Gene Symbol: EIR1
Description: Auxin efflux carrier family protein
Species: thale cress

Top Publications

  1. Michniewicz M, Zago M, Abas L, Weijers D, Schweighofer A, Meskiene I, et al. Antagonistic regulation of PIN phosphorylation by PP2A and PINOID directs auxin flux. Cell. 2007;130:1044-56 pubmed
    ..In the case of PIN proteins, this mechanism enables switches in the direction of intercellular auxin fluxes, which mediate differential growth, tissue patterning, and organogenesis. ..
  2. Kleine Vehn J, Leitner J, Zwiewka M, Sauer M, Abas L, Luschnig C, et al. Differential degradation of PIN2 auxin efflux carrier by retromer-dependent vacuolar targeting. Proc Natl Acad Sci U S A. 2008;105:17812-7 pubmed publisher
    ..In vivo visualization of PIN2 vacuolar targeting revealed its differential degradation in response to environmental signals, such as gravity...
  3. Shin H, Shin H, Guo Z, Blancaflor E, Masson P, Chen R. Complex regulation of Arabidopsis AGR1/PIN2-mediated root gravitropic response and basipetal auxin transport by cantharidin-sensitive protein phosphatases. Plant J. 2005;42:188-200 pubmed
    ..However, the regulatory mechanism underlying the function of AGR1/PIN2 is largely unknown...
  4. Shibasaki K, Uemura M, Tsurumi S, Rahman A. Auxin response in Arabidopsis under cold stress: underlying molecular mechanisms. Plant Cell. 2009;21:3823-38 pubmed publisher
    ..Microscopy of living cells revealed that trafficking of the auxin efflux carrier PIN2, which acts in basipetal auxin transport, was dramatically reduced by cold...
  5. Wisniewska J, Xu J, Seifertová D, Brewer P, Ruzicka K, Blilou I, et al. Polar PIN localization directs auxin flow in plants. Science. 2006;312:883 pubmed
    ..This finding provides a crucial piece in the puzzle of how auxin flow can be redirected via rapid changes in PIN polarity. ..
  6. Muller A, Guan C, Gälweiler L, Tänzler P, Huijser P, Marchant A, et al. AtPIN2 defines a locus of Arabidopsis for root gravitropism control. EMBO J. 1998;17:6903-11 pubmed
    ..We have isolated an Arabidopsis mutant of the AtPIN2 gene using transposon mutagenesis...
  7. Wan Y, Jasik J, Wang L, Hao H, Volkmann D, Menzel D, et al. The signal transducer NPH3 integrates the phototropin1 photosensor with PIN2-based polar auxin transport in Arabidopsis root phototropism. Plant Cell. 2012;24:551-65 pubmed publisher
    ..phototropin1 (phot1), the signal transducer NONPHOTOTROPIC HYPOCOTYL3 (NPH3), and the auxin efflux transporter PIN2 were essential for BL-induced auxin flux in the root apex transition zone...
  8. Leitner J, Petrasek J, Tomanov K, Retzer K, Parezová M, Korbei B, et al. Lysine63-linked ubiquitylation of PIN2 auxin carrier protein governs hormonally controlled adaptation of Arabidopsis root growth. Proc Natl Acad Sci U S A. 2012;109:8322-7 pubmed publisher
    ..Here, we describe the fate of Arabidopsis PIN2 protein, required for directional cellular efflux of the phytohormone auxin, and identify cis- and trans-acting ..
  9. Rahman A, Takahashi M, Shibasaki K, Wu S, Inaba T, Tsurumi S, et al. Gravitropism of Arabidopsis thaliana roots requires the polarization of PIN2 toward the root tip in meristematic cortical cells. Plant Cell. 2010;22:1762-76 pubmed publisher
    ..auxin transport, and hence gravitropism, depends on the polar deployment of the PIN-FORMED auxin efflux carrier PIN2. In Arabidopsis thaliana, PIN2 has the expected shootward localization in epidermis and lateral root cap; however, ..

