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Sterol uptake control protein 2

 UPC2_CANAL              Reviewed;         712 AA.
Q59QC7; A0A1D8PEC3;
01-APR-2015, integrated into UniProtKB/Swiss-Prot.
26-APR-2005, sequence version 1.
25-OCT-2017, entry version 89.
RecName: Full=Sterol uptake control protein 2;
Name=UPC2 {ECO:0000303|PubMed:15590814}; Synonyms=ECM22;
OrderedLocusNames=CAALFM_C108460CA; ORFNames=CaO19.391, CaO19.8021;
Candida albicans (strain SC5314 / ATCC MYA-2876) (Yeast).
Eukaryota; Fungi; Dikarya; Ascomycota; Saccharomycotina;
Saccharomycetes; Saccharomycetales; Debaryomycetaceae;
Candida/Lodderomyces clade; Candida.
NCBI_TaxID=237561;
[1]
NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
STRAIN=SC5314 / ATCC MYA-2876;
PubMed=15123810; DOI=10.1073/pnas.0401648101;
Jones T., Federspiel N.A., Chibana H., Dungan J., Kalman S.,
Magee B.B., Newport G., Thorstenson Y.R., Agabian N., Magee P.T.,
Davis R.W., Scherer S.;
"The diploid genome sequence of Candida albicans.";
Proc. Natl. Acad. Sci. U.S.A. 101:7329-7334(2004).
[2]
GENOME REANNOTATION.
STRAIN=SC5314 / ATCC MYA-2876;
PubMed=17419877; DOI=10.1186/gb-2007-8-4-r52;
van het Hoog M., Rast T.J., Martchenko M., Grindle S., Dignard D.,
Hogues H., Cuomo C., Berriman M., Scherer S., Magee B.B., Whiteway M.,
Chibana H., Nantel A., Magee P.T.;
"Assembly of the Candida albicans genome into sixteen supercontigs
aligned on the eight chromosomes.";
Genome Biol. 8:RESEARCH52.1-RESEARCH52.12(2007).
[3]
NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA], AND GENOME
REANNOTATION.
STRAIN=SC5314 / ATCC MYA-2876;
PubMed=24025428; DOI=10.1186/gb-2013-14-9-r97;
Muzzey D., Schwartz K., Weissman J.S., Sherlock G.;
"Assembly of a phased diploid Candida albicans genome facilitates
allele-specific measurements and provides a simple model for repeat
and indel structure.";
Genome Biol. 14:RESEARCH97.1-RESEARCH97.14(2013).
[4]
DISRUPTION PHENOTYPE, AND FUNCTION.
PubMed=15590814; DOI=10.1128/EC.3.6.1391-1397.2004;
Silver P.M., Oliver B.G., White T.C.;
"Role of Candida albicans transcription factor Upc2p in drug
resistance and sterol metabolism.";
Eukaryot. Cell 3:1391-1397(2004).
[5]
DISRUPTION PHENOTYPE, FUNCTION, AND DNA-BINDING.
PubMed=15855491; DOI=10.1128/AAC.49.5.1745-1752.2005;
MacPherson S., Akache B., Weber S., De Deken X., Raymond M.,
Turcotte B.;
"Candida albicans zinc cluster protein Upc2p confers resistance to
antifungal drugs and is an activator of ergosterol biosynthetic
genes.";
Antimicrob. Agents Chemother. 49:1745-1752(2005).
[6]
INDUCTION.
PubMed=16039996; DOI=10.1016/j.bbrc.2005.07.018;
Singh V., Sinha I., Sadhale P.P.;
"Global analysis of altered gene expression during morphogenesis of
Candida albicans in vitro.";
Biochem. Biophys. Res. Commun. 334:1149-1158(2005).
[7]
IDENTIFICATION.
PubMed=18629206; DOI=10.1002/cfg.492;
Maicas S., Moreno I., Nieto A., Gomez M., Sentandreu R., Valentin E.;
"In silico analysis for transcription factors with Zn(II)(2)C(6)
binuclear cluster DNA-binding domains in Candida albicans.";
Comp. Funct. Genomics 6:345-356(2005).
[8]
INDUCTION.
PubMed=15820985; DOI=10.1093/jac/dki088;
Copping V.M.S., Barelle C.J., Hube B., Gow N.A.R., Brown A.J.P.,
Odds F.C.;
"Exposure of Candida albicans to antifungal agents affects expression
of SAP2 and SAP9 secreted proteinase genes.";
J. Antimicrob. Chemother. 55:645-654(2005).
[9]
FUNCTION, AND MUTAGENESIS OF GLY-648.
