{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,9,30]],"date-time":"2025-09-30T00:07:39Z","timestamp":1759190859997},"reference-count":93,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2017,2,22]],"date-time":"2017-02-22T00:00:00Z","timestamp":1487721600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2017,2,22]],"date-time":"2017-02-22T00:00:00Z","timestamp":1487721600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"abstract":"<jats:title>Abstract<\/jats:title><jats:p>Sperm nuclear migration during fertilization in <jats:italic>Arabidopsis<\/jats:italic> and rice has recently been found to be actin-dependent, but the driving force behind this actin cytoskeleton-dependent motion is unclear. Here, we confirmed that the actin-dependent sperm nuclei migration during fertilization is a conserved mechanism in plants. Using <jats:italic>in vitro<\/jats:italic> fertilization systems, we showed that a functional actin is also essential in maize and tobacco for sperm nuclei migration after gamete membrane fusion. Cytoskeleton depolymerization inhibitor treatments supported the view that sperm nuclei migration is actin-dependent but microtubule-independent in both egg cell and central cell during double fertilization. We further revealed that the actin-based motor myosin is not the driving force for sperm nuclear migration in maize and tobacco. The WASP-Arp2\/3 complex signal cascade is shown here to be involved in the regulation of sperm nuclear migration in maize and tobacco. It is interesting that sperm nuclei migration within somatic cell also need WASP-Arp2\/3 complex signal cascade and actin, suggesting that the mechanism of sperm nuclear migration is not gamete specific.<\/jats:p>","DOI":"10.1038\/srep43161","type":"journal-article","created":{"date-parts":[[2017,2,22]],"date-time":"2017-02-22T11:37:32Z","timestamp":1487763452000},"update-policy":"http:\/\/dx.doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":16,"title":["The WASP-Arp2\/3 complex signal cascade is involved in actin-dependent sperm nuclei migration during double fertilization in tobacco and maize"],"prefix":"10.1038","volume":"7","author":[{"given":"Xiongbo","family":"Peng","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Tingting","family":"Yan","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Mengxiang","family":"Sun","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"297","published-online":{"date-parts":[[2017,2,22]]},"reference":[{"key":"BFsrep43161_CR1","unstructured":"Nawaschin, S. Resultate einer Revision der Befruchtungsvorgange bei Lilium martagon und Fritillaria tenella. \u0418\u0437\u0432\u0435\u0441\u0442\u0438\u044f\u0420\u043e\u0441\u0441\u0438\u0439\u0441\u043a\u043e\u0439 \u0430\u043a\u0430\u0434\u0435\u043c\u0438\u0438 \u043d\u0430\u0443\u043a. \u0421\u0435\u0440\u0438\u044f \u043c\u0430\u0442\u0435\u043c\u0430\u0442\u0438\u0447\u0435\u0441\u043a\u0430\u044f 9, 377\u2013382 (1898)."},{"key":"BFsrep43161_CR2","first-page":"129","volume":"11","author":"M Guignard","year":"1899","unstructured":"Guignard, M. Sur les antherozoides et la double copulation sexuelle chez les vegetaux angiosperms. Rev. Gen. Bot. 11, 129\u2013135 (1899).","journal-title":"Rev. Gen. Bot"},{"key":"BFsrep43161_CR3","doi-asserted-by":"crossref","first-page":"S107","DOI":"10.1105\/tpc.016873","volume":"16","author":"K Weterings","year":"2004","unstructured":"Weterings, K. & Russell, S. D. Experimental analysis of the fertilization process. Plant Cell 16 Suppl, S107\u2013118 (2004).","journal-title":"Plant Cell"},{"key":"BFsrep43161_CR4","doi-asserted-by":"crossref","first-page":"5139","DOI":"10.1093\/jxb\/erv275","volume":"66","author":"LJ Qu","year":"2015","unstructured":"Qu, L. J., Li, L., Lan, Z. & Dresselhaus, T. Peptide signalling during the pollen tube