A3AEM_PHYIT
ID A3AEM_PHYIT Reviewed; 147 AA.
AC D0NPN8;
DT 08-MAY-2019, integrated into UniProtKB/Swiss-Prot.
DT 15-DEC-2009, sequence version 1.
DT 03-AUG-2022, entry version 39.
DE RecName: Full=RxLR effector protein Avr3a {ECO:0000303|PubMed:16965554};
DE AltName: Full=Avirulence protein 3a {ECO:0000303|PubMed:15894622};
DE Flags: Precursor;
GN Name=Avr3a {ECO:0000303|PubMed:15894622};
GN Synonyms=Avr3a(EM) {ECO:0000303|PubMed:19245321},
GN Pex147 {ECO:0000303|PubMed:15894622}, PexRD7 {ECO:0000303|PubMed:19794118};
GN ORFNames=PITG_14371;
OS Phytophthora infestans (strain T30-4) (Potato late blight agent).
OC Eukaryota; Sar; Stramenopiles; Oomycota; Peronosporales; Peronosporaceae;
OC Phytophthora.
OX NCBI_TaxID=403677;
RN [1]
RP NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA], AND INDUCTION.
RC STRAIN=T30-4;
RX PubMed=19741609; DOI=10.1038/nature08358;
RG The Broad Institute Genome Sequencing Platform;
RA Haas B.J., Kamoun S., Zody M.C., Jiang R.H., Handsaker R.E., Cano L.M.,
RA Grabherr M., Kodira C.D., Raffaele S., Torto-Alalibo T., Bozkurt T.O.,
RA Ah-Fong A.M., Alvarado L., Anderson V.L., Armstrong M.R., Avrova A.,
RA Baxter L., Beynon J., Boevink P.C., Bollmann S.R., Bos J.I., Bulone V.,
RA Cai G., Cakir C., Carrington J.C., Chawner M., Conti L., Costanzo S.,
RA Ewan R., Fahlgren N., Fischbach M.A., Fugelstad J., Gilroy E.M., Gnerre S.,
RA Green P.J., Grenville-Briggs L.J., Griffith J., Grunwald N.J., Horn K.,
RA Horner N.R., Hu C.H., Huitema E., Jeong D.H., Jones A.M., Jones J.D.,
RA Jones R.W., Karlsson E.K., Kunjeti S.G., Lamour K., Liu Z., Ma L.,
RA Maclean D., Chibucos M.C., McDonald H., McWalters J., Meijer H.J.,
RA Morgan W., Morris P.F., Munro C.A., O'Neill K., Ospina-Giraldo M.,
RA Pinzon A., Pritchard L., Ramsahoye B., Ren Q., Restrepo S., Roy S.,
RA Sadanandom A., Savidor A., Schornack S., Schwartz D.C., Schumann U.D.,
RA Schwessinger B., Seyer L., Sharpe T., Silvar C., Song J., Studholme D.J.,
RA Sykes S., Thines M., van de Vondervoort P.J., Phuntumart V., Wawra S.,
RA Weide R., Win J., Young C., Zhou S., Fry W., Meyers B.C., van West P.,
RA Ristaino J., Govers F., Birch P.R., Whisson S.C., Judelson H.S.,
RA Nusbaum C.;
RT "Genome sequence and analysis of the Irish potato famine pathogen
RT Phytophthora infestans.";
RL Nature 461:393-398(2009).
RN [2]
RP FUNCTION, AND SUBCELLULAR LOCATION.
RX PubMed=15894622; DOI=10.1073/pnas.0500113102;
RA Armstrong M.R., Whisson S.C., Pritchard L., Bos J.I., Venter E.,
RA Avrova A.O., Rehmany A.P., Boehme U., Brooks K., Cherevach I., Hamlin N.,
RA White B., Fraser A., Lord A., Quail M.A., Churcher C., Hall N.,
RA Berriman M., Huang S., Kamoun S., Beynon J.L., Birch P.R.;
RT "An ancestral oomycete locus contains late blight avirulence gene Avr3a,
RT encoding a protein that is recognized in the host cytoplasm.";
RL Proc. Natl. Acad. Sci. U.S.A. 102:7766-7771(2005).
RN [3]
RP FUNCTION.