PubMed=18487346; DOI=10.1128/EC.00103-08;
Dunkel N., Liu T.T., Barker K.S., Homayouni R., Morschhauser J.,
Rogers P.D.;
"A gain-of-function mutation in the transcription factor Upc2p causes
upregulation of ergosterol biosynthesis genes and increased
fluconazole resistance in a clinical Candida albicans isolate.";
Eukaryot. Cell 7:1180-1190(2008).
[10]
INDUCTION.
PubMed=18757808; DOI=10.1099/mic.0.2008/017475-0;
Hoot S.J., Oliver B.G., White T.C.;
"Candida albicans UPC2 is transcriptionally induced in response to
antifungal drugs and anaerobicity through Upc2p-dependent and
-independent mechanisms.";
Microbiology 154:2748-2756(2008).
[11]
FUNCTION, AND MUTAGENESIS OF ALA-643.
PubMed=19884367; DOI=10.1128/AAC.01102-09;
Heilmann C.J., Schneider S., Barker K.S., Rogers P.D.,
Morschhaeuser J.;
"An A643T mutation in the transcription factor Upc2p causes
constitutive ERG11 upregulation and increased fluconazole resistance
in Candida albicans.";
Antimicrob. Agents Chemother. 54:353-359(2010).
[12]
INDUCTION, AND FUNCTION.
PubMed=20656915; DOI=10.1128/EC.00130-10;
Hoot S.J., Brown R.P., Oliver B.G., White T.C.;
"The UPC2 promoter in Candida albicans contains two cis-acting
elements that bind directly to Upc2p, resulting in transcriptional
autoregulation.";
Eukaryot. Cell 9:1354-1362(2010).
[13]
FUNCTION, AND MUTAGENESIS OF ALA-643.
PubMed=21078937; DOI=10.1128/AAC.00995-10;
Hoot S.J., Smith A.R., Brown R.P., White T.C.;
"An A643V amino acid substitution in Upc2p contributes to azole
resistance in well-characterized clinical isolates of Candida
albicans.";
Antimicrob. Agents Chemother. 55:940-942(2011).
[14]
FUNCTION, AND DISRUPTION PHENOTYPE.
PubMed=22006003; DOI=10.1128/AAC.00574-11;
Dhamgaye S., Devaux F., Manoharlal R., Vandeputte P., Shah A.H.,
Singh A., Blugeon C., Sanglard D., Prasad R.;
"In vitro effect of malachite green on Candida albicans involves
multiple pathways and transcriptional regulators UPC2 and STP2.";
Antimicrob. Agents Chemother. 56:495-506(2012).
[15]
FUNCTION, AND MUTAGENESIS OF TRP-478; TYR-642; ALA-643; ALA-646 AND
GLY-648.
PubMed=22923048; DOI=10.1128/EC.00215-12;
Flowers S.A., Barker K.S., Berkow E.L., Toner G., Chadwick S.G.,
Gygax S.E., Morschhauser J., Rogers P.D.;
"Gain-of-function mutations in UPC2 are a frequent cause of ERG11
upregulation in azole-resistant clinical isolates of Candida
albicans.";
Eukaryot. Cell 11:1289-1299(2012).
[16]
FUNCTION.
PubMed=25385116; DOI=10.1128/AAC.04448-14;
Schneider S., Morschhaeuser J.;
"Induction of Candida albicans drug resistance genes by hybrid zinc
cluster transcription factors.";
Antimicrob. Agents Chemother. 59:558-569(2015).
[17]
FUNCTION, DNA-BINDING, AND BIOPHYSICOCHEMICAL PROPERTIES.
PubMed=24145546; DOI=10.1128/AAC.01677-13;
Gallo-Ebert C., Donigan M., Stroke I.L., Swanson R.N., Manners M.T.,
Francisco J., Toner G., Gallagher D., Huang C.Y., Gygax S.E., Webb M.,
Nickels J.T. Jr.;
"Novel antifungal drug discovery based on targeting pathways
regulating the fungus-conserved Upc2 transcription factor.";
Antimicrob. Agents Chemother. 58:258-266(2014).
[18]
FUNCTION.
PubMed=24243794; DOI=10.1128/EC.00245-13;
Lohberger A., Coste A.T., Sanglard D.;
"Distinct roles of Candida albicans drug resistance transcription
factors TAC1, MRR1, and UPC2 in virulence.";
Eukaryot. Cell 13:127-142(2014).
[19]
FUNCTION, AND DISRUPTION PHENOTYPE.
PubMed=24376002; DOI=10.1128/EC.00336-13;
Wellington M., Koselny K., Sutterwala F.S., Krysan D.J.;
"Candida albicans triggers NLRP3-mediated pyroptosis in macrophages.";
Eukaryot. Cell 13:329-340(2014).
