Gene Symbol: PAD4
Description: alpha/beta-Hydrolases superfamily protein
Alias: ARABIDOPSIS PHYTOALEXIN DEFICIENT 4, ATPAD4, PHYTOALEXIN DEFICIENT 4, alpha/beta-Hydrolases superfamily protein
Species: thale cress

Top Publications

  1. Zhu S, Jeong R, Venugopal S, Lapchyk L, Navarre D, Kachroo A, et al. SAG101 forms a ternary complex with EDS1 and PAD4 and is required for resistance signaling against turnip crinkle virus. PLoS Pathog. 2011;7:e1002318 pubmed publisher
    EDS1, PAD4, and SAG101 are common regulators of plant immunity against many pathogens...
  2. Jirage D, Tootle T, Reuber T, Frost L, Feys B, Parker J, et al. Arabidopsis thaliana PAD4 encodes a lipase-like gene that is important for salicylic acid signaling. Proc Natl Acad Sci U S A. 1999;96:13583-8 pubmed
    The Arabidopsis PAD4 gene previously was found to be required for expression of multiple defense responses including camalexin synthesis and PR-1 gene expression in response to infection by the bacterial pathogen Pseudomonas syringae pv...
  3. Wiermer M, Feys B, Parker J. Plant immunity: the EDS1 regulatory node. Curr Opin Plant Biol. 2005;8:383-9 pubmed
    ENHANCED DISEASE SUSCEPTIBILITY 1 (EDS1) and its interacting partner, PHYTOALEXIN DEFICIENT 4 (PAD4), constitute a regulatory hub that is essential for basal resistance to invasive biotrophic and hemi-biotrophic pathogens...
  4. Feys B, Wiermer M, Bhat R, Moisan L, Medina Escobar N, Neu C, et al. Arabidopsis SENESCENCE-ASSOCIATED GENE101 stabilizes and signals within an ENHANCED DISEASE SUSCEPTIBILITY1 complex in plant innate immunity. Plant Cell. 2005;17:2601-13 pubmed
    ..Together with PHYTOALEXIN-DEFICIENT4 (PAD4), a known EDS1 interactor, SAG101 contributes intrinsic and indispensable signaling activity to EDS1-dependent ..
  5. Pegadaraju V, Knepper C, Reese J, Shah J. Premature leaf senescence modulated by the Arabidopsis PHYTOALEXIN DEFICIENT4 gene is associated with defense against the phloem-feeding green peach aphid. Plant Physiol. 2005;139:1927-34 pubmed
    ..In contrast, resistance against GPA was compromised in the phytoalexin deficient4 (pad4) mutant plant...
  6. Rietz S, Stamm A, Malonek S, Wagner S, Becker D, Medina Escobar N, et al. Different roles of Enhanced Disease Susceptibility1 (EDS1) bound to and dissociated from Phytoalexin Deficient4 (PAD4) in Arabidopsis immunity. New Phytol. 2011;191:107-19 pubmed publisher
    ..Arabidopsis EDS1 interacts with two related proteins, Phytoalexin Deficient4 (PAD4) and Senescence Associated Gene101 (SAG101), whose combined activities are essential for defense signaling...
  7. Nawrath C, Heck S, Parinthawong N, Metraux J. EDS5, an essential component of salicylic acid-dependent signaling for disease resistance in Arabidopsis, is a member of the MATE transporter family. Plant Cell. 2002;14:275-86 pubmed
    ..infection by certain pathogens as well as after UV-C light exposure depends on the pathogen response proteins EDS1, PAD4, and NDR1, indicating that the signal transduction pathways after UV-C light exposure and pathogen inoculation ..
  8. Glazebrook J, Zook M, Mert F, Kagan I, Rogers E, Crute I, et al. Phytoalexin-deficient mutants of Arabidopsis reveal that PAD4 encodes a regulatory factor and that four PAD genes contribute to downy mildew resistance. Genetics. 1997;146:381-92 pubmed
    ..Mutations in PAD1, PAD2, and PAD4 caused enhanced susceptibility to the bacterial pathogen Pseudomonas syringae pv...