RX PubMed=16965554; DOI=10.1111/j.1365-313x.2006.02866.x;
RA Bos J.I., Kanneganti T.D., Young C., Cakir C., Huitema E., Win J.,
RA Armstrong M.R., Birch P.R., Kamoun S.;
RT "The C-terminal half of Phytophthora infestans RXLR effector AVR3a is
RT sufficient to trigger R3a-mediated hypersensitivity and suppress INF1-
RT induced cell death in Nicotiana benthamiana.";
RL Plant J. 48:165-176(2006).
RN [4]
RP DOMAIN, AND SUBCELLULAR LOCATION.
RX PubMed=17914356; DOI=10.1038/nature06203;
RA Whisson S.C., Boevink P.C., Moleleki L., Avrova A.O., Morales J.G.,
RA Gilroy E.M., Armstrong M.R., Grouffaud S., van West P., Chapman S.,
RA Hein I., Toth I.K., Pritchard L., Birch P.R.;
RT "A translocation signal for delivery of oomycete effector proteins into
RT host plant cells.";
RL Nature 450:115-118(2007).
RN [5]
RP FUNCTION.
RX PubMed=19794118; DOI=10.1105/tpc.109.068247;
RA Oh S.K., Young C., Lee M., Oliva R., Bozkurt T.O., Cano L.M., Win J.,
RA Bos J.I., Liu H.Y., van Damme M., Morgan W., Choi D., Van der Vossen E.A.,
RA Vleeshouwers V.G., Kamoun S.;
RT "In planta expression screens of Phytophthora infestans RXLR effectors
RT reveal diverse phenotypes, including activation of the Solanum
RT bulbocastanum disease resistance protein Rpi-blb2.";
RL Plant Cell 21:2928-2947(2009).
RN [6]
RP FUNCTION.
RX PubMed=19245321; DOI=10.1094/mpmi-22-3-0269;
RA Bos J.I., Chaparro-Garcia A., Quesada-Ocampo L.M., McSpadden Gardener B.B.,
RA Kamoun S.;
RT "Distinct amino acids of the Phytophthora infestans effector AVR3a
RT condition activation of R3a hypersensitivity and suppression of cell
RT death.";
RL Mol. Plant Microbe Interact. 22:269-281(2009).
RN [7]
RP FUNCTION, AND INTERACTION WITH HOST E3 LIGASE CMPG1.
RX PubMed=20457921; DOI=10.1073/pnas.0914408107;
RA Bos J.I., Armstrong M.R., Gilroy E.M., Boevink P.C., Hein I., Taylor R.M.,
RA Zhendong T., Engelhardt S., Vetukuri R.R., Harrower B., Dixelius C.,
RA Bryan G., Sadanandom A., Whisson S.C., Kamoun S., Birch P.R.;
RT "Phytophthora infestans effector AVR3a is essential for virulence and
RT manipulates plant immunity by stabilizing host E3 ligase CMPG1.";
RL Proc. Natl. Acad. Sci. U.S.A. 107:9909-9914(2010).
RN [8]
RP FUNCTION.
RX PubMed=21348873; DOI=10.1111/j.1469-8137.2011.03643.x;
RA Gilroy E.M., Taylor R.M., Hein I., Boevink P., Sadanandom A., Birch P.R.;
RT "CMPG1-dependent cell death follows perception of diverse pathogen
RT elicitors at the host plasma membrane and is suppressed by Phytophthora
RT infestans RXLR effector AVR3a.";
RL New Phytol. 190:653-666(2011).
RN [9]
RP DOMAIN, AND SUBUNIT.
RX PubMed=22977236; DOI=10.1074/jbc.m112.395129;
RA Wawra S., Agacan M., Boddey J.A., Davidson I., Gachon C.M., Zanda M.,
RA Grouffaud S., Whisson S.C., Birch P.R., Porter A.J., van West P.;
RT "Avirulence protein 3a (AVR3a) from the potato pathogen Phytophthora
RT infestans forms homodimers through its predicted translocation region and
RT does not specifically bind phospholipids.";
RL J. Biol. Chem. 287:38101-38109(2012).
RN [10]
RP FUNCTION, AND INTERACTION WITH DRP2.
RX PubMed=26348328; DOI=10.1371/journal.pone.0137071;
RA Chaparro-Garcia A., Schwizer S., Sklenar J., Yoshida K., Petre B.,
RA Bos J.I., Schornack S., Jones A.M., Bozkurt T.O., Kamoun S.;
RT "Phytophthora infestans RXLR-WY Effector AVR3a Associates with Dynamin-
RT Related Protein 2 Required for Endocytosis of the Plant Pattern Recognition
RT Receptor FLS2.";
RL PLoS ONE 10:E0137071-E0137071(2015).