More Information


  1. Abas L, Benjamins R, Malenica N, Paciorek T, Wisniewska J, Wirniewska J, et al. Intracellular trafficking and proteolysis of the Arabidopsis auxin-efflux facilitator PIN2 are involved in root gravitropism. Nat Cell Biol. 2006;8:249-56 pubmed
    ..Here, we show that intracellular trafficking and proteasome activity combine to control PIN2 degradation during root gravitropism...
  2. Feraru E, Feraru M, Kleine Vehn J, Martinière A, Mouille G, Vanneste S, et al. PIN polarity maintenance by the cell wall in Arabidopsis. Curr Biol. 2011;21:338-43 pubmed publisher
  3. Dhonukshe P, Huang F, Galvan Ampudia C, Mähönen A, Kleine Vehn J, Xu J, et al. Plasma membrane-bound AGC3 kinases phosphorylate PIN auxin carriers at TPRXS(N/S) motifs to direct apical PIN recycling. Development. 2010;137:3245-55 pubmed publisher
  4. Petrasek J, Mravec J, Bouchard R, Blakeslee J, Abas M, Seifertová D, et al. PIN proteins perform a rate-limiting function in cellular auxin efflux. Science. 2006;312:914-8 pubmed
    ..This suggests a direct involvement of PINs in catalyzing cellular auxin efflux...
  5. Luschnig C, Gaxiola R, Grisafi P, Fink G. EIR1, a root-specific protein involved in auxin transport, is required for gravitropism in Arabidopsis thaliana. Genes Dev. 1998;12:2175-87 pubmed
    The EIR1 gene of Arabidopsis is a member of a family of plant genes with similarities to bacterial membrane transporters...
  6. Willige B, Isono E, Richter R, Zourelidou M, Schwechheimer C. Gibberellin regulates PIN-FORMED abundance and is required for auxin transport-dependent growth and development in Arabidopsis thaliana. Plant Cell. 2011;23:2184-95 pubmed publisher
    ..explained by a transcriptional regulation of the PIN genes but that GA deficiency promotes, at least in the case of PIN2, the targeting of PIN proteins for vacuolar degradation...
  7. Baster P, Robert S, Kleine Vehn J, Vanneste S, Kania U, Grunewald W, et al. SCF(TIR1/AFB)-auxin signalling regulates PIN vacuolar trafficking and auxin fluxes during root gravitropism. EMBO J. 2013;32:260-74 pubmed publisher
    ..We show that PIN2 turnover is differentially regulated on the upper and lower side of gravistimulated roots by distinct but partially ..
  8. Leitner J, Retzer K, Korbei B, Luschnig C. Dynamics in PIN2 auxin carrier ubiquitylation in gravity-responding Arabidopsis roots. Plant Signal Behav. 2012;7:1271-3 pubmed publisher
    ..1) (-) ( 6) Here we show dynamic adjustments in PIN2 ubiquitylation in gravity-stimulated roots, a response that coincides with establishment of a lateral PIN2 ..
  9. Karampelias M, Neyt P, De Groeve S, Aesaert S, Coussens G, Rolčík J, et al. ROTUNDA3 function in plant development by phosphatase 2A-mediated regulation of auxin transporter recycling. Proc Natl Acad Sci U S A. 2016;113:2768-73 pubmed publisher
    ..Our data indicate that RON3 contributes to auxin-mediated development by playing a role in PIN recycling and polarity establishment through regulation of the PP2A complex activity. ..
  10. Drdová E, Synek L, Pecenková T, Hála M, Kulich I, Fowler J, et al. The exocyst complex contributes to PIN auxin efflux carrier recycling and polar auxin transport in Arabidopsis. Plant J. 2013;73:709-19 pubmed publisher
    ..The recycling of PIN1 and PIN2 proteins from brefeldin-A compartments is delayed after the brefeldin-A washout in exo70A1 and sec8 exocyst mutants...
  11. Ambrose C, Ruan Y, Gardiner J, Tamblyn L, Catching A, Kirik V, et al. CLASP interacts with sorting nexin 1 to link microtubules and auxin transport via PIN2 recycling in Arabidopsis thaliana. Dev Cell. 2013;24:649-59 pubmed publisher
    ..Consistent with SNX1's role in trafficking of the auxin efflux carrier PIN-FORMED2 (PIN2), clasp-1 mutant plants have enhanced PIN2 degradation, and PIN2 movement to lytic vacuoles is rapidly induced by ..
