[{"language":[{"iso":"eng"}],"isi":1,"acknowledgement":"The authors sincerely thank Dr Barbara Kloeckener Gruissem’s time and efforts in critical reading and constructive advice on the manuscript. The authors gratefully acknowledge Dr. Eva Sundberg for generously providing transgenic plants to support this study.\r\nThis work was supported by the European Research Council Advanced Grant (ETAP-742985 to H.T. and J.F.) and the Taiwan National Science and Technology Council (NSTC 112-2311-B-005-008 to H.T. and L.-H.C.).","date_created":"2025-03-19T09:44:19Z","article_type":"original","year":"2025","date_published":"2025-03-05T00:00:00Z","month":"03","abstract":[{"text":"Auxin and its PIN-FORMED (PIN) exporters are essential for tissue repair and regeneration in flowering plants. To gain insight into the evolution of this mechanism, we investigated their roles in leaves excised from Physcomitrium patens, a bryophyte known for its remarkable cell reprogramming capacity. We used various approaches to manipulate auxin levels, including exogenous application, pharmacological manipulations, and auxin biosynthesis mutants. We observed no significant effect on the rate of cell reprogramming. Rather, our analysis of auxin dynamics revealed a decrease in auxin levels upon excision, which was followed by a local increase before the reprogramming process began. Mutant analysis revealed that PpPINs are required for effective cell reprogramming, and endogenously expressed PpPINA-GFP accumulates polarly at sites that will develop into future filamentous stem cells. In addition, hyperpolarized PpPINA variants carrying mutated phosphorylation sites showed a marked delay in reprogramming, whereas endogenous or nonpolar versions do not have this effect. These results underscore that both the levels and the polarity of PpPINA are important for efficient cell reprogramming. Overall, these findings highlight the pivotal role of PIN polarity in plant regeneration. Furthermore, they suggest that understanding polarity mechanisms could have broader implications for improving regenerative processes across various plant species.","lang":"eng"}],"project":[{"_id":"261099A6-B435-11E9-9278-68D0E5697425","call_identifier":"H2020","grant_number":"742985","name":"Tracing Evolution of Auxin Transport and Polarity in Plants"}],"author":[{"last_name":"Tang","first_name":"Han","id":"19BDF720-25A0-11EA-AC6E-928F3DDC885E","full_name":"Tang, Han","orcid":"0000-0001-6152-6637"},{"full_name":"Chen, L","first_name":"L","last_name":"Chen"},{"last_name":"Friml","first_name":"Jiří","id":"4159519E-F248-11E8-B48F-1D18A9856A87","orcid":"0000-0002-8302-7596","full_name":"Friml, Jiří"}],"pmid":1,"article_number":"pcaf008","publication_status":"published","corr_author":"1","external_id":{"isi":["001436802900001"],"pmid":["39829340"]},"department":[{"_id":"JiFr"}],"publisher":"Oxford University Press","citation":{"ama":"Tang H, Chen L, Friml J. Auxin fluctuation and PIN polarization in moss leaf cell reprogramming. <i>Plant and Cell Physiology</i>. 2025. doi:<a href=\"https://doi.org/10.1093/pcp/pcaf008\">10.1093/pcp/pcaf008</a>","apa":"Tang, H., Chen, L., &#38; Friml, J. (2025). Auxin fluctuation and PIN polarization in moss leaf cell reprogramming. <i>Plant and Cell Physiology</i>. Oxford University Press. <a href=\"https://doi.org/10.1093/pcp/pcaf008\">https://doi.org/10.1093/pcp/pcaf008</a>","mla":"Tang, Han, et al. “Auxin Fluctuation and PIN Polarization in Moss Leaf Cell Reprogramming.” <i>Plant and Cell Physiology</i>, pcaf008, Oxford University Press, 2025, doi:<a href=\"https://doi.org/10.1093/pcp/pcaf008\">10.1093/pcp/pcaf008</a>.","ieee":"H. Tang, L. Chen, and J. Friml, “Auxin fluctuation and PIN polarization in moss leaf cell reprogramming.,” <i>Plant and Cell Physiology</i>. Oxford University Press, 2025.","short":"H. Tang, L. Chen, J. Friml, Plant and Cell Physiology (2025).","ista":"Tang H, Chen L, Friml J. 2025. Auxin fluctuation and PIN polarization in moss leaf cell reprogramming. Plant and Cell Physiology., pcaf008.","chicago":"Tang, Han, L Chen, and Jiří Friml. “Auxin Fluctuation and PIN Polarization in Moss Leaf Cell Reprogramming.” <i>Plant and Cell Physiology</i>. Oxford University Press, 2025. <a href=\"https://doi.org/10.1093/pcp/pcaf008\">https://doi.org/10.1093/pcp/pcaf008</a>."