  9. Pegadaraju V, Louis J, Singh V, Reese J, Bautor J, Feys B, et al. Phloem-based resistance to green peach aphid is controlled by Arabidopsis PHYTOALEXIN DEFICIENT4 without its signaling partner ENHANCED DISEASE SUSCEPTIBILITY1. Plant J. 2007;52:332-41 pubmed
    ..Previously, it has been shown that Arabidopsis thaliana PHYTOALEXIN DEFICIENT4 (PAD4), which is expressed at elevated levels in response to GPA infestation, is required for resistance to GPA in the ..

More Information


  1. Feys B, Moisan L, Newman M, Parker J. Direct interaction between the Arabidopsis disease resistance signaling proteins, EDS1 and PAD4. EMBO J. 2001;20:5400-11 pubmed
    The Arabidopsis EDS1 and PAD4 genes encode lipase-like proteins that function in resistance (R) gene-mediated and basal plant disease resistance...
  2. Bartsch M, Gobbato E, Bednarek P, Debey S, Schultze J, Bautor J, et al. Salicylic acid-independent ENHANCED DISEASE SUSCEPTIBILITY1 signaling in Arabidopsis immunity and cell death is regulated by the monooxygenase FMO1 and the Nudix hydrolase NUDT7. Plant Cell. 2006;18:1038-51 pubmed
    ..In both responses, EDS1, assisted by its interacting partner, PHYTOALEXIN-DEFICIENT4 (PAD4), regulates accumulation of the phenolic defense molecule salicylic acid (SA) and other as yet unidentified signal ..
  3. Wagner S, Stuttmann J, Rietz S, Guerois R, Brunstein E, Bautor J, et al. Structural basis for signaling by exclusive EDS1 heteromeric complexes with SAG101 or PAD4 in plant innate immunity. Cell Host Microbe. 2013;14:619-30 pubmed publisher
    ..1 (EDS1) and its sequence-related interaction partners, senescence-associated gene 101 (SAG101) and phytoalexin deficient 4 (PAD4)...
  4. Ke Y, Liu H, Li X, Xiao J, Wang S. Rice OsPAD4 functions differently from Arabidopsis AtPAD4 in host-pathogen interactions. Plant J. 2014;78:619-31 pubmed publisher
    The extensively studied Arabidopsis phytoalexin deficient 4 (AtPAD4) gene plays an important role in Arabidopsis disease resistance; however, the function of its sequence ortholog in rice is unknown...
  5. Cui H, Gobbato E, Kracher B, Qiu J, Bautor J, Parker J. A core function of EDS1 with PAD4 is to protect the salicylic acid defense sector in Arabidopsis immunity. New Phytol. 2017;213:1802-1817 pubmed publisher
    ..SA accumulation is promoted by Enhanced Disease Susceptibility1 with its co-regulator Phytoalexin Deficient4 (EDS1/PAD4). Current models position EDS1/PAD4 upstream of SA but their functional relationship remains unclear...
  6. Youssef R, MacDonald M, Brewer E, Bauchan G, Kim K, Matthews B. Ectopic expression of AtPAD4 broadens resistance of soybean to soybean cyst and root-knot nematodes. BMC Plant Biol. 2013;13:67 pubmed publisher
    ..maculicola. AtPAD4 (Arabidopsis thaliana PAD4) encodes a lipase-like protein that plays a regulatory role mediating salicylic acid ..
  7. Hillmer R, Tsuda K, Rallapalli G, Asai S, Truman W, Papke M, et al. The highly buffered Arabidopsis immune signaling network conceals the functions of its components. PLoS Genet. 2017;13:e1006639 pubmed publisher
    ..regulatory mechanisms show unexpected patterns for how the jasmonate (JA), ethylene (ET), phytoalexin-deficient 4 (PAD4), and salicylate (SA) signaling sectors control the transcriptional response to flg22...