RN [11]
RP INDUCTION.
RX PubMed=28228125; DOI=10.1186/s12864-017-3585-x;
RA Ah-Fong A.M., Kim K.S., Judelson H.S.;
RT "RNA-seq of life stages of the oomycete Phytophthora infestans reveals
RT dynamic changes in metabolic, signal transduction, and pathogenesis genes
RT and a major role for calcium signaling in development.";
RL BMC Genomics 18:198-198(2017).
RN [12]
RP CLEAVAGE, ACETYLATION AT LYS-48, SUBUNIT, AND GLYCOSYLATION.
RX PubMed=28522546; DOI=10.1105/tpc.16.00552;
RA Wawra S., Trusch F., Matena A., Apostolakis K., Linne U., Zhukov I.,
RA Stanek J., Kozminski W., Davidson I., Secombes C.J., Bayer P., van West P.;
RT "The RxLR motif of the host targeting effector AVR3a of Phytophthora
RT infestans is cleaved before secretion.";
RL Plant Cell 29:1184-1195(2017).
RN [13]
RP INDUCTION, AND SUBCELLULAR LOCATION.
RX PubMed=29312401; DOI=10.3389/fpls.2017.02155;
RA Yin J., Gu B., Huang G., Tian Y., Quan J., Lindqvist-Kreuze H., Shan W.;
RT "Conserved RXLR effector genes of Phytophthora infestans expressed at the
RT early stage of potato infection are suppressive to host defense.";
RL Front. Plant Sci. 8:2155-2155(2017).
CC -!- FUNCTION: Multifunctional effector that can suppress host BAK1/SERK3-
CC mediated immunity through at least two different pathways
CC (PubMed:19794118, PubMed:20457921, PubMed:21348873, PubMed:26348328).
CC Manipulates plant immunity by targeting and stabilizing host E3 ligase
CC CMPG1. By preventing the normal 26S proteasome-dependent degradation of
CC potato CMPG1, and thus potentially of its protein substrates in the
CC host cell, Avr3a further abolishes host cell death during the
CC biotrophic phase of infection (PubMed:19794118, PubMed:20457921,
CC PubMed:21348873). Associates with and affects also the function of the
CC dynamin-related protein 2 (DRP2), a plant GTPase involved in immune
CC receptor-mediated endocytosis (PubMed:26348328). The Avr3a(EM) form
CC evades recognition by R3a, thus does not trigger R3a-mediated
CC hypersensitivity and does not suppress INF1-induced cell death
CC (PubMed:15894622, PubMed:16965554, PubMed:19245321).
CC {ECO:0000269|PubMed:15894622, ECO:0000269|PubMed:16965554,
CC ECO:0000269|PubMed:19245321, ECO:0000269|PubMed:19794118,
CC ECO:0000269|PubMed:20457921, ECO:0000269|PubMed:21348873,
CC ECO:0000269|PubMed:26348328}.
CC -!- SUBUNIT: Forms homodimers via the RxLR-dEER motif (PubMed:22977236).
CC Interacts with host E3 ligase CMPG1 (PubMed:20457921). Interacts with
CC host DRP2 (PubMed:26348328). {ECO:0000269|PubMed:20457921,
CC ECO:0000269|PubMed:22977236, ECO:0000269|PubMed:26348328}.
CC -!- SUBCELLULAR LOCATION: Secreted {ECO:0000269|PubMed:15894622,
CC ECO:0000269|PubMed:16965554, ECO:0000269|PubMed:17914356,
CC ECO:0000269|PubMed:29312401}. Host cytoplasm
CC {ECO:0000269|PubMed:15894622, ECO:0000269|PubMed:16965554,
CC ECO:0000269|PubMed:17914356, ECO:0000269|PubMed:29312401}.
CC -!- INDUCTION: Expression is induced during host plant infection.
CC {ECO:0000269|PubMed:19741609, ECO:0000269|PubMed:28228125,
CC ECO:0000269|PubMed:29312401}.
CC -!- DOMAIN: The RxLR-dEER motif is required for the delivery of the
CC effector to the host cell cytoplasm but does not bind
CC phosphatidylinositol monophosphates (PubMed:17914356, PubMed:22977236).
CC The motif is cleaved after the RxLR sequence (PubMed:28522546). The
CC RxLR motif (residues 44 to 47) plays a crucial role in the
CC intracellular processing before secretion (PubMed:28522546). The Glu-
CC rich part localized just after the cleavage site (residues 48 to 59) is
CC required for homodimerization (PubMed:28522546).