journey and double fertilization. J Exp Bot 66, 5139\u20135150 (2015).","journal-title":"J Exp Bot"},{"key":"BFsrep43161_CR5","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1016\/j.pbi.2015.10.006","volume":"28","author":"MM Kanaoka","year":"2015","unstructured":"Kanaoka, M. M. & Higashiyama, T. Peptide signaling in pollen tube guidance. Curr Opin Plant Biol 28, 127\u2013136 (2015).","journal-title":"Curr Opin Plant Biol"},{"key":"BFsrep43161_CR6","doi-asserted-by":"crossref","first-page":"907","DOI":"10.1016\/j.cell.2015.03.018","volume":"161","author":"D Maruyama","year":"2015","unstructured":"Maruyama, D. et al. Rapid Elimination of the Persistent Synergid through a Cell Fusion Mechanism. Cell 161, 907\u2013918 (2015).","journal-title":"Cell"},{"key":"BFsrep43161_CR7","doi-asserted-by":"crossref","first-page":"4722","DOI":"10.1038\/ncomms5722","volume":"5","author":"Y Hamamura","year":"2014","unstructured":"Hamamura, Y. et al. Live imaging of calcium spikes during double fertilization in Arabidopsis. Nat Commun 5, 4722 (2014).","journal-title":"Nat Commun"},{"key":"BFsrep43161_CR8","doi-asserted-by":"crossref","first-page":"4645","DOI":"10.1038\/ncomms5645","volume":"5","author":"P Denninger","year":"2014","unstructured":"Denninger, P. et al. Male-female communication triggers calcium signatures during fertilization in Arabidopsis. Nat Commun 5, 4645 (2014).","journal-title":"Nat Commun"},{"key":"BFsrep43161_CR9","doi-asserted-by":"crossref","unstructured":"Kawashima, T. et al. Dynamic F-actin movement is essential for fertilization in Arabidopsis thaliana. Elife 3 (2014).","DOI":"10.7554\/eLife.04501"},{"key":"BFsrep43161_CR10","doi-asserted-by":"crossref","first-page":"82","DOI":"10.1016\/S1360-1385(96)80039-2","volume":"1","author":"E Kranz","year":"1996","unstructured":"Kranz, E. & Dresselhaus, T. In vitro fertilization with isolated higher plant gametes. Trends plant sci 1, 82\u201389 (1996).","journal-title":"Trends plant sci"},{"key":"BFsrep43161_CR11","doi-asserted-by":"crossref","first-page":"178","DOI":"10.1007\/BF00193715","volume":"15","author":"M Kovacs","year":"1995","unstructured":"Kovacs, M., Barnabas, B. & Kranz, E. Electro-fused isolated wheat (Triticum aestivum L.) gametes develop into multicellular structures. Plant Cell Rep 15, 178\u2013180 (1995).","journal-title":"Plant Cell Rep"},{"key":"BFsrep43161_CR12","doi-asserted-by":"crossref","first-page":"125","DOI":"10.1017\/S0967199400001878","volume":"2","author":"E Kranz","year":"1994","unstructured":"Kranz, E. & Lorz, H. In vitro fertilisation of maize by single egg and sperm cell protoplast fusion mediated by high calcium and high pH. Zygote 2, 125\u2013128 (1994).","journal-title":"Zygote"},{"key":"BFsrep43161_CR13","doi-asserted-by":"crossref","first-page":"734","DOI":"10.1038\/sj.cr.7290343","volume":"15","author":"XB Peng","year":"2005","unstructured":"Peng, X. B., Sun, M. X. & Yang, H. Y. A novel in vitro system for gamete fusion in maize. Cell Res 15, 734\u2013738 (2005).","journal-title":"Cell Res"},{"key":"BFsrep43161_CR14","doi-asserted-by":"crossref","first-page":"1598","DOI":"10.1126\/science.263.5153.1598","volume":"263","author":"JE Faure","year":"1994","unstructured":"Faure, J. E., Digonnet, C. & Dumas, C. An in Vitro System for Adhesion and Fusion of Maize Gametes. Science 263, 1598\u20131600 (1994).","journal-title":"Science"},{"key":"BFsrep43161_CR15","doi-asserted-by":"crossref","first-page":"581","DOI":"10.1007\/s00425-007-0506-2","volume":"226","author":"T Uchiumi","year":"2007","unstructured":"Uchiumi, T., Uemura, I. & Okamoto, T. Establishment of an in vitro fertilization system in rice (Oryza sativa L.). Planta 226, 581\u2013589 (2007).","journal-title":"Planta"},{"key":"BFsrep43161_CR16","doi-asserted-by":"crossref","first-page":"657","DOI":"10.1007\/BF01275510","volume":"16","author":"HQ Tian","year":"1997","unstructured":"Tian, H. Q. & Russell, S. D. Micromanipulation of male and female gametes of Nicotiana tabacum: II. Preliminary attempts for in vitro fertilization and egg cell culture. Plant cell rep 16, 657\u2013661 (1997).","journal-title":"Plant cell rep"},{"key":"BFsrep43161_CR17","first-page":"12","volume":"4","author":"E Kranz","year":"1991","unstructured":"Kranz, E., Bautor, J. & L\u00f6rz, H. In vitro fertilization of single, isolated gametes of maize mediated by electrofusion. Sex Plant Reprod 4, 12\u201316 (1991).","journal-title":"Sex Plant Reprod"},{"key":"BFsrep43161_CR18","first-page":"489","volume":"36","author":"M Sun","year":"1995","unstructured":"Sun, M., Yang, H. & Zhou, C. Single-pair fusion of various combinations between female gametoplasts and other protoplasts in Nicotiana tabacum. Acta Bot Sin 36, 489\u201393 (1995).","journal-title":"Acta Bot Sin"},{"key":"BFsrep43161_CR19","doi-asserted-by":"crossref","first-page":"1120","DOI":"10.1038\/ncb1201-1120","volume":"3","author":"AF Antoine","year":"2001","unstructured":"Antoine, A. F., Faure, J. E., Dumas, C. & Feijo, J. A. Differential contribution of cytoplasmic Ca2+ and Ca2+ influx to gamete fusion and egg activation in maize. Nat Cell Biol 3, 1120\u20131123 (2001).","journal-title":"Nat Cell Biol"},{"key":"BFsrep43161_CR20","doi-asserted-by":"crossref","first-page":"10643","DOI":"10.1073\/pnas.180243697","volume":"97","author":"AF Antoine","year":"2000","unstructured":"Antoine, A. F. et al. A calcium influx is triggered and propagates in the zygote as a wavefront during in vitro fertilization of flowering plants. Proc Natl Acad Sci USA 97, 10643\u201310648 (2000).","journal-title":"Proc Natl Acad Sci USA"},{"key":"BFsrep43161_CR21","doi-asserted-by":"crossref","first-page":"2867","DOI":"10.1242\/dev.124.15.2867","volume":"124","author":"C Digonnet","year":"1997","unstructured":"Digonnet, C., Aldon, D., Leduc, N., Dumas, C. & Rougier, M. First evidence of a calcium transient in flowering plants at fertilization. Development 124, 2867\u20132874 (1997).","journal-title":"Development"},{"key":"BFsrep43161_CR22","doi-asserted-by":"crossref","first-page":"267","DOI":"10.1007\/s004970050194","volume":"12","author":"M-X Sun","year":"2000","unstructured":"Sun, M.-X., Moscatelli, A., Yang, H.-Y. & Cresti, M. In vitro double fertilization in Nicotiana tabacum (L.):polygamy compared with selected single pair somatic protoplastand chloroplast fusions. Sex Plant Reprod 12, 267\u2013275 (2000).","journal-title":"Sex Plant Reprod"},{"key":"BFsrep43161_CR23","doi-asserted-by":"crossref","first-page":"1051","DOI":"10.1046\/j.1365-313X.2003.01692.x","volume":"33","author":"JE Faure","year":"2003","unstructured":"Faure, J. E. et al. Double fertilization in maize: the two male gametes from a pollen grain have the ability to fuse with egg cells. Plant J 33, 1051\u20131062 (2003).","journal-title":"Plant J"},{"key":"BFsrep43161_CR24","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1046\/j.1365-313X.1995.08010009.x","volume":"8","author":"E Kranz","year":"1995","unstructured":"Kranz, E., Wiegen, P. & L\u00f6rz, H. Early cytological events after induction of cell division in egg cells and zygote development following in vitro fertilization with angiosperm gametes. Plant J 8, 9\u201323 (1995).","journal-title":"Plant J"},{"key":"BFsrep43161_CR25","doi-asserted-by":"crossref","first-page":"267","DOI":"10.1007\/s004970050194","volume":"12","author":"M-X Sun","year":"2000","unstructured":"Sun, M.-X., Moscatelli, A., Yang, H.