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A3AEM_PHYIN
ID   A3AEM_PHYIN             Reviewed;         147 AA.
AC   Q572D3;
DT   08-MAY-2019, integrated into UniProtKB/Swiss-Prot.
DT   10-MAY-2005, sequence version 1.
DT   25-MAY-2022, entry version 51.
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:19794118}, PexRD7 {ECO:0000303|PubMed:19794118};
GN   ORFNames=PI35.0050, PI49.0530;
OS   Phytophthora infestans (Potato late blight agent) (Botrytis infestans).
OC   Eukaryota; Sar; Stramenopiles; Oomycota; Peronosporales; Peronosporaceae;
OC   Phytophthora.
OX   NCBI_TaxID=4787;
RN   [1]
RP   NUCLEOTIDE SEQUENCE [GENOMIC DNA], 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   [2]
RP   NUCLEOTIDE SEQUENCE [GENOMIC DNA].
RX   PubMed=21303555; DOI=10.1186/1471-2156-12-23;
RA   Cardenas M., Grajales A., Sierra R., Rojas A., Gonzalez-Almario A.,
RA   Vargas A., Marin M., Fermin G., Lagos L.E., Grunwald N.J., Bernal A.,
RA   Salazar C., Restrepo S.;
RT   "Genetic diversity of Phytophthora infestans in the Northern Andean
RT   region.";
RL   BMC Genet. 12:23-23(2011).
RN   [3]
RP   NUCLEOTIDE SEQUENCE [GENOMIC DNA].
RC   STRAIN=SV1;
RX   PubMed=30731720; DOI=10.1094/pdis-11-10-0853;
RA   Cardenas M.E., Medina E., Tabima J., Vargas A., Lopera C., Bernal A.,
RA   Restrepo S.;
RT   "First Report of Phytophthora infestans Causing Late Blight on Solanum
RT   viarum in Colombia.";
RL   Plant Dis. 95:875-875(2011).
RN   [4]
RP   NUCLEOTIDE SEQUENCE [GENOMIC DNA].
RC   STRAIN=Race 1, Race 1.2, Race 1.2.3, Race 1.2.4, and Race 1.3;
RA   Pankin A.A., Kinash E.A., Kozlovskaya I.N., Kuznetsova M.A., Khavkin E.E.;
RT   "Are simple Phytophthora infestans races really that simple?";
RL   PPO Spec. Rep. 15:205-211(2012).
RN   [5]
RP   NUCLEOTIDE SEQUENCE [GENOMIC DNA].
RC   STRAIN=2404, 2474, 2518, 2524, 2557, and Z3-2;
RX   PubMed=23036748; DOI=10.1016/j.riam.2012.09.005;
RA   Cespedes M.C., Cardenas M.E., Vargas A.M., Rojas A., Morales J.G.,
RA   Jimenez P., Bernal A.J., Restrepo S.;
RT   "Physiological and molecular characterization of Phytophthora infestans
RT   isolates from the Central Colombian Andean Region.";
RL   Rev. Iberoam. Micol. 30:81-87(2013).
RN   [6]
RP   NUCLEOTIDE SEQUENCE [GENOMIC DNA].
RC   STRAIN=Race 1, Race 131, and Race EKSL 11.12;
RX   DOI=10.3103/S1068367414060068;
RA   Beketova M., Sokolova E., Malyuchenko O., Alekseev Y., Kuznetsova M.,
RA   Kozlovsky B., Rogozina E., Khavkin E.;
RT   "On molecular identification of Phytophthora infestans genotypes.";
RL   Russ. Agricult. Sci. 40:435-438(2014).
RN   [7]
RP   NUCLEOTIDE SEQUENCE [GENOMIC DNA].
RC   STRAIN=FJ-F041, GS-Pd11330, GZ-GA007, and NX-Pd21418;
RX   PubMed=30151044; DOI=10.1111/eva.12629;
RA   Yang L., Ouyang H.B., Fang Z.G., Zhu W., Wu E.J., Luo G.H., Shang L.P.,
RA   Zhan J.;
RT   "Evidence for intragenic recombination and selective sweep in an effector
RT   gene of Phytophthora infestans.";
RL   Evol. Appl. 11:1342-1353(2018).
RN   [8]
RP   NUCLEOTIDE SEQUENCE [GENOMIC DNA].
RC   STRAIN=EG-005, EG-006, and EG-011;
RA   El-Ganainy S.M., Ahmed Y.M., Soliman M.S., Squires J.N., Cooke D.E.;
RT   "Diversity of RXLR effector genes in Phytophthora infestans isolates from
RT   Egypt.";
RL   Submitted (FEB-2018) to the EMBL/GenBank/DDBJ databases.