  8. Mühlenbock P, Plaszczyca M, Plaszczyca M, Mellerowicz E, Karpinski S. Lysigenous aerenchyma formation in Arabidopsis is controlled by LESION SIMULATING DISEASE1. Plant Cell. 2007;19:3819-30 pubmed
    ..regulators LESION SIMULATING DISEASE1 (LSD1), ENHANCED DISEASE SUSCEPIBILITY1 (EDS1), and PHYTOALEXIN DEFICIENT4 (PAD4) that operate upstream of ethylene and reactive oxygen species production...
  9. Wang J, Shine M, Gao Q, Navarre D, Jiang W, Liu C, et al. Enhanced Disease Susceptibility1 Mediates Pathogen Resistance and Virulence Function of a Bacterial Effector in Soybean. Plant Physiol. 2014;165:1269-1284 pubmed
    Enhanced disease susceptibility1 (EDS1) and phytoalexin deficient4 (PAD4) are well-known regulators of both basal and resistance (R) protein-mediated plant defense...
  10. Wituszynska W, Slesak I, Vanderauwera S, Szechynska Hebda M, Kornas A, Van Der Kelen K, et al. Lesion simulating disease1, enhanced disease susceptibility1, and phytoalexin deficient4 conditionally regulate cellular signaling homeostasis, photosynthesis, water use efficiency, and seed yield in Arabidopsis. Plant Physiol. 2013;161:1795-805 pubmed publisher
    ..the role of lesion simulating disease1 (LSD1), enhanced disease susceptibility1 (EDS1), and phytoalexin deficient4 (PAD4) in the regulation of photosynthesis, water use efficiency, reactive oxygen species/hormonal homeostasis, and seed ..
  11. Louis J, Gobbato E, Mondal H, Feys B, Parker J, Shah J. Discrimination of Arabidopsis PAD4 activities in defense against green peach aphid and pathogens. Plant Physiol. 2012;158:1860-72 pubmed publisher
    The Arabidopsis (Arabidopsis thaliana) lipase-like protein PHYTOALEXIN DEFICIENT4 (PAD4) is essential for defense against green peach aphid (GPA; Myzus persicae) and the pathogens Pseudomonas syringae and Hyaloperonospora arabidopsidis...
  12. Wang B, Wang N, Song N, Wang W, Wang J, Wang X, et al. Overexpression of AtPAD4 in transgenic Brachypodium distachyon enhances resistance to Puccinia brachypodii. Plant Biol (Stuttg). 2017;19:868-874 pubmed publisher
    ..b>Phytoalexin deficient 4 (PAD4) plays a regulatory role in mediating expression of genes involved in plant defence...
  13. Makandar R, Nalam V, Chowdhury Z, Sarowar S, Klossner G, Lee H, et al. The Combined Action of ENHANCED DISEASE SUSCEPTIBILITY1, PHYTOALEXIN DEFICIENT4, and SENESCENCE-ASSOCIATED101 Promotes Salicylic Acid-Mediated Defenses to Limit Fusarium graminearum Infection in Arabidopsis thaliana. Mol Plant Microbe Interact. 2015;28:943-53 pubmed publisher
    ..that, in Arabidopsis, the defense regulator EDS1 (ENHANCED DISEASE SUSCEPTIBILITY1) and its interacting partners, PAD4 (PHYTOALEXIN-DEFICIENT4) and SAG101 (SENESCENCE-ASSOCIATED GENE101), promote SA accumulation to curtail F...
  14. Gupta V, Willits M, Glazebrook J. Arabidopsis thaliana EDS4 contributes to salicylic acid (SA)-dependent expression of defense responses: evidence for inhibition of jasmonic acid signaling by SA. Mol Plant Microbe Interact. 2000;13:503-11 pubmed
    ..Two mutations that cause reduced SA levels, eds4 and pad4, cause heightened responses to inducers of JA-dependent gene expression, providing genetic evidence to support the ..