},"publication":"Plant and Cell Physiology","title":"Auxin fluctuation and PIN polarization in moss leaf cell reprogramming.","_id":"19420","OA_type":"closed access","date_updated":"2025-09-30T11:05:55Z","day":"05","ec_funded":1,"status":"public","quality_controlled":"1","publication_identifier":{"eissn":["1471-9053"],"issn":["0032-0781"]},"doi":"10.1093/pcp/pcaf008","user_id":"317138e5-6ab7-11ef-aa6d-ffef3953e345","oa_version":"None","scopus_import":"1","article_processing_charge":"No","type":"journal_article"},{"article_type":"original","year":"2022","date_published":"2022-01-21T00:00:00Z","date_created":"2021-12-28T11:44:18Z","acknowledgement":"The authors thank Ralf Stracke (Bielefeld University, Bielefeld, Germany) for providing the myb mutants and their colleagues Bert De Rybel for the tmo5t;mo5l1 double mutant, Boris Parizot for tips on the RNA-seq analysis, Veronique Storme for statistical help on both the RNA-seq and lateral root density, and Martine De Cock for help in preparing the manuscript.","oa":1,"issue":"1","isi":1,"language":[{"iso":"eng"}],"intvolume":"        63","abstract":[{"lang":"eng","text":"The synthetic strigolactone (SL) analog, rac-GR24, has been instrumental in studying the role of SLs as well as karrikins because it activates the receptors DWARF14 (D14) and KARRIKIN INSENSITIVE 2 (KAI2) of their signaling pathways, respectively. Treatment with rac-GR24 modifies the root architecture at different levels, such as decreasing the lateral root density (LRD), while promoting root hair elongation or flavonol accumulation. Previously, we have shown that the flavonol biosynthesis is transcriptionally activated in the root by rac-GR24 treatment, but, thus far, the molecular players involved in that response have remained unknown. To get an in-depth insight into the changes that occur after the compound is perceived by the roots, we compared the root transcriptomes of the wild type and the more axillary growth2 (max2) mutant, affected in both SL and karrikin signaling pathways, with and without rac-GR24 treatment. Quantitative reverse transcription (qRT)-PCR, reporter line analysis and mutant phenotyping indicated that the flavonol response and the root hair elongation are controlled by the ELONGATED HYPOCOTYL 5 (HY5) and MYB12 transcription factors, but HY5, in contrast to MYB12, affects the LRD as well. Furthermore, we identified the transcription factors TARGET OF MONOPTEROS 5 (TMO5) and TMO5 LIKE1 as negative and the Mediator complex as positive regulators of the rac-GR24 effect on LRD. Altogether, hereby, we get closer toward understanding the molecular mechanisms that underlay the rac-GR24 responses in the root."}],"month":"01","volume":63,"external_id":{"pmid":["34791413"],"isi":["000877899400009"]},"pmid":1,"publication_status":"published","author":[{"full_name":"Struk, Sylwia","last_name":"Struk","first_name":"Sylwia"},{"full_name":"Braem, Lukas","last_name":"Braem","first_name":"Lukas"},{"full_name":"Matthys, Cedrick","last_name":"Matthys","first_name":"Cedrick"},{"first_name":"Alan","last_name":"Walton","full_name":"Walton, Alan"},{"full_name":"Vangheluwe, Nick","first_name":"Nick","last_name":"Vangheluwe"},{"first_name":"Stan","last_name":"Van Praet","full_name":"Van Praet, Stan"},{"last_name":"Jiang","first_name":"Lingxiang","full_name":"Jiang, Lingxiang"},{"id":"3028BD74-F248-11E8-B48F-1D18A9856A87","full_name":"Baster, Pawel","last_name":"Baster","first_name":"Pawel"},{"full_name":"De Cuyper, Carolien","first_name":"Carolien","last_name":"De Cuyper"},{"full_name":"Boyer, Francois-Didier","first_name":"Francois-Didier","last_name":"Boyer"},{"last_name":"Stes","first_name":"Elisabeth","full_name":"Stes, Elisabeth"},{"last_name":"Beeckman","first_name":"Tom","full_name":"Beeckman, Tom"},{"last_name":"Friml","first_name":"Jiří","id":"4159519E-F248-11E8-B48F-1D18A9856A87","orcid":"0000-0002-8302-7596","full_name":"Friml, Jiří"},{"first_name":"Kris","last_name":"Gevaert","full_name":"Gevaert, Kris"},{"full_name":"Goormachtig, Sofie","last_name":"Goormachtig","first_name":"Sofie"}],"_id":"10583","title":"Transcriptional analysis in the Arabidopsis roots reveals new regulators that link rac-GR24 treatment with changes in flavonol accumulation, root hair elongation and lateral root density","citation":{"short":"S. Struk, L. Braem, C. Matthys, A. Walton, N. Vangheluwe, S. Van Praet, L. Jiang, P. Baster, C. De Cuyper, F.-D. Boyer, E. Stes, T. Beeckman, J. Friml, K. Gevaert, S. Goormachtig, Plant &#38; Cell Physiology 63 (2022) 104–119.","chicago":"Struk, Sylwia, Lukas Braem, Cedrick Matthys, Alan Walton, Nick Vangheluwe, Stan Van Praet, Lingxiang Jiang, et al. “Transcriptional Analysis in the Arabidopsis Roots Reveals New Regulators That Link Rac-GR24 Treatment with Changes in Flavonol Accumulation, Root Hair Elongation and Lateral Root Density.” <i>Plant &#38; Cell Physiology</i>. Oxford University Press, 2022. <a href=\"https://doi.org/10.1093/pcp/pcab149\">https://doi.org/10.1093/pcp/pcab149</a>.","ista":"Struk S, Braem L, Matthys C, Walton A, Vangheluwe N, Van Praet S, Jiang L, Baster P, De Cuyper C, Boyer F-D, Stes E, Beeckman T, Friml J, Gevaert K, Goormachtig S. 2022. Transcriptional analysis in the Arabidopsis roots reveals new regulators that link rac-GR24 treatment with changes in flavonol accumulation, root hair elongation and lateral root density. Plant &#38; Cell Physiology. 63(1), 104–119.","ama":"Struk S, Braem L, Matthys C, et al. Transcriptional analysis in the Arabidopsis roots reveals new regulators that link rac-GR24 treatment with changes in flavonol accumulation, root hair elongation and lateral root density. <i>Plant &#38; Cell Physiology</i>. 2022;63(1):104-119. doi:<a href=\"https://doi.org/10.1093/pcp/pcab149\">10.1093/pcp/pcab149</a>","ieee":"S. Struk <i>et al.