[20]
FUNCTION, AND DISRUPTION PHENOTYPE.
PubMed=24659578; DOI=10.1128/EC.00221-13;
Vasicek E.M., Berkow E.L., Flowers S.A., Barker K.S., Rogers P.D.;
"UPC2 is universally essential for azole antifungal resistance in
Candida albicans.";
Eukaryot. Cell 13:933-946(2014).
[21]
FUNCTION.
PubMed=25084864; DOI=10.1128/EC.00096-14;
Van Hauwenhuyse F., Fiori A., Van Dijck P.;
"Ascorbic acid inhibition of Candida albicans Hsp90-mediated
morphogenesis occurs via the transcriptional regulator Upc2.";
Eukaryot. Cell 13:1278-1289(2014).
-!- FUNCTION: Transcription factor involved in the regulation of
ergosterol biosynthetic genes such as ERG2 and ERG11 through
direct binding to sterol response elements (SREs) in the
promoters. Binds also to its own promoter on 2 cis-acting elements
to promote autoregulation. Regulates sterol uptake across the
plasma membrane. Acts as a major regulator of ascorbic acid-
induced response. Plays a role in the triggering of pyroptosis, an
inflammasome-mediated programmed cell death pathway in
macrophages, allowing macrophages escaping.
{ECO:0000269|PubMed:15590814, ECO:0000269|PubMed:15855491,
ECO:0000269|PubMed:18487346, ECO:0000269|PubMed:19884367,
ECO:0000269|PubMed:20656915, ECO:0000269|PubMed:21078937,
ECO:0000269|PubMed:22006003, ECO:0000269|PubMed:22923048,
ECO:0000269|PubMed:24145546, ECO:0000269|PubMed:24243794,
ECO:0000269|PubMed:24376002, ECO:0000269|PubMed:24659578,
ECO:0000269|PubMed:25084864, ECO:0000269|PubMed:25385116}.
-!- BIOPHYSICOCHEMICAL PROPERTIES:
Kinetic parameters:
KM=50 nM for DNA sterol response element (SRE)
{ECO:0000269|PubMed:24145546};
-!- SUBCELLULAR LOCATION: Nucleus {ECO:0000255|PROSITE-
ProRule:PRU00227}.
-!- INDUCTION: Expression is up-regulated by ergosterol depletion, by
azoles, and in anaerobic conditions. Transcript is repressed
during co-incubated with macrophages, a condition that induces
filamentous growth. Promotes positive auto-regulation through
binding to its own promoter. {ECO:0000269|PubMed:15820985,
ECO:0000269|PubMed:16039996, ECO:0000269|PubMed:18757808,
ECO:0000269|PubMed:20656915}.
-!- DISRUPTION PHENOTYPE: Shows increased suceptibility to the azole
drugs ketoconazole, itraconazole, and fluconazole; drugs that act
on ergosterol biosynthesis such as terbinafine, fenpropimorph, and
lovastatin; as well as malachite green. Leads to lower ergosterol
levels. Decreases pyroptosis but has little effect on
filamentation in the macrophage. {ECO:0000269|PubMed:15590814,
ECO:0000269|PubMed:15855491, ECO:0000269|PubMed:22006003,
ECO:0000269|PubMed:24376002, ECO:0000269|PubMed:24659578}.
-!- MISCELLANEOUS: Gain-of-function mutations in UPC2 are more
prevalent among clinical isolates than previously thought and make
a significant contribution to azole antifungal resistance.
{ECO:0000269|PubMed:22923048}.
-----------------------------------------------------------------------
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-----------------------------------------------------------------------
EMBL; CP017623; AOW26484.1; -; Genomic_DNA.
RefSeq; XP_711879.1; XM_706786.2.
ProteinModelPortal; Q59QC7; -.
SMR; Q59QC7; -.
PRIDE; Q59QC7; -.
EnsemblFungi; KHC83595; KHC83595; W5Q_00814.
EnsemblFungi; KHC89031; KHC89031; I503_00822.
GeneID; 3646489; -.
KEGG; cal:CAALFM_C108460CA; -.
CGD; CAL0000191743; UPC2.
OrthoDB; EOG092C3B2O; -.
PRO; PR:Q59QC7; -.
Proteomes; UP000000559; Chromosome 1.
GO; GO:0005634; C:nucleus; IEA:UniProtKB-SubCell.
GO; GO:0003677; F:DNA binding; IEA:UniProtKB-KW.
GO; GO:0000982; F:transcription factor activity, RNA polymerase II core promoter proximal region sequence-specific binding; IDA:CGD.