  15. Louis J, Leung Q, Pegadaraju V, Reese J, Shah J. PAD4-dependent antibiosis contributes to the ssi2-conferred hyper-resistance to the green peach aphid. Mol Plant Microbe Interact. 2010;23:618-27 pubmed publisher
    ..Arabidopsis thaliana responds to GPA infestation with elevated expression of the PHYTOALEXIN DEFICIENT4 (PAD4) gene...
  16. Gao M, Wang X, Wang D, Xu F, Ding X, Zhang Z, et al. Regulation of cell death and innate immunity by two receptor-like kinases in Arabidopsis. Cell Host Microbe. 2009;6:34-44 pubmed publisher
    ..Our data suggest that BIR1 negatively regulates multiple plant resistance signaling pathways, one of which is the SOBIR1-dependent pathway identified here. ..
  17. Mühlenbock P, Szechynska Hebda M, Plaszczyca M, Baudo M, Mateo A, Mullineaux P, et al. Chloroplast signaling and LESION SIMULATING DISEASE1 regulate crosstalk between light acclimation and immunity in Arabidopsis. Plant Cell. 2008;20:2339-56 pubmed publisher
    ..death depends on the plant defense regulators ENHANCED DISEASE SUSCEPTIBILITY1 (EDS1) and PHYTOALEXIN DEFICIENT4 (PAD4). We find that EDS1 and PAD4 operate upstream of ethylene and ROS production in the EEE response...
  18. Wang Y, Zhang Y, Wang Z, Zhang X, Yang S. A missense mutation in CHS1, a TIR-NB protein, induces chilling sensitivity in Arabidopsis. Plant J. 2013;75:553-65 pubmed publisher
    ..The chilling sensitivity of chs1 was fully rescued by pad4 and eds1, but not by ndr1. The overexpression of the TIR and NB domains can suppress the chs1-conferred phenotypes...
  19. Consonni C, Humphry M, Hartmann H, Livaja M, Durner J, Westphal L, et al. Conserved requirement for a plant host cell protein in powdery mildew pathogenesis. Nat Genet. 2006;38:716-20 pubmed
  20. Kohorn B, Kohorn S, Saba N, Martinez V. Requirement for pectin methyl esterase and preference for fragmented over native pectins for wall-associated kinase-activated, EDS1/PAD4-dependent stress response in Arabidopsis. J Biol Chem. 2014;289:18978-86 pubmed publisher
    ..of WAK2 constitutively activates the stress response, and we show here that the effect is dependent upon EDS1 and PAD4, transcriptional activators involved in the pathogen response...
  21. Gao F, Dai R, Pike S, Qiu W, Gassmann W. Functions of EDS1-like and PAD4 genes in grapevine defenses against powdery mildew. Plant Mol Biol. 2014;86:381-93 pubmed publisher
    ..acid (SA) pathway, including ENHANCED DISEASE SUSCEPTIBILITY 1 (EDS1), EDS1-LIKE 2 (EDL2), EDL5 and PHYTOALEXIN DEFICIENT 4 (PAD4) of two grapevine species, Vitis vinifera cv. Cabernet Sauvignon, which is susceptible to E...
  22. Mukherjee M, Larrimore K, Ahmed N, Bedick T, Barghouthi N, Traw M, et al. Ascorbic acid deficiency in arabidopsis induces constitutive priming that is dependent on hydrogen peroxide, salicylic acid, and the NPR1 gene. Mol Plant Microbe Interact. 2010;23:340-51 pubmed publisher
    ..We also studied double mutants defective in the AA-biosynthetic gene VTC1 and the SA signaling pathway genes PAD4, EDS5, and NPR1, respectively. All vtc mutants were more resistant to P. syringae than the wild type...