  12. Löfke C, Scheuring D, Dünser K, Schöller M, Luschnig C, Kleine Vehn J. Tricho- and atrichoblast cell files show distinct PIN2 auxin efflux carrier exploitations and are jointly required for defined auxin-dependent root organ growth. J Exp Bot. 2015;66:5103-12 pubmed publisher
    ..The pharmacologic, genetic, and live-cell imaging approaches reveal that PIN2 auxin efflux carriers are differentially controlled in tricho- and atrichoblast cells...
  13. Eysholdt Derzsó E, Sauter M. Root Bending Is Antagonistically Affected by Hypoxia and ERF-Mediated Transcription via Auxin Signaling. Plant Physiol. 2017;175:412-423 pubmed publisher
    ..The protein abundance of the auxin efflux carrier PIN2 is reduced at hypoxic conditions, a response that is suppressed by RAP2...
  14. Jasik J, Bokor B, Stuchlík S, Micieta K, Turna J, Schmelzer E. Effects of Auxins on PIN-FORMED2 (PIN2) Dynamics Are Not Mediated by Inhibiting PIN2 Endocytosis. Plant Physiol. 2016;172:1019-1031 pubmed
    ..and 1-N-naphthylphthalamic acid, were able to inhibit endocytosis of the putative auxin transporter PIN-FORMED2 (PIN2) in Arabidopsis (Arabidopsis thaliana) root epidermis cells...
  15. Honda H, Hamasaki R, Ejima C, Shimizu N, Kiyohara S, Sawa S. MM31/EIR1 promotes lateral root formation in Arabidopsis. Plant Signal Behav. 2011;6:968-73 pubmed
    ..Based on genetic and physiological analyses, we propose that MM31/EIR1 modulates lateral root formation by regulating the IAA polar transport system, and that auxin transport from the ..
  16. Pan J, Fujioka S, Peng J, Chen J, Li G, Chen R. The E3 ubiquitin ligase SCFTIR1/AFB and membrane sterols play key roles in auxin regulation of endocytosis, recycling, and plasma membrane accumulation of the auxin efflux transporter PIN2 in Arabidopsis thaliana. Plant Cell. 2009;21:568-80 pubmed publisher
    ..Auxin inhibits PIN2 endocytosis and promotes its PM localization. However, the underlying mechanisms remain elusive...
  17. Mei Y, Jia W, Chu Y, Xue H. Arabidopsis phosphatidylinositol monophosphate 5-kinase 2 is involved in root gravitropism through regulation of polar auxin transport by affecting the cycling of PIN proteins. Cell Res. 2012;22:581-97 pubmed publisher
    ..vesicle trafficking caused by PtdIns(4,5)P(2) reduction, which hence results in suppressed cycling of PIN proteins (PIN2 and 3), and delayed redistribution of PIN2 and auxin under gravistimulation in pip5k2 roots...
  18. Dai M, Zhang C, Kania U, Chen F, Xue Q, McCray T, et al. A PP6-type phosphatase holoenzyme directly regulates PIN phosphorylation and auxin efflux in Arabidopsis. Plant Cell. 2012;24:2497-514 pubmed publisher
    ..Thus, an Arabidopsis PP6-type phosphatase holoenzyme acts antagonistically with PID to direct auxin transport polarity and plant development by directly regulating PIN phosphorylation...
  19. Sassi M, Lu Y, Zhang Y, Wang J, Dhonukshe P, Blilou I, et al. COP1 mediates the coordination of root and shoot growth by light through modulation of PIN1- and PIN2-dependent auxin transport in Arabidopsis. Development. 2012;139:3402-12 pubmed publisher
    ..elongation and adapts auxin transport and cell proliferation in the root apical meristem by modulating PIN1 and PIN2 intracellular distribution in the root in a COP1-dependent fashion, thus permitting a rapid and precise tuning of ..
