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The galS protein is a member of the GalR-LacI family, the same family in which the GalR protein belongs. GalS is 53% identical and 85% similar with GalR. As a result, GalS represses transcription of the operon in an analagous manner to GalR, forming the repressosome and inducing HU protein binding. However, it represses transcription much less efficiently in comparison to GalR<ref name="Weickert&Adhya" />.
The galS protein is a member of the GalR-LacI family, the same family in which the GalR protein belongs. GalS is 53% identical and 85% similar with GalR. As a result, GalS represses transcription of the operon in an analagous manner to GalR, forming the repressosome and inducing HU protein binding. However, it represses transcription much less efficiently in comparison to GalR<ref name="Weickert&Adhya" />.


==References==
1. The galactose regulon of Escherichia coli- <ref name="Weickert&Adhya" />


2. Regulation of open complex formation at the Escherichia coli galactose operon promoters: Simultaneous interaction of RNA polymerase, gal repressor and CAP/cAMP- <ref name="Goodrich&McClure" />


3. Regulation of the gal Operon of Escherichia coli by the capR Gene*- <ref>{{cite journal|last1=Mackie|first1=George|last2=Wilson|first2=David B.|title=Regulation of the gal Operon of Escherichia coli by the capR Gene*|journal=The Journal of Biological Chemistry|date=May 25, 1972|volume=247|issue=10|page=2973-2978|url=http://www.jbc.org/content/247/10/2973.short}}</ref>


4. A control element within a structural gene: The gal operon of Escherichia coli- <ref name="Meher&Orosz&Adhya" />

5. Control of transcription termination- <ref>{{cite journal|last1=Adhya|first1=Sankar|last2=Gottesman|first2=Max|title=Control of transcription termination|journal=Annual Review of Biochemistry|date=1978|volume=47|page=967-996|doi=10.1146/annurev.bi.47.070178.004535|url=http://www.annualreviews.org/doi/abs/10.1146/annurev.bi.47.070178.004535}}</ref>

6. Regulatory sequences involved in the promotion and termination of RNA transcription- <ref>{{cite journal|last1=Rosenberg|first1=Martin|last2=Court|first2=Donald|title=Regulatory sequences involved in the promotion and termination of RNA transcription|journal=Annual Review of Biochemistry|date=1979|volume=13|page=319-353|doi=10.1146/annurev.ge.13.120179.001535|url=http://www.annualreviews.org/doi/abs/10.1146/annurev.ge.13.120179.001535}}</ref>

7. Two promoters, one inducible and one constitutive, control transcription of the Streptomyces lividans galactose operon-
<ref>{{cite journal|last1=Fornwald|first1=James A.|last2=Schmidt|first2=Francis J.|last3=Adams|first3=Craig W.|last4=Rosenberg|first4=Martin|last5=Brawner|first5=Mary E.|title=Two promoters, one inducible and one constitutive, control transcription of the Streptomyces lividans galactose operon|journal=Proceedings of the National Academy of Sciences|date=April 1987|volume=84|page=2130-2134|pmid=3031664|url=http://www.pnas.org/content/84/8/2130.full.pdf}}</ref>

8. Galactose Operon- <ref name="Spektrum" />

9. Quantification of the galactose-operon mRNAs 5 bases different in their 5’-ends<ref name="Ji&Jeon" />


{{reflist}}
{{reflist}}

Revision as of 15:03, 4 December 2015

Remember to keep to an encyclopedic style that is accessible. Good references but try to find something more recent too. BenjaminLaufer (talk) 15:49, 4 November 2015 (UTC)

The gal operon (galactose operon) is a prokaryotic operon necessary for galactose transport and metabolism in Escherichia coli, and many other bacteria including those within the Streptomyces genus. Although galactose is not the preferred carbon source in bacteria, D-galactose (one of two possible isomers) is important in E. coli as a building block for other cellular pathways[1] (ie. lactose synthesis, glucose conversion via the Leloir pathway, etc.). The gal operon contains genes coding for the enzymes necessary in this galactose to glucose conversion as well as the controls necessary for this process[2].



Gal Operon Structure

Structural Genes

Bacterial operons are polycistronic, meaning multiple gene products can be translated from one mRNA transcript[1]. The gal operon possesses four structural genes which appear in the order: galE, galT, galk, and galM[3][1]. As a result, whenever galactose metabolism is needed, the gal operon will be transcribed and the three gene products will then be translated. The three genes and their products are as follows:

  • galE is the first gene to appear in the gal operon and codes for the enzyme UDP-Galactose-4-Epimerase, or GALE, which assists the conversion of UDP-galactose to UDP-glucose
  • galT is the second gene to appear in the gal operon and codes for the enzyme Galactose-1-Phosphat-Uridyl-Transferase, or GALT, which assists the conversion of Galactose-1-P to UDP-Galactose
  • galK is the third gene to appear in the gal operon and codes for the enzyme Galactokinase, which phosphorylates Galactose to Galactose-1-P[2].
  • galM is the final gene to appear the gal operon and codes for the enzyme Mutarotase, which interconverts D-α-galactose and D-β-galactose.
The gal operon; the three structural genes, their products and how they associate with the galactose metabolic pathway.
The gal operon; the three structural genes, their products and how they associate with the galactose metabolic pathway.