</i>, “Transcriptional analysis in the Arabidopsis roots reveals new regulators that link rac-GR24 treatment with changes in flavonol accumulation, root hair elongation and lateral root density,” <i>Plant &#38; Cell Physiology</i>, vol. 63, no. 1. Oxford University Press, pp. 104–119, 2022.","apa":"Struk, S., Braem, L., Matthys, C., Walton, A., Vangheluwe, N., Van Praet, S., … Goormachtig, S. (2022). Transcriptional analysis in the Arabidopsis roots reveals new regulators that link rac-GR24 treatment with changes in flavonol accumulation, root hair elongation and lateral root density. <i>Plant &#38; Cell Physiology</i>. Oxford University Press. <a href=\"https://doi.org/10.1093/pcp/pcab149\">https://doi.org/10.1093/pcp/pcab149</a>","mla":"Struk, Sylwia, et al. “Transcriptional Analysis in the Arabidopsis Roots Reveals New Regulators That Link Rac-GR24 Treatment with Changes in Flavonol Accumulation, Root Hair Elongation and Lateral Root Density.” <i>Plant &#38; Cell Physiology</i>, vol. 63, no. 1, Oxford University Press, 2022, pp. 104–19, doi:<a href=\"https://doi.org/10.1093/pcp/pcab149\">10.1093/pcp/pcab149</a>."},"keyword":["flavonols","MAX2","rac-Gr24","RNA-seq","root development","transcriptional regulation"],"publication":"Plant & Cell Physiology","publisher":"Oxford University Press","department":[{"_id":"JiFr"}],"main_file_link":[{"open_access":"1","url":"https://doi.org/10.1093/pcp/pcab149"}],"day":"21","date_updated":"2026-06-18T08:43:19Z","page":"104-119","user_id":"2DF688A6-F248-11E8-B48F-1D18A9856A87","scopus_import":"1","oa_version":"Published Version","quality_controlled":"1","doi":"10.1093/pcp/pcab149","publication_identifier":{"eissn":["1471-9053"],"issn":["0032-0781"]},"status":"public","type":"journal_article","article_processing_charge":"No","ddc":["580"]},{"volume":60,"abstract":[{"text":"Abiotic stress poses constant challenges for plant survival and is a serious problem for global agricultural productivity. On a molecular level, stress conditions result in elevation of reactive oxygen species (ROS) production causing oxidative stress associated with oxidation of proteins and nucleic acids as well as impairment of membrane functions. Adaptation of root growth to ROS accumulation is facilitated through modification of auxin and cytokinin hormone homeostasis. Here, we report that in Arabidopsis root meristem, ROS-induced changes of auxin levels correspond to decreased abundance of PIN auxin efflux carriers at the plasma membrane (PM). Specifically, increase in H2O2 levels affects PIN2 endocytic recycling. We show that the PIN2 intracellular trafficking during adaptation to oxidative stress requires the function of the ADP-ribosylation factor (ARF)-guanine-nucleotide exchange factor (GEF) BEN1, an actin-associated regulator of the trafficking from the PM to early endosomes and, presumably, indirectly, trafficking to the vacuoles. We propose that H2O2 levels affect the actin dynamics thus modulating ARF-GEF-dependent trafficking of PIN2. This mechanism provides a way how root growth acclimates to stress and adapts to a changing environment.","lang":"eng"}],"month":"02","issue":"2","language":[{"iso":"eng"}],"isi":1,"intvolume":"        60","year":"2019","date_published":"2019-02-01T00:00:00Z","date_created":"2019-03-17T22:59:14Z","author":[{"last_name":"Zwiewka","first_name":"Marta","full_name":"Zwiewka, Marta"},{"full_name":"Bielach, Agnieszka","last_name":"Bielach","first_name":"Agnieszka"},{"last_name":"Tamizhselvan","first_name":"Prashanth","full_name":"Tamizhselvan, Prashanth"},{"first_name":"Sharmila","last_name":"Madhavan","full_name":"Madhavan, Sharmila"},{"full_name":"Ryad, Eman Elrefaay","last_name":"Ryad","first_name":"Eman Elrefaay"},{"first_name":"Shutang","last_name":"Tan","full_name":"Tan, Shutang","orcid":"0000-0002-0471-8285","id":"2DE75584-F248-11E8-B48F-1D18A9856A87"},{"first_name":"Mónika","last_name":"Hrtyan","full_name":"Hrtyan, Mónika","id":"45A71A74-F248-11E8-B48F-1D18A9856A87"},{"full_name":"Dobrev, Petre","last_name":"Dobrev","first_name":"Petre"},{"last_name":"Vanková","first_name":"Radomira","full_name":"Vanková, Radomira"},{"last_name":"Friml","first_name":"Jiří","id":"4159519E-F248-11E8-B48F-1D18A9856A87","orcid":"0000-0002-8302-7596","full_name":"Friml, Jiří"},{"last_name":"Tognetti","first_name":"Vanesa B.","full_name":"Tognetti, Vanesa B."}],"external_id":{"isi":["000459634300002"],"pmid":["30668780"]},"pmid":1,"publication_status":"published","date_updated":"2023-08-25T08:05:28Z","day":"01","citation":{"short":"M. Zwiewka, A. Bielach, P. Tamizhselvan, S. Madhavan, E.E. Ryad, S. Tan, M. Hrtyan, P. Dobrev, R. Vanková, J. Friml, V.B. Tognetti, Plant and Cell Physiology 60 (2019) 255–273.","chicago":"Zwiewka, Marta, Agnieszka Bielach, Prashanth Tamizhselvan, Sharmila Madhavan, Eman Elrefaay Ryad, Shutang Tan, Mónika Hrtyan, et al. “Root Adaptation to H2O2-Induced Oxidative Stress by ARF-GEF BEN1- and Cytoskeleton-Mediated PIN2 Trafficking.” <i>Plant and Cell Physiology</i>. Oxford University Press, 2019. <a href=\"https://doi.org/10.1093/pcp/pcz001\">https://doi.org/10.1093/pcp/pcz001</a>.","ista":"Zwiewka M, Bielach A, Tamizhselvan P, Madhavan S, Ryad EE, Tan S, Hrtyan M, Dobrev P, Vanková R, Friml J, Tognetti VB. 2019. Root adaptation to H2O2-induced oxidative stress by ARF-GEF BEN1- and cytoskeleton-mediated PIN2 trafficking. Plant and Cell Physiology. 