  20. Katz E, Nisani S, Sela M, Behar H, Chamovitz D. The effect of indole-3-carbinol on PIN1 and PIN2 in Arabidopsis roots. Plant Signal Behav. 2015;10:e1062200 pubmed publisher
    ..Brief treatment with indole-3-carbinol led to a reduction in the amount of PIN1 and to mislocalization of PIN2.
  21. Jasik J, Boggetti B, Baluska F, Volkmann D, Gensch T, Rutten T, et al. PIN2 turnover in Arabidopsis root epidermal cells explored by the photoconvertible protein Dendra2. PLoS ONE. 2013;8:e61403 pubmed publisher
    ..Dendra2 was fused to the PIN2 protein, an auxin transporter in the root tip, and by time-lapse imaging and assessment of red and green signal ..
  22. Li K, Kamiya T, Fujiwara T. Differential Roles of PIN1 and PIN2 in Root Meristem Maintenance Under Low-B Conditions in Arabidopsis thaliana. Plant Cell Physiol. 2015;56:1205-14 pubmed publisher
    ..Among them, only the pin2/eir1-1 mutant showed a significantly shorter root under low-B conditions than under control conditions...
  23. Kleine Vehn J, Langowski L, Wisniewska J, Dhonukshe P, Brewer P, Friml J. Cellular and molecular requirements for polar PIN targeting and transcytosis in plants. Mol Plant. 2008;1:1056-66 pubmed publisher
    ..The resulting dynamic PIN positioning to different sides of cells defines a three-dimensional pattern of auxin fluxes within plant tissues. ..
  24. Kumar M, Pandya Kumar N, Dam A, Haor H, Mayzlish Gati E, Belausov E, et al. Arabidopsis response to low-phosphate conditions includes active changes in actin filaments and PIN2 polarization and is dependent on strigolactone signalling. J Exp Bot. 2015;66:1499-510 pubmed publisher
    ..At least part of SL-signalling execution in roots involves MAX2-dependent effects on PIN2 polar localization in the plasma membrane (PM) and actin bundling and dynamics...
  25. Lin D, Nagawa S, Chen J, Cao L, Chen X, Xu T, et al. A ROP GTPase-dependent auxin signaling pathway regulates the subcellular distribution of PIN2 in Arabidopsis roots. Curr Biol. 2012;22:1319-25 pubmed publisher
    ..gene whose loss-of-function mutations increased lateral root density and retarded gravitropic responses, as do pin2 knockout mutations. SPK1 belongs to the conserved DHR2-Dock family of Rho guanine nucleotide exchange factors...
  26. Diener A. Visualizing and quantifying Fusarium oxysporum in the plant host. Mol Plant Microbe Interact. 2012;25:1531-41 pubmed publisher
    ..were measured in the roots of susceptible agb1 and rfo1 while a reduced level was detected in the resistant eir1. In contrast, disease severity and F. oxysporum infection were uncoupled in tir3...
  27. Na X, Hu Y, Yue K, Lu H, Jia P, Wang H, et al. Narciclasine modulates polar auxin transport in Arabidopsis roots. J Plant Physiol. 2011;168:1149-56 pubmed publisher
    ..Expressions of auxin transport genes were affected and the polar localization of PIN2 protein was altered under NCS treatment...
  28. Xu W, Jia L, Baluska F, Ding G, Shi W, Ye N, et al. PIN2 is required for the adaptation of Arabidopsis roots to alkaline stress by modulating proton secretion. J Exp Bot. 2012;63:6105-14 pubmed publisher
    ..It was found that alkaline stress increased auxin transport and PIN2 (an auxin efflux transporter) abundance in the root tip of wild-type Arabidopsis plants (WT)...
  29. Zourelidou M, Müller I, Willige B, Nill C, Jikumaru Y, Li H, et al. The polarly localized D6 PROTEIN KINASE is required for efficient auxin transport in Arabidopsis thaliana. Development. 2009;136:627-36 pubmed publisher
    ..D6PK localizes to the basal (lower) membrane of Arabidopsis root cells, where it colocalizes with PIN1, PIN2 and PIN4. D6PK and PIN1 interact genetically, and D6PK phosphorylates PIN proteins in vitro and in vivo...