RN   [9]
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   [10]
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   [11]
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   [12]
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   [13]
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   [14]
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   [15]
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   [16]
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   [17]
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   [18]
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   [19]
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. Preventing the normal 26S proteasome-dependent degradation of
CC       potato CMPG1, and thus potentially of its protein substrates in the
CC       host cell, further abolishes host cell death during the biotrophic
CC       phase of infection (PubMed:19794118, PubMed:20457921, PubMed:21348873).
CC       Associates with and affects also the function of the dynamin-related
CC       protein 2 (DRP2), a plant GTPase involved in immune receptor-mediated
CC       endocytosis (PubMed:26348328). The Avr3a(EM) form evades recognition by
CC       R3a, thus does not trigger R3a-mediated hypersensitivity and does not
CC       suppress INF1-induced cell death (PubMed:15894622, PubMed:16965554,
CC       PubMed:19245321). {ECO:0000269|PubMed:15894622,
CC       ECO:0000269|PubMed:16965554, ECO:0000269|PubMed:19245321,
CC       ECO:0000269|PubMed:19794118, ECO:0000269|PubMed:20457921,
CC       ECO:0000269|PubMed:21348873, 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:28228125, 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 domain, 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; GU258052; ADC96691.1; -; Genomic_DNA.
DR   EMBL; HQ639930; AEH27535.1; -; Genomic_DNA.
DR   EMBL; JN849402; AFQ32411.1; -; Genomic_DNA.
DR   EMBL; JN849403; AFQ32412.1; -; Genomic_DNA.
DR   EMBL; JN849404; AFQ32413.1; -; Genomic_DNA.
DR   EMBL; JN849405; AFQ32414.1; -; Genomic_DNA.
DR   EMBL; JN849406; AFQ32415.1; -; Genomic_DNA.
DR   EMBL; JN849407; AFQ32416.1; -; Genomic_DNA.
DR   EMBL; KF154421; AGV54925.1; -; Genomic_DNA.
DR   EMBL; KF154423; AGV54927.1; -; Genomic_DNA.
DR   EMBL; KF154424; AGV54928.1; -; Genomic_DNA.
DR   EMBL; KF154429; AGV54933.1; -; Genomic_DNA.
DR   EMBL; KF154430; AGV54934.1; -; Genomic_DNA.
DR   EMBL; KP317569; AKH40250.1; -; Genomic_DNA.
DR   EMBL; KP317575; AKH40255.1; -; Genomic_DNA.
DR   EMBL; KP317576; AKH40256.1; -; Genomic_DNA.
DR   EMBL; KP317585; AKH40263.1; -; Genomic_DNA.
DR   EMBL; MH043151; AXU25232.1; -; Genomic_DNA.
DR   EMBL; MH043159; AXU25240.1; -; Genomic_DNA.
DR   EMBL; MH043178; AXU25259.1; -; Genomic_DNA.
DR   EMBL; MH043185; AXU25266.1; -; Genomic_DNA.
DR   EMBL; MG976602; AYP64705.1; -; Genomic_DNA.
DR   EMBL; MG976603; AYP64706.1; -; Genomic_DNA.
DR   EMBL; MG976604; AYP64707.1; -; Genomic_DNA.
DR   EMBL; AJ893356; CAI72254.1; -; Genomic_DNA.
DR   EMBL; AJ893357; CAI72345.1; -; Genomic_DNA.
DR   AlphaFoldDB; Q572D3; -.
DR   BMRB; Q572D3; -.
DR   SMR; Q572D3; -.
DR   iPTMnet; Q572D3; -.
DR   VEuPathDB; FungiDB:PITG_14371; -.
DR   OMA; VMNDSPK; -.
DR   PHI-base; PHI:473; -.
DR   PHI-base; PHI:4958; -.
DR   PHI-base; PHI:6374; -.
DR   GO; GO:0005576; C:extracellular region; IEA:UniProtKB-SubCell.
DR   GO; GO:0030430; C:host cell cytoplasm; IEA:UniProtKB-SubCell.
DR   GO; GO:0080185; P:effector-mediated induction of plant hypersensitive response by symbiont; IDA:GO_Central.
DR   GO; GO:0044414; P:suppression of host defenses by symbiont; IDA:PAMGO_VMD.
DR   InterPro; IPR031825; RXLR.
DR   Pfam; PF16810; RXLR; 1.
PE   1: Evidence at protein level;
KW   Acetylation; Glycoprotein; Host cytoplasm; Secreted; Signal; Virulence.
FT   SIGNAL          1..21
FT                   /evidence="ECO:0000255"
FT   CHAIN           22..147
FT                   /note="RxLR effector protein Avr3a"
FT                   /id="PRO_5013531960"
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
 
 
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