-Y. & Cresti, M. In vitro double fertilization in Nicotiana tabacum (L.): fusion behavior and gamete interaction traced by video-enhanced microscopy. Sex Plant Reprod 12, 267\u2013275 (2000).","journal-title":"Sex Plant Reprod"},{"key":"BFsrep43161_CR26","doi-asserted-by":"crossref","first-page":"747","DOI":"10.2307\/3869612","volume":"5","author":"JE Faure","year":"1993","unstructured":"Faure, J. E., Mogensen, H. L., Dumas, C., Lorz, H. & Kranz, E. Karyogamy after Electrofusion of Single Egg and Sperm Cell Protoplasts from Maize: Cytological Evidence and Time Course. Plant Cell 5, 747\u2013755 (1993).","journal-title":"Plant Cell"},{"key":"BFsrep43161_CR27","doi-asserted-by":"crossref","first-page":"1533","DOI":"10.1104\/pp.114.236059","volume":"165","author":"Y Ohnishi","year":"2014","unstructured":"Ohnishi, Y., Hoshino, R. & Okamoto, T. Dynamics of Male and Female Chromatin during Karyogamy in Rice Zygotes. Plant Physiol 165, 1533\u20131543 (2014).","journal-title":"Plant Physiol"},{"key":"BFsrep43161_CR28","doi-asserted-by":"crossref","first-page":"e989021","DOI":"10.4161\/15592324.2014.989021","volume":"10","author":"Y Ohnishi","year":"2015","unstructured":"Ohnishi, Y. & Okamoto, T. Karyogamy in rice zygotes: Actin filament-dependent migration of sperm nucleus, chromatin dynamics, and de novo gene expression. Plant Signal Behav 10, e989021 (2015).","journal-title":"Plant Signal Behav"},{"key":"BFsrep43161_CR29","doi-asserted-by":"crossref","first-page":"455","DOI":"10.1083\/jcb.94.2.455","volume":"94","author":"G Schatten","year":"1982","unstructured":"Schatten, G., Schatten, H., Bestor, T. H. & Balczon, R. Taxol inhibits the nuclear movements during fertilization and induces asters in unfertilized sea urchin eggs. J Cell Biol 94, 455\u2013465 (1982).","journal-title":"J Cell Biol"},{"key":"BFsrep43161_CR30","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1016\/0092-8674(83)90203-9","volume":"35","author":"S Strome","year":"1983","unstructured":"Strome, S. & Wood, W. B. Generation of asymmetry and segregation of germ-line granules in early C. elegans embryos. Cell 35, 15\u201325 (1983).","journal-title":"Cell"},{"key":"BFsrep43161_CR31","doi-asserted-by":"crossref","first-page":"539","DOI":"10.1083\/jcb.69.3.539","volume":"69","author":"FJ Longo","year":"1976","unstructured":"Longo, F. J. Sperm aster in rabbit zygotes: its structure and function. J Cell Biol 69, 539\u2013547 (1976).","journal-title":"J Cell Biol"},{"key":"BFsrep43161_CR32","first-page":"239","volume":"48","author":"P Le Guen","year":"1989","unstructured":"Le Guen, P. & Crozet, N. Microtubule and centrosome distribution during sheep fertilization. Eur J Cell Biol 48, 239\u2013249 (1989).","journal-title":"Eur J Cell Biol"},{"key":"BFsrep43161_CR33","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1006\/dbio.1994.1064","volume":"162","author":"CS Navara","year":"1994","unstructured":"Navara, C. S., First, N. L. & Schatten, G. Microtubule organization in the cow during fertilization, polyspermy, parthenogenesis, and nuclear transfer: the role of the sperm aster. Dev Biol 162, 29\u201340 (1994).","journal-title":"Dev Biol"},{"key":"BFsrep43161_CR34","doi-asserted-by":"crossref","first-page":"481","DOI":"10.1002\/mrd.1113","volume":"60","author":"QY Sun","year":"2001","unstructured":"Sun, Q. Y. et al. Microtubule assembly after treatment of pig oocytes with taxol: correlation with chromosomes, gamma-tubulin, and MAP kinase. Mol Reprod Dev 60, 481\u2013490 (2001).","journal-title":"Mol Reprod Dev"},{"key":"BFsrep43161_CR35","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1530\/rep.1.00847","volume":"131","author":"QY Sun","year":"2006","unstructured":"Sun, Q. Y. & Schatten, H. Regulation of dynamic events by microfilaments during oocyte maturation and fertilization. Reproduction 131, 193\u2013205 (2006).","journal-title":"Reproduction"},{"key":"BFsrep43161_CR36","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1104\/pp.125.1.119","volume":"125","author":"CR