60(2), 255–273.","ama":"Zwiewka M, Bielach A, Tamizhselvan P, et al. Root adaptation to H2O2-induced oxidative stress by ARF-GEF BEN1- and cytoskeleton-mediated PIN2 trafficking. <i>Plant and Cell Physiology</i>. 2019;60(2):255-273. doi:<a href=\"https://doi.org/10.1093/pcp/pcz001\">10.1093/pcp/pcz001</a>","ieee":"M. Zwiewka <i>et al.</i>, “Root adaptation to H2O2-induced oxidative stress by ARF-GEF BEN1- and cytoskeleton-mediated PIN2 trafficking,” <i>Plant and Cell Physiology</i>, vol. 60, no. 2. Oxford University Press, pp. 255–273, 2019.","apa":"Zwiewka, M., Bielach, A., Tamizhselvan, P., Madhavan, S., Ryad, E. E., Tan, S., … Tognetti, V. B. (2019). Root adaptation to H2O2-induced oxidative stress by ARF-GEF BEN1- and cytoskeleton-mediated PIN2 trafficking. <i>Plant and Cell Physiology</i>. Oxford University Press. <a href=\"https://doi.org/10.1093/pcp/pcz001\">https://doi.org/10.1093/pcp/pcz001</a>","mla":"Zwiewka, Marta, et al. “Root Adaptation to H2O2-Induced Oxidative Stress by ARF-GEF BEN1- and Cytoskeleton-Mediated PIN2 Trafficking.” <i>Plant and Cell Physiology</i>, vol. 60, no. 2, Oxford University Press, 2019, pp. 255–73, doi:<a href=\"https://doi.org/10.1093/pcp/pcz001\">10.1093/pcp/pcz001</a>."},"publication":"Plant and Cell Physiology","publisher":"Oxford University Press","department":[{"_id":"JiFr"}],"title":"Root adaptation to H2O2-induced oxidative stress by ARF-GEF BEN1- and cytoskeleton-mediated PIN2 trafficking","_id":"6104","type":"journal_article","article_processing_charge":"No","quality_controlled":"1","publication_identifier":{"issn":["0032-0781"],"eissn":["1471-9053"]},"doi":"10.1093/pcp/pcz001","status":"public","page":"255-273","scopus_import":"1","oa_version":"None","user_id":"4359f0d1-fa6c-11eb-b949-802e58b17ae8"},{"_id":"799","title":"BEN3/BIG2 ARF GEF is involved in brefeldin a-sensitive trafficking at the trans-Golgi network/early endosome in Arabidopsis thaliana","file":[{"relation":"main_file","date_created":"2019-04-17T07:52:34Z","access_level":"open_access","creator":"dernst","file_size":1352913,"content_type":"application/pdf","checksum":"bd3e3a94d55416739cbb19624bb977f8","file_id":"6333","file_name":"2017_PlantCellPhysio_Kitakura.pdf","date_updated":"2020-07-14T12:48:06Z"}],"department":[{"_id":"JiFr"}],"publisher":"Oxford University Press","publication":"Plant and Cell Physiology","has_accepted_license":"1","citation":{"mla":"Kitakura, Saeko, et al. “BEN3/BIG2 ARF GEF Is Involved in Brefeldin a-Sensitive Trafficking at the Trans-Golgi Network/Early Endosome in Arabidopsis Thaliana.” <i>Plant and Cell Physiology</i>, vol. 58, no. 10, 1801–1811, Oxford University Press, 2017, doi:<a href=\"https://doi.org/10.1093/pcp/pcx118\">10.1093/pcp/pcx118</a>.","apa":"Kitakura, S., Adamowski, M., Matsuura, Y., Santuari, L., Kouno, H., Arima, K., … Tanaka, H. (2017). BEN3/BIG2 ARF GEF is involved in brefeldin a-sensitive trafficking at the trans-Golgi network/early endosome in Arabidopsis thaliana. <i>Plant and Cell Physiology</i>. Oxford University Press. <a href=\"https://doi.org/10.1093/pcp/pcx118\">https://doi.org/10.1093/pcp/pcx118</a>","ieee":"S. Kitakura <i>et al.</i>, “BEN3/BIG2 ARF GEF is involved in brefeldin a-sensitive trafficking at the trans-Golgi network/early endosome in Arabidopsis thaliana,” <i>Plant and Cell Physiology</i>, vol. 58, no. 10. Oxford University Press, 2017.","ama":"Kitakura S, Adamowski M, Matsuura Y, et al. BEN3/BIG2 ARF GEF is involved in brefeldin a-sensitive trafficking at the trans-Golgi network/early endosome in Arabidopsis thaliana. <i>Plant and Cell Physiology</i>. 2017;58(10). doi:<a href=\"https://doi.org/10.1093/pcp/pcx118\">10.1093/pcp/pcx118</a>","ista":"Kitakura S, Adamowski M, Matsuura Y, Santuari L, Kouno H, Arima K, Hardtke C, Friml J, Kakimoto T, Tanaka H. 2017. BEN3/BIG2 ARF GEF is involved in brefeldin a-sensitive trafficking at the trans-Golgi network/early endosome in Arabidopsis thaliana. Plant and Cell Physiology. 58(10), 1801–1811.","chicago":"Kitakura, Saeko, Maciek Adamowski, Yuki Matsuura, Luca Santuari, Hirotaka Kouno, Kohei Arima, Christian Hardtke, Jiří Friml, Tatsuo Kakimoto, and Hirokazu Tanaka. “BEN3/BIG2 ARF GEF Is Involved in Brefeldin a-Sensitive Trafficking at the Trans-Golgi Network/Early Endosome in Arabidopsis Thaliana.” <i>Plant and Cell Physiology</i>. Oxford University Press, 2017. <a href=\"https://doi.org/10.1093/pcp/pcx118\">https://doi.org/10.1093/pcp/pcx118</a>.","short":"S. Kitakura, M. Adamowski, Y. Matsuura, L. Santuari, H. Kouno, K. Arima, C. Hardtke, J. Friml, T. Kakimoto, H. Tanaka, Plant and Cell Physiology 58 (2017)."