GO; GO:0008270; F:zinc ion binding; IEA:InterPro.
GO; GO:0035690; P:cellular response to drug; IMP:CGD.
GO; GO:0008204; P:ergosterol metabolic process; IMP:CGD.
GO; GO:0045892; P:negative regulation of transcription, DNA-templated; IMP:CGD.
GO; GO:0009405; P:pathogenesis; IEA:UniProtKB-KW.
GO; GO:0045944; P:positive regulation of transcription from RNA polymerase II promoter; IDA:CGD.
GO; GO:0016125; P:sterol metabolic process; IMP:CGD.
GO; GO:0006351; P:transcription, DNA-templated; IEA:UniProtKB-KW.
CDD; cd00067; GAL4; 1.
Gene3D; 4.10.240.10; -; 1.
InterPro; IPR021858; Fun_TF.
InterPro; IPR001138; Zn2-C6_fun-type_DNA-bd.
InterPro; IPR036864; Zn2-C6_fun-type_DNA-bd_sf.
Pfam; PF11951; Fungal_trans_2; 1.
Pfam; PF00172; Zn_clus; 1.
SMART; SM00066; GAL4; 1.
SUPFAM; SSF57701; SSF57701; 1.
PROSITE; PS00463; ZN2_CY6_FUNGAL_1; 1.
PROSITE; PS50048; ZN2_CY6_FUNGAL_2; 1.
1: Evidence at protein level;
Activator; Complete proteome; DNA-binding; Metal-binding; Nucleus;
Reference proteome; Transcription; Transcription regulation;
Virulence; Zinc.
CHAIN 1 712 Sterol uptake control protein 2.
/FTId=PRO_0000431801.
DNA_BIND 54 81 Zn(2)-C6 fungal-type.
{ECO:0000255|PROSITE-ProRule:PRU00227}.
MUTAGEN 478 478 W->C: Leads to increased ERG11
expression, increased cellular
ergosterol, and decreased susceptibility
to fluconazole.
{ECO:0000269|PubMed:22923048}.
MUTAGEN 642 642 Y->F: Leads to increased ERG11
expression, increased cellular
ergosterol, and decreased susceptibility
to fluconazole.
{ECO:0000269|PubMed:22923048}.
MUTAGEN 643 643 A->T,V: Leads to increased ERG11
expression, increased cellular
ergosterol, and decreased susceptibility
to fluconazole.
{ECO:0000269|PubMed:22923048}.
MUTAGEN 646 646 A->V: Leads to increased ERG11
expression, increased cellular
ergosterol, and decreased susceptibility
to fluconazole.
{ECO:0000269|PubMed:22923048}.
MUTAGEN 648 648 G->D,S: Leads to increased ERG11
expression, increased cellular
ergosterol, and decreased susceptibility
to fluconazole.
{ECO:0000269|PubMed:22923048}.
SEQUENCE 712 AA; 77035 MW; F6AED0926B01AF61 CRC64;
MMMTVKQESP NSTLNTSEFS SDENLKTNNS EPPKKVSKSS TGKRKYHQKS RNGCSTCKKR
RVKCDEQRPV CGNCTKLKLD CGYLHEPLEN ILNTKKDIAN NEPPSKKRKR KVSTVSAASD
SESTTQQATP SLTPSPNHSQ DIKTQPVIPP TNPLSALSSG LLSAGNLNNL NVAHLVNNLS
GLGDLSNLAS LGNLASLSNL ASLAQLPIDL SNLGSLLDSP AASNIAASFL GSAAATTVPP
TTNSEFKESN QRKSQTQMPP QPTVPITSMG AATTTSSHQQ ANMPSRSKPQ PETLQSSIPA
TTSGSPGMSY PGCPSNSDPF GRSSDKSLPN ISPNMSIPAN PLSDPLTQGM RSNLNMLDLK
LMFHYTSVVA NTITGAGISD TNIWNCDIPK LAFEHPFLMH SILAFSATHL SRTEKGLDQC
VTCHRGDALR LLREAVLNIN ADNTDALVAS ALILIMDSLA NASFPSSTSP KSLPASAWIF
HVKGAATILT AVWPLTEASR FYKFISVDLG DLGDIINQGV NMNKSKGIDR ENSAYYTDLE
CHDADIADLF PVLLDSPYLI TLAYLNKLHK ERYKSDFILR IFAFPALLDK QFMGLLMSGD
VKAMRIMRSY YKLLRSFTTE MKDKVWFLEG VSQVLPVNVE EYAGGAGGMH MMMDFLGGGP
AIVDDNEIDA EITKFDPSGT LTNKLIDTDN LPSVLTSNLD LMQGDNGFMN MK


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