A3AKI_PHYIN
ID A3AKI_PHYIN Reviewed; 147 AA.
AC E2DWQ7;
DT 08-MAY-2019, integrated into UniProtKB/Swiss-Prot.
DT 30-NOV-2010, sequence version 1.
DT 25-MAY-2022, entry version 37.
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(KI) {ECO:0000303|PubMed:19245321};
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].
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 [2]
RP NUCLEOTIDE SEQUENCE [GENOMIC DNA].
RC STRAIN=Race 1.2.3;
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 [3]
RP NUCLEOTIDE SEQUENCE [GENOMIC DNA].
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 [4]
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 [5]
RP FUNCTION, AND DOMAIN.
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 [6]
RP DOMAIN, SUBCELLULAR LOCATION, AND MUTAGENESIS OF 44-ARG--ARG-49 AND
RP 57-GLU--ARG-59.
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 [7]
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 [8]
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 [9]
RP FUNCTION, INTERACTION WITH HOST E3 LIGASE CMPG1, AND MUTAGENESIS OF
RP TYR-147.
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 [10]
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 [11]
RP DOMAIN, MUTAGENESIS OF ARG-81; LYS-85; LYS-86 AND LYS-89, AND
RP PHOSPHATIDYLINOSITOL MONOPHOSPHATE-BINDING.
RX PubMed=21821794; DOI=10.1073/pnas.1106002108;
RA Yaeno T., Li H., Chaparro-Garcia A., Schornack S., Koshiba S., Watanabe S.,
RA Kigawa T., Kamoun S., Shirasu K.;
RT "Phosphatidylinositol monophosphate-binding interface in the oomycete RXLR
RT effector AVR3a is required for its stability in host cells to modulate
RT plant immunity.";
RL Proc. Natl. Acad. Sci. U.S.A. 108:14682-14687(2011).
RN [12]
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 [13]
RP SUBCELLULAR LOCATION.
RX PubMed=23243124; DOI=10.1105/tpc.112.104992;
RA Engelhardt S., Boevink P.C., Armstrong M.R., Ramos M.B., Hein I.,
RA Birch P.R.;
RT "Relocalization of late blight resistance protein R3a to endosomal
RT compartments is associated with effector recognition and required for the
RT immune response.";
RL Plant Cell 24:5142-5158(2012).
RN [14]
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 [15]
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 [16]
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).
RN [17] {ECO:0007744|PDB:2NAR}
RP STRUCTURE BY NMR OF 61-147, CLEAVAGE, ACETYLATION AT LYS-48, SUBUNIT, AND
RP 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).
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 also with the dynamin-related protein 2 (DRP2), a plant
CC GTPase involved in immune receptor-mediated endocytosis
CC (PubMed:26348328). The Avr3A(KI) form is recognized by R3a which
CC triggers R3a-mediated hypersensitivity and suppresses INF1-induced cell
CC death (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 PubMed:28522546). Interacts with host E3 ligase CMPG1
CC (PubMed:20457921). Interacts with host DRP2 (PubMed:26348328).
CC {ECO:0000269|PubMed:20457921, ECO:0000269|PubMed:22977236,
CC ECO:0000269|PubMed:26348328, ECO:0000269|PubMed:28522546}.
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}. Host
CC endosome {ECO:0000269|PubMed:23243124}. Note=Avr3A(KI), but not
CC Avr3A(EM), is relocalized to endosomes in the presence of R3a.
CC {ECO:0000269|PubMed:23243124}.
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 PubMed:21821794). The motif is cleaved after the RxLR sequence
CC (PubMed:28522546). The RxLR motif (residues 44 to 47) plays a crucial
CC role in the intracellular processing before secretion
CC (PubMed:28522546). The Glu-rich part localized just after the cleavage
CC site (residues 48 to 59) is required for homodimerization
CC (PubMed:28522546). {ECO:0000269|PubMed:17914356,
CC ECO:0000269|PubMed:21821794, 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:0000269|PubMed:21821794}.
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}.
CC ---------------------------------------------------------------------------
CC Copyrighted by the UniProt Consortium, see https://www.uniprot.org/terms
CC Distributed under the Creative Commons Attribution (CC BY 4.0) License
CC ---------------------------------------------------------------------------
DR EMBL; GU258055; ADC96694.1; -; Genomic_DNA.
DR EMBL; KF154426; AGV54930.1; -; Genomic_DNA.
DR EMBL; KP317568; AKH40249.1; -; Genomic_DNA.
DR PDB; 2NAR; NMR; -; A=61-147.
DR PDBsum; 2NAR; -.
DR AlphaFoldDB; E2DWQ7; -.
DR SMR; E2DWQ7; -.
DR iPTMnet; E2DWQ7; -.
DR VEuPathDB; FungiDB:PITG_14371; -.
DR GO; GO:0005576; C:extracellular region; IEA:UniProtKB-SubCell.
DR GO; GO:0044174; C:host cell endosome; IEA:UniProtKB-SubCell.
DR InterPro; IPR031825; RXLR.
DR Pfam; PF16810; RXLR; 1.
PE 1: Evidence at protein level;
KW 3D-structure; Acetylation; Glycoprotein; Host cytoplasm; Host endosome;
KW Secreted; Signal; Virulence.
FT SIGNAL 1..21
FT /evidence="ECO:0000255"
FT CHAIN 22..147
FT /note="RxLR effector protein Avr3a"
FT /id="PRO_5013522507"
FT REGION 77..147
FT /note="Effector domain"
FT /evidence="ECO:0000305|PubMed:21821794"
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"
FT MUTAGEN 44..47
FT /note="RLLR->AAAA: Prevents delivery of Avr3a into host
FT cells."
FT /evidence="ECO:0000269|PubMed:17914356"
FT MUTAGEN 57..59
FT /note="EER->AAA: Prevents delivery of Avr3a into host
FT cells."
FT /evidence="ECO:0000269|PubMed:17914356"
FT MUTAGEN 81
FT /note="R->E: Does not affect PIP-binding."
FT /evidence="ECO:0000269|PubMed:21821794"
FT MUTAGEN 85
FT /note="K->E: Reduces the PIP-binding ability."
FT /evidence="ECO:0000269|PubMed:21821794"
FT MUTAGEN 86
FT /note="K->E: Does not affect PIP-binding."
FT /evidence="ECO:0000269|PubMed:21821794"
FT MUTAGEN 89
FT /note="K->E: Reduces the PIP-binding ability."
FT /evidence="ECO:0000269|PubMed:21821794"
FT MUTAGEN 147
FT /note="Missing: Prevents the interaction with host CMPG1."
FT /evidence="ECO:0000269|PubMed:20457921"
FT TURN 75..77
FT /evidence="ECO:0007829|PDB:2NAR"
FT HELIX 82..93
FT /evidence="ECO:0007829|PDB:2NAR"
FT HELIX 95..107
FT /evidence="ECO:0007829|PDB:2NAR"
FT HELIX 112..122
FT /evidence="ECO:0007829|PDB:2NAR"
FT STRAND 125..127
FT /evidence="ECO:0007829|PDB:2NAR"
FT HELIX 131..142
FT /evidence="ECO:0007829|PDB:2NAR"
SQ SEQUENCE 147 AA; 16658 MW; 1D9D02E2F11E8C45 CRC64;
MRLAIMLSAT AVAINFATCS AIDQTKVLVY GTPAHYIHDS AGRRLLRKNE ENEETSEERA
PNFNLANLNE EMFNVAALTK RADAKKLAKQ LMGNDKLADA AYIWWQHNRV TLDQIDTFLK
LASRKTQGAK YNQIYNSYMM HLGLTGY