},"day":"21","date_updated":"2025-07-10T11:54:55Z","user_id":"2DF688A6-F248-11E8-B48F-1D18A9856A87","oa_version":"Submitted Version","scopus_import":"1","status":"public","publication_identifier":{"issn":["0032-0781"]},"doi":"10.1093/pcp/pcx118","quality_controlled":"1","article_processing_charge":"No","type":"journal_article","ddc":["581"],"date_created":"2018-12-11T11:48:34Z","date_published":"2017-08-21T00:00:00Z","file_date_updated":"2020-07-14T12:48:06Z","publist_id":"6854","year":"2017","intvolume":"        58","language":[{"iso":"eng"}],"oa":1,"issue":"10","isi":1,"month":"08","abstract":[{"text":"Membrane traffic at the trans-Golgi network (TGN) is crucial for correctly distributing various membrane proteins to their destination. Polarly localized auxin efflux proteins, including PIN-FORMED1 (PIN1), are dynamically transported between the endosomes and the plasma membrane (PM) in the plant cells. The intracellular trafficking of PIN1 protein is sensitive to a fungal toxin brefeldin A (BFA), which is known to inhibit guanine-nucleotide exchange factors for ADP ribosylation factors (ARF GEFs) such as GNOM. However, the molecular details of the BFA-sensitive trafficking pathway have not been revealed fully. In a previous study, we have identified an Arabidopsis mutant BFA-visualized endocytic trafficking defective 3 (ben3) which exhibited reduced sensitivity to BFA in terms of BFA-induced intracellular PIN1 agglomeration. Here, we show that BEN3 encodes a member of BIG family ARF GEFs, BIG2. Fluorescent proteins tagged BEN3/BIG2 co-localized with markers for TGN / early endosome (EE). Inspection of conditionally induced de novo synthesized PIN1 confirmed that its secretion to the PM is BFA-sensitive and established BEN3/BIG2 as a crucial component of this BFA action at the level of TGN/EE. Furthermore, ben3 mutation alleviated BFA-induced agglomeration of another TGN-localized ARF GEF BEN1/MIN7. Taken together our results suggest that BEN3/BIG2 is an ARF GEF component, which confers BFA sensitivity to the TGN/EE in Arabidopsis.","lang":"eng"}],"volume":58,"publication_status":"published","article_number":"1801-1811","pubrep_id":"1009","pmid":1,"external_id":{"pmid":["29016942"],"isi":["000413220400019"]},"author":[{"full_name":"Kitakura, Saeko","last_name":"Kitakura","first_name":"Saeko"},{"first_name":"Maciek","last_name":"Adamowski","full_name":"Adamowski, Maciek","orcid":"0000-0001-6463-5257","id":"45F536D2-F248-11E8-B48F-1D18A9856A87"},{"full_name":"Matsuura, Yuki","first_name":"Yuki","last_name":"Matsuura"},{"full_name":"Santuari, Luca","first_name":"Luca","last_name":"Santuari"},{"first_name":"Hirotaka","last_name":"Kouno","full_name":"Kouno, Hirotaka"},{"last_name":"Arima","first_name":"Kohei","full_name":"Arima, Kohei"},{"first_name":"Christian","last_name":"Hardtke","full_name":"Hardtke, Christian"},{"id":"4159519E-F248-11E8-B48F-1D18A9856A87","orcid":"0000-0002-8302-7596","full_name":"Friml, Jirí","last_name":"Friml","first_name":"Jirí"},{"first_name":"Tatsuo","last_name":"Kakimoto","full_name":"Kakimoto, Tatsuo"},{"full_name":"Tanaka, Hirokazu","last_name":"Tanaka","first_name":"Hirokazu"}]},{"author":[{"full_name":"Johnson, Kaeli C.M.","first_name":"Kaeli C.M.","last_name":"Johnson"},{"full_name":"Xia, Shitou","first_name":"Shitou","last_name":"Xia"},{"first_name":"Xiaoqi","last_name":"Feng","orcid":"0000-0002-4008-1234","full_name":"Feng, Xiaoqi","id":"e0164712-22ee-11ed-b12a-d80fcdf35958"},{"full_name":"Li, Xin","last_name":"Li","first_name":"Xin"}],"external_id":{"pmid":["26063389"]},"publication_status":"published","pmid":1,"acknowledgement":"This work was supported by the National Sciences and Engineering Research Council of Canada [Canada Graduate\r\nScholarship–Doctoral to K.J.; Discovery Grant to X.L.]; the department of Botany at the University of f British Columbia\r\n[the Dewar Cooper Memorial Fund to X.L.].The authors would like to thank Dr. Yuelin Zhang and Ms. Yan Li for their assistance with next-generation sequencing, and Mr. Charles Copeland for critical reading of the manuscript.","language":[{"iso":"eng"}],"issue":"8","intvolume":"        56","date_published":"2015-08-01T00:00:00Z","article_type":"original","year":"2015","date_created":"2023-01-16T09:20:22Z","volume":56,"abstract":[{"text":"SNC1 (SUPPRESSOR OF NPR1, CONSTITUTIVE 1) is one of a suite of intracellular Arabidopsis NOD-like receptor (NLR) proteins which, upon activation, result in the induction of defense responses. However, the molecular mechanisms underlying NLR activation and the subsequent provocation of immune responses are only partially characterized. To identify negative regulators of NLR-mediated immunity, a forward genetic screen was undertaken to search for enhancers of the dwarf, autoimmune gain-of-function snc1 mutant. To avoid lethality resulting from severe dwarfism, the screen was conducted using mos4 (modifier of snc1, 4) snc1 plants, which display wild-type-like morphology and resistance. M2 progeny were screened for mutant, snc1-enhancing (muse) mutants displaying a reversion to snc1-like phenotypes. The muse9 mos4 snc1 triple mutant was found to exhibit