  30. Kim S, Xu Z, Song K, Kim D, Kang H, Reichardt I, et al. Adaptor protein complex 2-mediated endocytosis is crucial for male reproductive organ development in Arabidopsis. Plant Cell. 2013;25:2970-85 pubmed publisher
    ..a lypophilic dye used as an endocytosis marker, and polar localization of auxin-efflux carrier PIN FORMED2 (PIN2) in the stamen filaments...
  31. Chen R, Hilson P, Sedbrook J, Rosen E, Caspar T, Masson P. The arabidopsis thaliana AGRAVITROPIC 1 gene encodes a component of the polar-auxin-transport efflux carrier. Proc Natl Acad Sci U S A. 1998;95:15112-7 pubmed
    ..1996) Current Biol. 6, 1589-1591]. Here we show that mutations in the Arabidopsis thaliana AGRAVITROPIC 1 (AGR1) gene involved in root gravitropism confer increased root-growth sensitivity to auxin and decreased sensitivity to ..
  32. Sukumar P, Edwards K, Rahman A, DeLong A, Muday G. PINOID kinase regulates root gravitropism through modulation of PIN2-dependent basipetal auxin transport in Arabidopsis. Plant Physiol. 2009;150:722-35 pubmed publisher
    ..Furthermore, the effect of staurosporine is lost in pinformed2 (pin2)...
  33. Leitner J, Luschnig C. Ubiquitylation-mediated control of polar auxin transport: analysis of Arabidopsis PIN2 auxin transport protein. Methods Mol Biol. 2014;1209:233-49 pubmed publisher
    ..demonstrated ubiquitylation of a number of plasma membrane proteins, including the auxin efflux carrier protein PIN2. By using solubilized membrane protein immunoprecipitation assays, we established quantitative approaches, suitable ..
  34. Sun J, Chen Q, Qi L, Jiang H, Li S, Xu Y, et al. Jasmonate modulates endocytosis and plasma membrane accumulation of the Arabidopsis PIN2 protein. New Phytol. 2011;191:360-75 pubmed publisher
    ..of CORONATINE INSENSITIVE 1 (COI1)- and AUXIN RESISTANT 1 (AXR1)-dependent methyl jasmonate (MeJA) effects on PIN2 subcellular distribution: at lower concentration (5 ?M), MeJA inhibits PIN2 endocytosis, whereas, at higher ..
  35. Moschou P, Smertenko A, Minina E, Fukada K, Savenkov E, Robert S, et al. The caspase-related protease separase (extra spindle poles) regulates cell polarity and cytokinesis in Arabidopsis. Plant Cell. 2013;25:2171-86 pubmed publisher
    ..Separase facilitates polar targeting of the auxin efflux carrier PIN-formed2 (PIN2) to the rootward side of the root cortex cells...
  36. Pandya Kumar N, Shema R, Kumar M, Mayzlish Gati E, Levy D, Zemach H, et al. Strigolactone analog GR24 triggers changes in PIN2 polarity, vesicle trafficking and actin filament architecture. New Phytol. 2014;202:1184-96 pubmed publisher
    ..the effects of GR24 (a synthetic, biologically active SL analog) on localization of the auxin efflux transporter PIN2, endosomal trafficking, and F-actin architecture and dynamics in the plasma membrane (PM) of epidermal cells of the ..
  37. Santelia D, Henrichs S, Vincenzetti V, Sauer M, Bigler L, Klein M, et al. Flavonoids redirect PIN-mediated polar auxin fluxes during root gravitropic responses. J Biol Chem. 2008;283:31218-26 pubmed publisher
    ..Here we show that the root elongation zone of agravitropic pin2/eir1/wav6/agr1 has an altered pattern and amount of flavonol glycosides...
  38. Tsugeki R, Ditengou F, Sumi Y, Teale W, Palme K, Okada K. NO VEIN mediates auxin-dependent specification and patterning in the Arabidopsis embryo, shoot, and root. Plant Cell. 2009;21:3133-51 pubmed publisher
    ..expression of PIN7 in leaf primordia, cell type-specific expression of PIN3, PIN4, and PIN7 in the root, and PIN2 polarity in the root cortex...
