770-8. PMID: 16166540

Mouslim03: Mouslim C, Latifi T, Groisman EA (2003). "Signal-dependent requirement for the co-activator protein RcsA in transcription of the RcsB-regulated ugd gene." J Biol Chem 278(50);50588-95. PMID: 14514676

Ogasawara07a: Ogasawara H, Hasegawa A, Kanda E, Miki T, Yamamoto K, Ishihama A (2007). "Genomic SELEX search for target promoters under the control of the PhoQP-RstBA signal relay cascade." J Bacteriol 189(13);4791-9. PMID: 17468243

Pao95: Pao GM, Saier MH (1995). "Response regulators of bacterial signal transduction systems: selective domain shuffling during evolution." J Mol Evol 1995;40(2);136-54. PMID: 7699720

Parkinson92: Parkinson JS, Kofoid EC (1992). "Communication modules in bacterial signaling proteins." Annu Rev Genet 1992;26;71-112. PMID: 1482126

Parkinson93: Parkinson JS (1993). "Signal transduction schemes of bacteria." Cell 1993;73(5);857-71. PMID: 8098993

Sledjeski95: Sledjeski D, Gottesman S (1995). "A small RNA acts as an antisilencer of the H-NS-silenced rcsA gene of Escherichia coli." Proc Natl Acad Sci U S A 1995;92(6);2003-7. PMID: 7534408

Sledjeski96: Sledjeski DD, Gottesman S (1996). "Osmotic shock induction of capsule synthesis in Escherichia coli K-12." J Bacteriol 1996;178(4);1204-6. PMID: 8576059

Stock90: Stock JB, Stock AM, Mottonen JM (1990). "Signal transduction in bacteria." Nature 1990;344(6265);395-400. PMID: 2157156

Stout90: Stout V, Gottesman S (1990). "RcsB and RcsC: a two-component regulator of capsule synthesis in Escherichia coli." J Bacteriol 1990;172(2);659-69. PMID: 2404948

Stout91: Stout V, Torres-Cabassa A, Maurizi MR, Gutnick D, Gottesman S (1991). "RcsA, an unstable positive regulator of capsular polysaccharide synthesis." J Bacteriol 173(5);1738-47. PMID: 1999391

Stout94: Stout V "Regulation of capsule synthesis includes interactions of the RcsC/RcsB regulatory pair." Res Microbiol 1994;145(5-6);389-92. PMID: 7855424

Stout96: Stout V (1996). "Identification of the promoter region for the colanic acid polysaccharide biosynthetic genes in Escherichia coli K-12." J Bacteriol 178(14);4273-80. PMID: 8763957

TorresCaba87: Torres-Cabassa AS, Gottesman S (1987). "Capsule synthesis in Escherichia coli K-12 is regulated by proteolysis." J Bacteriol 169(3);981-9. PMID: 3029041

UniProt08: UniProt Consortium (2008). "UniProt version 14.6 released on 2008-12-16." Database.

Vianney05: Vianney A, Jubelin G, Renault S, Dorel C, Lejeune P, Lazzaroni JC (2005). "Escherichia coli tol and rcs genes participate in the complex network affecting curli synthesis." Microbiology 151(Pt 7);2487-97. PMID: 16000739

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