dwarf morphology, elevated expression of the pPR2-GUS defense marker reporter gene and enhanced resistance to the oomycete pathogen Hyaloperonospora arabidopsidis Noco2. Via map-based cloning and Illumina sequencing, it was determined that the muse9 mutation is in the gene encoding the SWI/SNF chromatin remodeler SYD (SPLAYED), and was thus renamed syd-10. The syd-10 single mutant has no observable alteration from wild-type-like resistance, although the syd-4 T-DNA insertion allele displays enhanced resistance to the bacterial pathogen Pseudomonas syringae pv. maculicola ES4326. Transcription of SNC1 is increased in both syd-4 and syd-10. These data suggest that SYD plays a subtle, specific role in the regulation of SNC1 expression and SNC1-mediated immunity. SYD may work with other proteins at the chromatin level to repress SNC1 transcription; such regulation is important for fine-tuning the expression of NLR-encoding genes to prevent unpropitious autoimmunity.","lang":"eng"}],"month":"08","doi":"10.1093/pcp/pcv087","publication_identifier":{"issn":["0032-0781","1471-9053"]},"quality_controlled":"1","status":"public","scopus_import":"1","oa_version":"None","user_id":"2DF688A6-F248-11E8-B48F-1D18A9856A87","page":"1616-1623","type":"journal_article","extern":"1","article_processing_charge":"No","publication":"Plant and Cell Physiology","citation":{"ama":"Johnson KCM, Xia S, Feng X, Li X. The chromatin remodeler SPLAYED negatively regulates SNC1-mediated immunity. <i>Plant and Cell Physiology</i>. 2015;56(8):1616-1623. doi:<a href=\"https://doi.org/10.1093/pcp/pcv087\">10.1093/pcp/pcv087</a>","mla":"Johnson, Kaeli C. M., et al. “The Chromatin Remodeler SPLAYED Negatively Regulates SNC1-Mediated Immunity.” <i>Plant and Cell Physiology</i>, vol. 56, no. 8, Oxford University Press, 2015, pp. 1616–23, doi:<a href=\"https://doi.org/10.1093/pcp/pcv087\">10.1093/pcp/pcv087</a>.","apa":"Johnson, K. C. M., Xia, S., Feng, X., &#38; Li, X. (2015). The chromatin remodeler SPLAYED negatively regulates SNC1-mediated immunity. <i>Plant and Cell Physiology</i>. Oxford University Press. <a href=\"https://doi.org/10.1093/pcp/pcv087\">https://doi.org/10.1093/pcp/pcv087</a>","ieee":"K. C. M. Johnson, S. Xia, X. Feng, and X. Li, “The chromatin remodeler SPLAYED negatively regulates SNC1-mediated immunity,” <i>Plant and Cell Physiology</i>, vol. 56, no. 8. Oxford University Press, pp. 1616–1623, 2015.","short":"K.C.M. Johnson, S. Xia, X. Feng, X. Li, Plant and Cell Physiology 56 (2015) 1616–1623.","ista":"Johnson KCM, Xia S, Feng X, Li X. 2015. The chromatin remodeler SPLAYED negatively regulates SNC1-mediated immunity. Plant and Cell Physiology. 56(8), 1616–1623.","chicago":"Johnson, Kaeli C.M., Shitou Xia, Xiaoqi Feng, and Xin Li. “The Chromatin Remodeler SPLAYED Negatively Regulates SNC1-Mediated Immunity.” <i>Plant and Cell Physiology</i>. Oxford University Press, 2015. <a href=\"https://doi.org/10.1093/pcp/pcv087\">https://doi.org/10.1093/pcp/pcv087</a>."},"keyword":["Cell Biology","Plant Science","Physiology","General Medicine"],"publisher":"Oxford University Press","department":[{"_id":"XiFe"}],"title":"The chromatin remodeler SPLAYED negatively regulates SNC1-mediated immunity","_id":"12196","date_updated":"2023-05-08T11:03:23Z"},{"author":[{"full_name":"Naramoto, Satoshi","first_name":"Satoshi","last_name":"Naramoto"},{"last_name":"Nodzyński","first_name":"Tomasz","full_name":"Nodzyński, Tomasz"},{"last_name":"Dainobu","first_name":"Tomoko","full_name":"Dainobu, Tomoko"},{"full_name":"Takatsuka, Hirotomo","last_name":"Takatsuka","first_name":"Hirotomo"},{"full_name":"Okada, Teruyo","last_name":"Okada","first_name":"Teruyo"},{"id":"4159519E-F248-11E8-B48F-1D18A9856A87","full_name":"Friml, Jirí","orcid":"0000-0002-8302-7596","last_name":"Friml","first_name":"Jirí"},{"first_name":"Hiroo","last_name":"Fukuda","full_name":"Fukuda, Hiroo"}],"external_id":{"isi":["000334679500009"]},"publication_status":"published","project":[{"_id":"25716A02-B435-11E9-9278-68D0E5697425","grant_number":"282300","call_identifier":"FP7","name":"Polarity and subcellular dynamics in plants"}],"volume":55,"abstract":[{"lang":"eng","text":"Leaf venation develops complex patterns in angiosperms, but the mechanism underlying this process is largely unknown. To elucidate the molecular mechanisms governing vein pattern formation, we previously isolated vascular network defective (van) mutants that displayed venation discontinuities. Here, we report the phenotypic analysis of van4 mutants, and we identify and characterize the VAN4 gene. Detailed phenotypic analysis shows that van4 mutants are defective in procambium cell differentiation and subsequent vascular cell differentiation. Reduced shoot and root cell growth is observed in van4 mutants, suggesting that VAN4 function is important for cell growth and the establishment of venation continuity. Consistent with these phenotypes, the VAN4 gene is strongly expressed in vascular and meristematic cells. VAN4 encodes a putative TRS120, which is a known guanine nucleotide exchange factor (GEF) for Rab GTPase involved in regulating vesicle transport, and a known tethering factor that determines the specificity of membrane fusion. VAN4 protein localizes at the trans-Golgi network/early endosome (TGN/EE). Aberrant recycling of the auxin efflux carrier PIN proteins is observed in van4 mutants. These results suggest that VAN4-mediated exocytosis at the TGN plays important roles in plant vascular development and cell growth in shoot and root. Our identification of VAN4 as a putative TRS120 shows that Rab GTPases are crucial (in addition to ARF GTPases) for continuous vascular development, and provides further evidence for the importance of vesicle transport in leaf vascular formation."