  23. Szechyńska Hebda M, Czarnocka W, Hebda M, Bernacki M, Karpiński S. PAD4, LSD1 and EDS1 regulate drought tolerance, plant biomass production, and cell wall properties. Plant Cell Rep. 2016;35:527-39 pubmed publisher
    ..g. in areas with poor irrigation. In this study, we performed functional analysis of proteins encoded by PHYTOALEXIN DEFICIENT 4 (PAD4), LESION SIMULATING DISEASE 1 (LSD1) and ENHANCED DISEASE SUSCEPTIBILITY 1 (EDS1) genes to explain ..
  24. Mishina T, Zeier J. Pathogen-associated molecular pattern recognition rather than development of tissue necrosis contributes to bacterial induction of systemic acquired resistance in Arabidopsis. Plant J. 2007;50:500-13 pubmed
    ..Our data indicate that PAMPs significantly contribute to SAR initiation in Arabidopsis and that tissue necroses at inoculation sites are dispensable for SAR activation. ..
  25. Singh V, Louis J, Ayre B, Reese J, Pegadaraju V, Shah J. TREHALOSE PHOSPHATE SYNTHASE11-dependent trehalose metabolism promotes Arabidopsis thaliana defense against the phloem-feeding insect Myzus persicae. Plant J. 2011;67:94-104 pubmed publisher
    ..Our results provide a framework for the signaling function of TPS11-dependent trehalose in plant stress responses, and also reveal an important contribution of starch in controlling the severity of aphid infestation. ..
  26. Carstens M, McCrindle T, Adams N, Diener A, Guzha D, Murray S, et al. Increased resistance to biotrophic pathogens in the Arabidopsis constitutive induced resistance 1 mutant is EDS1 and PAD4-dependent and modulated by environmental temperature. PLoS ONE. 2014;9:e109853 pubmed publisher
    ..SA-signalling branch of the defence network, ENHANCED DISEASE SUSCEPTIBILITY1 (EDS1) and PHYTOALEXIN DEFICIENT4 (PAD4)...
  27. Vogelmann K, Drechsel G, Bergler J, Subert C, Philippar K, Soll J, et al. Early senescence and cell death in Arabidopsis saul1 mutants involves the PAD4-dependent salicylic acid pathway. Plant Physiol. 2012;159:1477-87 pubmed publisher
    ..Double mutant analyses showed that PHYTOALEXIN DEFICIENT4 (PAD4) but not NONEXPRESSER OF PR GENES1 (NPR1) is essential for saul1 phenotypes...
  28. Brodersen P, Petersen M, Bjørn Nielsen H, Zhu S, Newman M, Shokat K, et al. Arabidopsis MAP kinase 4 regulates salicylic acid- and jasmonic acid/ethylene-dependent responses via EDS1 and PAD4. Plant J. 2006;47:532-46 pubmed
    ..and ET/JA-(co)activating functions depend on MPK4 kinase activity and involve the defense regulators EDS1 and PAD4, as mutations in these genes suppress de-repression of the SA pathway and suppress the block of the ET/JA pathway ..
  29. Zhou N, Tootle T, Tsui F, Klessig D, Glazebrook J. PAD4 functions upstream from salicylic acid to control defense responses in Arabidopsis. Plant Cell. 1998;10:1021-30 pubmed
    The Arabidopsis PAD4 gene was previously shown to be required for synthesis of camalexin in response to infection by the virulent bacterial pathogen Pseudomonas syringae pv maculicola ES4326 but not in response to challenge by the non-..
  30. Kim T, Kunz H, Bhattacharjee S, Hauser F, Park J, Engineer C, et al. Natural variation in small molecule-induced TIR-NB-LRR signaling induces root growth arrest via EDS1- and PAD4-complexed R protein VICTR in Arabidopsis. Plant Cell. 2012;24:5177-92 pubmed publisher
    ..DFPM) that triggers rapid inhibition of early abscisic acid signal transduction via PHYTOALEXIN DEFICIENT4 (PAD4)- and ENHANCED DISEASE SUSCEPTIBILITY1 (EDS1)-dependent immune signaling mechanisms...