The order of the genes, however, do not align with the order in which the respective gene products function in the galactose metabolic pathway. Instead, the order in which the gene products function is Mutarotase (galM), Galactokinase (galK), GALT (galT), then finally GALE (galE). Thus the gene products of the three structural genes in the gal operon function in reverse order in which they appear.

Regulatory Structures

Upstream of the three aforementioned structural genes, the gal operon has two overlapping promoters, which appear in the order: P2 and P1[3]. Both of the promoters have an associated transcription initiation site that appear five base pairs apart and are different in sequence.[4].

As well, the gal operon is associated with two operators, which appear in the order: OE and OI where E represents external and I represents internal. OI, the internal operator, is a structure that occurs within the structural gene galE. These two operators are separated by 113 base pairs, straddling both the promoters and transcription initiation sites with OE appearing at the beginning of the operon, and OI appearing a few base pairs upstream of the first structural gene galE[4].

A schematic presentation of the gal operon, including positioning of both promoters, both transcription initiation sites, as well as both operators
A schematic presentation of the gal operon, including positioning of both promoters, both transcription initiation sites, as well as both operators

Regulatory Genes

Regulatory genes code for gene products that alter the expression of other genes, usually either inhibiting or activating expression. There are three regulatory genes that are not part of the gal operon but associate with it in a regulatory fashion:

  • galR codes for the gal repressor, which regulates both P1 and P2
  • galS codes for the gal isorepressor which is similar to the gal repressor and also regulates both P1 and P2
  • crp codes for cyclic AMP receptor protein that when bound together with cAMP, forms a cAMP-CRP complex which stimulates transcription at P1 and represses transcription at P2[3]

Regulation

The cAMP-crp Complex

The cAMP-crp complex binds upstream of the two promoters and causes preferential transcription starting at P1 rather than from both P1 and P2. In the absence of the cAMP-crp complex, transcription initiation of the galoperon is approximately proportionate between the sites. However, when the cAMP-crp complex is present, transcription initiation predominantly begins at P1 (>95% of transcription intiations)[5][6]. The cAMP-crp complex alters transcription initiation preference by (1) increasing RNA polymerase affinity to P1, thereby increasing RNA polymerase binding to P1, and (2) elevating the rate of conversion of the closed DNA complex to an open DNA complex that is stable and ready to be transcribed[7][6].

The galR Protein

The galR protein works to repress both P1 and P2 by forming a repressosome. GalR proteins come together to form dimers, of which each monomer component binds simultaneously to either OE or OI. This interaction of dimers to operators is required for full repression and alters DNA conformation by forming a loop- the repressosome. [8] This repressosome is stabilized by the binding of an HU protein to the HBS (HU binding site) located between the promoters and OI. The galR protein, however, if binding to OE alone, stimulates transcription initiation from P2 and partially represses transcription initiation from P1[8].

The galS Protein

The galS protein is a member of the GalR-LacI family, the same family in which the GalR protein belongs. GalS is 53% identical and 85% similar with GalR. As a result, GalS represses transcription of the operon in an analagous manner to GalR, forming the repressosome and inducing HU protein binding. However, it represses transcription much less efficiently in comparison to GalR[1].



  1. ^ a b c d Weickert, Michael J.; Adhya, Sankar (1993). "The galactose regulon of Escherichia coli". Molecular Microbiology. 10 (2): 245-251. doi:10.1111/j.1365-2958.1993.tb01950.x.
  2. ^ a b {cite web|title=Galactose Operon|url=http://www.spektrum.de/lexikon/biologie/galactose-operon/26298%7Cwebsite=Spektrum der Wissenschaft|publisher=Spektrum Akademischer Verlag|accessdate=11 November 2015}}
  3. ^ a b c Irani, Meher H.; Orosz, Laszlo; Adhya, Sankar (March 1983). "A control element within a structural gene: The gal operon of Escherichia coli". Cell. 32 (3): 783-788. doi:10.1016/0092-8674(83)90064-8.
  4. ^ a b Ji, Sang Chun; Jeon, Heung Jin; Yun, Sang Hoon; Lee, Hee Jung; Lim, Heon M. (July 2010). "Quantification of the galactose-operon mRNAs 5 bases different in their 5'-ends" (PDF). BMB Reports. 43 (7): 474-479. PMID 20663408.
  5. ^ Cite error: The named reference Weickert&Adhya) was invoked but never defined (see the help page).
  6. ^ a b Goodrich, James A.; McClure, William R. (1992). "Regulation of open complex formation at the Escherichia coli galactose operon promoters: Simultaneous interaction of RNA polymerase, gal repressor and CAP/cAMP". Journal of Molecular Biology. 224 (1): 15-29. doi:10.1016/0022-2836(92)90573-3.
  7. ^ Cite error: The named reference Weickert&Adhya" was invoked but never defined (see the help page).
  8. ^ a b Semsey, Szabolcs; Virnik, Konstantin; Adhya, Sankar (2006). "Three-stage Regulation of the Amphibolic gal Operon: From Repressosome to GalR-free DNA" (PDF). Journal of Molecular Biology. 358: 355-363. doi:10.1016/j.jmb.2006.02.022.
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