}],"month":"04","isi":1,"language":[{"iso":"eng"}],"issue":"4","intvolume":"        55","publist_id":"4742","year":"2014","date_published":"2014-04-01T00:00:00Z","date_created":"2018-12-11T11:56:24Z","type":"journal_article","article_processing_charge":"No","quality_controlled":"1","doi":"10.1093/pcp/pcu012","publication_identifier":{"issn":["0032-0781"]},"ec_funded":1,"status":"public","page":"750 - 763","scopus_import":"1","oa_version":"None","user_id":"317138e5-6ab7-11ef-aa6d-ffef3953e345","date_updated":"2025-09-29T11:28:27Z","day":"01","citation":{"ieee":"S. Naramoto <i>et al.</i>, “VAN4 encodes a putative TRS120 that is required for normal cell growth and vein development in arabidopsis,” <i>Plant and Cell Physiology</i>, vol. 55, no. 4. Oxford University Press, pp. 750–763, 2014.","apa":"Naramoto, S., Nodzyński, T., Dainobu, T., Takatsuka, H., Okada, T., Friml, J., &#38; Fukuda, H. (2014). VAN4 encodes a putative TRS120 that is required for normal cell growth and vein development in arabidopsis. <i>Plant and Cell Physiology</i>. Oxford University Press. <a href=\"https://doi.org/10.1093/pcp/pcu012\">https://doi.org/10.1093/pcp/pcu012</a>","mla":"Naramoto, Satoshi, et al. “VAN4 Encodes a Putative TRS120 That Is Required for Normal Cell Growth and Vein Development in Arabidopsis.” <i>Plant and Cell Physiology</i>, vol. 55, no. 4, Oxford University Press, 2014, pp. 750–63, doi:<a href=\"https://doi.org/10.1093/pcp/pcu012\">10.1093/pcp/pcu012</a>.","ama":"Naramoto S, Nodzyński T, Dainobu T, et al. VAN4 encodes a putative TRS120 that is required for normal cell growth and vein development in arabidopsis. <i>Plant and Cell Physiology</i>. 2014;55(4):750-763. doi:<a href=\"https://doi.org/10.1093/pcp/pcu012\">10.1093/pcp/pcu012</a>","chicago":"Naramoto, Satoshi, Tomasz Nodzyński, Tomoko Dainobu, Hirotomo Takatsuka, Teruyo Okada, Jiří Friml, and Hiroo Fukuda. “VAN4 Encodes a Putative TRS120 That Is Required for Normal Cell Growth and Vein Development in Arabidopsis.” <i>Plant and Cell Physiology</i>. Oxford University Press, 2014. <a href=\"https://doi.org/10.1093/pcp/pcu012\">https://doi.org/10.1093/pcp/pcu012</a>.","ista":"Naramoto S, Nodzyński T, Dainobu T, Takatsuka H, Okada T, Friml J, Fukuda H. 2014. VAN4 encodes a putative TRS120 that is required for normal cell growth and vein development in arabidopsis. Plant and Cell Physiology. 55(4), 750–763.","short":"S. Naramoto, T. Nodzyński, T. Dainobu, H. Takatsuka, T. Okada, J. Friml, H. Fukuda, Plant and Cell Physiology 55 (2014) 750–763."},"publication":"Plant and Cell Physiology","department":[{"_id":"JiFr"}],"publisher":"Oxford University Press","_id":"2222","title":"VAN4 encodes a putative TRS120 that is required for normal cell growth and vein development in arabidopsis"},{"intvolume":"        55","language":[{"iso":"eng"}],"oa":1,"issue":"4","isi":1,"date_created":"2018-12-11T11:56:25Z","publist_id":"4741","year":"2014","file_date_updated":"2020-07-14T12:45:34Z","date_published":"2014-04-01T00:00:00Z","volume":55,"month":"04","abstract":[{"lang":"eng","text":"Correct positioning of membrane proteins is an essential process in eukaryotic organisms. The plant hormone auxin is distributed through intercellular transport and triggers various cellular responses. Auxin transporters of the PIN-FORMED (PIN) family localize asymmetrically at the plasma membrane (PM) and mediate the directional transport of auxin between cells. A fungal toxin, brefeldin A (BFA), inhibits a subset of guanine nucleotide exchange factors for ADP-ribosylation factor small GTPases (ARF GEFs) including GNOM, which plays a major role in localization of PIN1 predominantly to the basal side of the PM. The Arabidopsis genome encodes 19 ARF-related putative GTPases. However, ARF components involved in PIN1 localization have been genetically poorly defined. Using a fluorescence imaging-based forward genetic approach, we identified an Arabidopsis mutant, bfa-visualized exocytic trafficking defective1 (bex1), in which PM localization of PIN1-green fluorescent protein (GFP) as well as development is hypersensitive to BFA. We found that in bex1 a member of the ARF1 gene family, ARF1A1C, was mutated. ARF1A1C localizes to the trans-Golgi network/early endosome and Golgi apparatus, acts synergistically to BEN1/MIN7 ARF GEF and is important for PIN recycling to the PM. Consistent with the developmental importance of PIN proteins, functional interference with ARF1 resulted in an impaired auxin response gradient and various developmental defects including embryonic patterning defects and growth arrest. Our results show that ARF1A1C is essential for recycling of PIN auxin transporters and for various auxin-dependent developmental processes."