CC {ECO:0000269|PubMed:17914356, ECO:0000269|PubMed:22977236,
CC ECO:0000269|PubMed:28522546}.
CC -!- DOMAIN: The conserved, positively charged effector domain (residues 77
CC to 147), rather than the RxLR-dEER motif, is required for binding to
CC phosphatidylinositol monophosphates (PIPs). PIP binding is necessary
CC for accumulation of CMPG1 and Avr3a in host plants.
CC {ECO:0000250|UniProtKB:E2DWQ7}.
CC -!- PTM: Proteolytically cleaved. The cleavage site directly after the RxLR
CC sequence and the high conservation among other effector proteins
CC suggest that the RxLR motif might play a crucial role in the
CC intracellular processing before secretion.
CC {ECO:0000269|PubMed:28522546}.
CC -!- PTM: glycosylated. {ECO:0000269|PubMed:28522546}.
CC -!- PTM: N-acetylated at Lys-48 after cleavage.
CC {ECO:0000269|PubMed:28522546}.
CC -!- MISCELLANEOUS: The AVR3a protein of Phytophthora infestans is a
CC polymorphic member of the RXLR class of cytoplasmic effectors with dual
CC functions. AVR3a(KI) but not AVR3a(EM) activates innate immunity
CC triggered by the potato resistance protein R3a and is a strong
CC suppressor of the cell-death response induced by INF1 elicitin, a
CC secreted P.infestans protein that has features of pathogen-associated
CC molecular patterns. The 2 polymorphic residues between Avr3A(KI) and
CC Avr3a(EM) are localized at positions 80 and 103, respectively.
CC {ECO:0000269|PubMed:15894622, ECO:0000269|PubMed:16965554}.
CC -!- SIMILARITY: Belongs to the RxLR effector family. {ECO:0000305}.
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DR EMBL; DS028151; EEY62600.1; -; Genomic_DNA.
DR RefSeq; XP_002898842.1; XM_002898796.1.
DR AlphaFoldDB; D0NPN8; -.
DR SMR; D0NPN8; -.
DR STRING; 4787.PITG_14371T0; -.
DR iPTMnet; D0NPN8; -.
DR EnsemblProtists; PITG_14371T0; PITG_14371T0; PITG_14371.
DR GeneID; 9479695; -.
DR KEGG; pif:PITG_14371; -.
DR VEuPathDB; FungiDB:PITG_14371; -.
DR eggNOG; ENOG502RGQI; Eukaryota.
DR HOGENOM; CLU_1762379_0_0_1; -.
DR OMA; VMNDSPK; -.
DR OrthoDB; 1629495at2759; -.
DR Proteomes; UP000006643; Partially assembled WGS sequence.
DR GO; GO:0005576; C:extracellular region; IEA:UniProtKB-SubCell.
DR GO; GO:0030430; C:host cell cytoplasm; IEA:UniProtKB-SubCell.
DR InterPro; IPR031825; RXLR.
DR Pfam; PF16810; RXLR; 1.
PE 1: Evidence at protein level;
KW Acetylation; Glycoprotein; Host cytoplasm; Reference proteome; Secreted;
KW Signal; Virulence.
FT SIGNAL 1..21
FT /evidence="ECO:0000255"
FT CHAIN 22..147
FT /note="RxLR effector protein Avr3a"
FT /id="PRO_5003012271"
FT REGION 77..147
FT /note="Effector domain"
FT /evidence="ECO:0000250|UniProtKB:E2DWQ7"
FT MOTIF 44..59
FT /note="RxLR-dEER"
FT /evidence="ECO:0000305|PubMed:17914356"
FT SITE 47..48
FT /note="Cleavage"
FT /evidence="ECO:0000269|PubMed:28522546"
FT MOD_RES 48
FT /note="N6-acetyllysine"
FT /evidence="ECO:0000269|PubMed:28522546"
SQ SEQUENCE 147 AA; 16661 MW; 24CD19EADC7B8B39 CRC64;
MRLAIMLSAT AVAINFATSS AIDQTKVLVY GTPAHYIHDS AGRRLLRKNE ENEETSEERA
PNFNLANLNE EMFNVAALTE RADAKKLAKQ LMGNDKLADA AYMWWQHNRV TLDQIDTFLK
LASRKTQGAK YNQIYNSYMM HLGLTGY