  39. Armengot L, Marquès Bueno M, Soria Garcia A, Müller M, Munné Bosch S, Martinez M. Functional interplay between protein kinase CK2 and salicylic acid sustains PIN transcriptional expression and root development. Plant J. 2014;78:411-23 pubmed publisher
    ..Thus, unlike the outcome in mammals, an excess of CK2 in plant cells does not produce neoplasia, but it might be advantageous for plant fitness. ..
  40. Li G, Song H, Li B, Kronzucker H, Shi W. Auxin Resistant1 and PIN-FORMED2 Protect Lateral Root Formation in Arabidopsis under Iron Stress. Plant Physiol. 2015;169:2608-23 pubmed publisher
    ..Furthermore, Fe stress significantly reduced PIN-FORMED2 (PIN2)-green fluorescent protein (GFP) expression in root tips, and pin2-1 mutants exhibited significantly fewer LR ..
  41. Rigó G, Ayaydin F, Tietz O, Zsigmond L, Kovacs H, Pay A, et al. Inactivation of plasma membrane-localized CDPK-RELATED KINASE5 decelerates PIN2 exocytosis and root gravitropic response in Arabidopsis. Plant Cell. 2013;25:1592-608 pubmed publisher
    ..Whereas AUX1, PIN1, PIN3, PIN4, and PIN7 display normal localization, PIN2 is depleted from apical membranes of epidermal cells and shows basal to apical relocalization in the cortex of the ..
  42. Chen Z, Noir S, Kwaaitaal M, Hartmann H, Wu M, Mudgil Y, et al. Two seven-transmembrane domain MILDEW RESISTANCE LOCUS O proteins cofunction in Arabidopsis root thigmomorphogenesis. Plant Cell. 2009;21:1972-91 pubmed publisher
    ..Moreover, addition of auxin transport inhibitors or the loss of EIR1/AGR1/PIN2 function abolishes root curling of mlo4, mlo11, and wild-type seedlings...
  43. Kleine Vehn J, Wabnik K, Martinière A, Łangowski Ł, Willig K, Naramoto S, et al. Recycling, clustering, and endocytosis jointly maintain PIN auxin carrier polarity at the plasma membrane. Mol Syst Biol. 2011;7:540 pubmed publisher
    ..Moreover, our study suggests that the regulation of lateral diffusion and spatially defined endocytosis, but not super-polar exocytosis have primary importance for PIN polarity maintenance. ..
  44. Hanzawa T, Shibasaki K, Numata T, Kawamura Y, Gaude T, Rahman A. Cellular auxin homeostasis under high temperature is regulated through a sorting NEXIN1-dependent endosomal trafficking pathway. Plant Cell. 2013;25:3424-33 pubmed publisher
    ..counterbalance the elevated level of intracellular auxin by promoting shootward auxin efflux in a PIN-FORMED2 (PIN2)-dependent manner...
  45. Belin C, Megies C, Hauserová E, Lopez Molina L. Abscisic acid represses growth of the Arabidopsis embryonic axis after germination by enhancing auxin signaling. Plant Cell. 2009;21:2253-68 pubmed publisher
    ..We show that aux1 and pin2 mutants are insensitive to ABA-dependent repression of embryonic axis (hypocotyl and radicle) elongation...
  46. Blakeslee J, Bandyopadhyay A, Lee O, Mravec J, Titapiwatanakun B, Sauer M, et al. Interactions among PIN-FORMED and P-glycoprotein auxin transporters in Arabidopsis. Plant Cell. 2007;19:131-47 pubmed
    ..Here, we show that PGP1 and PGP19 colocalized with PIN1 in the shoot apex in Arabidopsis thaliana and with PIN1 and PIN2 in root tissues. Specific PGP-PIN interactions were seen in yeast two-hybrid and coimmunoprecipitation assays...
  47. Mishra B, Singh M, Aggrawal P, Laxmi A. Glucose and auxin signaling interaction in controlling Arabidopsis thaliana seedlings root growth and development. PLoS ONE. 2009;4:e4502 pubmed publisher
    ..Our findings implicate an important role of the glucose interacting with auxin signaling and transport machinery to control seedling root growth and development in changing nutrient conditions. ..