}],"project":[{"_id":"25716A02-B435-11E9-9278-68D0E5697425","grant_number":"282300","call_identifier":"FP7","name":"Polarity and subcellular dynamics in plants"},{"name":"Innovationsförderung in der Grenzregion Österreich – Tschechische Republik durch die Schaffung von Synergien im Bereich der Forschungsinfrastruktur","_id":"256BDAB0-B435-11E9-9278-68D0E5697425"}],"author":[{"full_name":"Tanaka, Hirokazu","first_name":"Hirokazu","last_name":"Tanaka"},{"first_name":"Tomasz","last_name":"Nodzyński","full_name":"Nodzyński, Tomasz"},{"first_name":"Saeko","last_name":"Kitakura","full_name":"Kitakura, Saeko"},{"first_name":"Mugurel","last_name":"Feraru","full_name":"Feraru, Mugurel"},{"first_name":"Michiko","last_name":"Sasabe","full_name":"Sasabe, Michiko"},{"full_name":"Ishikawa, Tomomi","last_name":"Ishikawa","first_name":"Tomomi"},{"first_name":"Jürgen","last_name":"Kleine Vehn","full_name":"Kleine Vehn, Jürgen"},{"full_name":"Kakimoto, Tatsuo","first_name":"Tatsuo","last_name":"Kakimoto"},{"last_name":"Friml","first_name":"Jirí","id":"4159519E-F248-11E8-B48F-1D18A9856A87","orcid":"0000-0002-8302-7596","full_name":"Friml, Jirí"}],"tmp":{"legal_code_url":"https://creativecommons.org/licenses/by-nc/4.0/legalcode","image":"/images/cc_by_nc.png","name":"Creative Commons Attribution-NonCommercial 4.0 International (CC BY-NC 4.0)","short":"CC BY-NC (4.0)"},"pmid":1,"pubrep_id":"431","publication_status":"published","external_id":{"isi":["000334679500008"],"pmid":["24369434"]},"department":[{"_id":"JiFr"}],"publisher":"Oxford University Press","file":[{"file_size":2028111,"content_type":"application/pdf","creator":"system","access_level":"open_access","relation":"main_file","date_created":"2018-12-12T10:14:25Z","date_updated":"2020-07-14T12:45:34Z","file_name":"IST-2016-431-v1+1_Plant_Cell_Physiol-2014-Tanaka-737-49.pdf","file_id":"5076","checksum":"b781a76b32ac35a520256453c3ba9433"}],"citation":{"short":"H. Tanaka, T. Nodzyński, S. Kitakura, M. Feraru, M. Sasabe, T. Ishikawa, J. Kleine Vehn, T. Kakimoto, J. Friml, Plant and Cell Physiology 55 (2014) 737–749.","chicago":"Tanaka, Hirokazu, Tomasz Nodzyński, Saeko Kitakura, Mugurel Feraru, Michiko Sasabe, Tomomi Ishikawa, Jürgen Kleine Vehn, Tatsuo Kakimoto, and Jiří Friml. “BEX1/ARF1A1C Is Required for BFA-Sensitive Recycling of PIN Auxin Transporters and Auxin-Mediated Development in Arabidopsis.” <i>Plant and Cell Physiology</i>. Oxford University Press, 2014. <a href=\"https://doi.org/10.1093/pcp/pct196\">https://doi.org/10.1093/pcp/pct196</a>.","ista":"Tanaka H, Nodzyński T, Kitakura S, Feraru M, Sasabe M, Ishikawa T, Kleine Vehn J, Kakimoto T, Friml J. 2014. BEX1/ARF1A1C is required for BFA-sensitive recycling of PIN auxin transporters and auxin-mediated development in arabidopsis. Plant and Cell Physiology. 55(4), 737–749.","ama":"Tanaka H, Nodzyński T, Kitakura S, et al. BEX1/ARF1A1C is required for BFA-sensitive recycling of PIN auxin transporters and auxin-mediated development in arabidopsis. <i>Plant and Cell Physiology</i>. 2014;55(4):737-749. doi:<a href=\"https://doi.org/10.1093/pcp/pct196\">10.1093/pcp/pct196</a>","ieee":"H. Tanaka <i>et al.</i>, “BEX1/ARF1A1C is required for BFA-sensitive recycling of PIN auxin transporters and auxin-mediated development in arabidopsis,” <i>Plant and Cell Physiology</i>, vol. 55, no. 4. Oxford University Press, pp. 737–749, 2014.","mla":"Tanaka, Hirokazu, et al. “BEX1/ARF1A1C Is Required for BFA-Sensitive Recycling of PIN Auxin Transporters and Auxin-Mediated Development in Arabidopsis.” <i>Plant and Cell Physiology</i>, vol. 55, no. 4, Oxford University Press, 2014, pp. 737–49, doi:<a href=\"https://doi.org/10.1093/pcp/pct196\">10.1093/pcp/pct196</a>.","apa":"Tanaka, H., Nodzyński, T., Kitakura, S., Feraru, M., Sasabe, M., Ishikawa, T., … Friml, J. (2014). BEX1/ARF1A1C is required for BFA-sensitive recycling of PIN auxin transporters and auxin-mediated development in arabidopsis. <i>Plant and Cell Physiology</i>. Oxford University Press. <a href=\"https://doi.org/10.1093/pcp/pct196\">https://doi.org/10.1093/pcp/pct196</a>"},"publication":"Plant and Cell Physiology","has_accepted_license":"1","_id":"2223","title":"BEX1/ARF1A1C is required for BFA-sensitive recycling of PIN auxin transporters and auxin-mediated development in arabidopsis","date_updated":"2025-09-29T11:27:52Z","day":"01","status":"public","ec_funded":1,"quality_controlled":"1","publication_identifier":{"issn":["0032-0781"]},"doi":"10.1093/pcp/pct196","page":"737 - 749","oa_version":"Published Version","user_id":"317138e5-6ab7-11ef-aa6d-ffef3953e345","scopus_import":"1","ddc":["570"],"article_processing_charge":"No","type":"journal_article"}]