Hawes","year":"2001","unstructured":"Hawes, C. R. & Satiat-Jeunemaitre, B. Trekking along the cytoskeleton. Plant Physiol 125, 119\u2013122 (2001).","journal-title":"Plant Physiol"},{"key":"BFsrep43161_CR37","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1111\/jipb.12289","volume":"57","author":"G Cai","year":"2015","unstructured":"Cai, G., Parrotta, L. & Cresti, M. Organelle trafficking, the cytoskeleton, and pollen tube growth. J Integr Plant Biol 57, 63\u201378 (2015).","journal-title":"J Integr Plant Biol"},{"key":"BFsrep43161_CR38","doi-asserted-by":"crossref","first-page":"19534","DOI":"10.1074\/jbc.M611802200","volume":"282","author":"K Ito","year":"2007","unstructured":"Ito, K. et al. Kinetic mechanism of the fastest motor protein, Chara myosin. J Biol Chem 282, 19534\u201319545 (2007).","journal-title":"J Biol Chem"},{"key":"BFsrep43161_CR39","doi-asserted-by":"crossref","first-page":"21585","DOI":"10.1073\/pnas.0910787106","volume":"106","author":"K Ito","year":"2009","unstructured":"Ito, K., Yamaguchi, Y., Yanase, K., Ichikawa, Y. & Yamamoto, K. Unique charge distribution in surface loops confers high velocity on the fast motor protein Chara myosin. Proc Natl Acad Sci USA 106, 21585\u201321590 (2009).","journal-title":"Proc Natl Acad Sci USA"},{"key":"BFsrep43161_CR40","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1111\/jipb.12282","volume":"57","author":"L Buchnik","year":"2015","unstructured":"Buchnik, L., Abu-Abied, M. & Sadot, E. Role of plant myosins in motile organelles: is a direct interaction required? J Integr Plant Biol 57, 23\u201330 (2015).","journal-title":"J Integr Plant Biol"},{"key":"BFsrep43161_CR41","doi-asserted-by":"crossref","first-page":"571","DOI":"10.1093\/pcp\/41.5.571","volume":"41","author":"J \u0160amaj","year":"2000","unstructured":"\u0160amaj, J., Peters, M., Volkmann, D. & Balu\u0161ka, F. Effects of myosin ATPase inhibitor 2, 3-butanedione 2-monoxime on distributions of myosins, F-actin, microtubules, and cortical endoplasmic reticulum in maize root apices. Plant Cell Physiol 41, 571\u2013582 (2000).","journal-title":"Plant Cell Physiol"},{"key":"BFsrep43161_CR42","doi-asserted-by":"crossref","first-page":"613","DOI":"10.1046\/j.1365-313X.1997.11030613.x","volume":"11","author":"RH K\u00f6hler","year":"1997","unstructured":"K\u00f6hler, R. H., Zipfel, W. R., Webb, W. W. & Hanson, M. R. The green fluorescent protein as a marker to visualize plant mitochondria in vivo. Plant J 11, 613\u2013621 (1997).","journal-title":"Plant J"},{"key":"BFsrep43161_CR43","doi-asserted-by":"crossref","first-page":"659","DOI":"10.1093\/jexbot\/53.369.659","volume":"53","author":"K Van Gestel","year":"2002","unstructured":"Van Gestel, K., Kohler, R. H. & Verbelen, J. P. Plant mitochondria move on F-actin, but their positioning in the cortical cytoplasm depends on both F-actin and microtubules. J Exp Bot 53, 659\u2013667 (2002).","journal-title":"J Exp Bot"},{"key":"BFsrep43161_CR44","doi-asserted-by":"crossref","first-page":"606","DOI":"10.1002\/cm.20379","volume":"66","author":"AS Paul","year":"2009","unstructured":"Paul, A. S. & Pollard, T. D. Review of the mechanism of processive actin filament elongation by formins. Cell Motil Cytoskeleton 66, 606\u2013617 (2009).","journal-title":"Cell Motil Cytoskeleton"},{"key":"BFsrep43161_CR45","doi-asserted-by":"crossref","first-page":"451","DOI":"10.1146\/annurev.biophys.35.040405.101936","volume":"36","author":"TD Pollard","year":"2007","unstructured":"Pollard, T. D. Regulation of actin filament assembly by Arp2\/3 complex and formins. Annu Rev Biophys Biomol Struct 36, 451\u2013477 (2007).","journal-title":"Annu Rev Biophys Biomol Struct"},{"key":"BFsrep43161_CR46","doi-asserted-by":"crossref","first-page":"1007","DOI":"10.1038\/35010008","volume":"404","author":"L Blanchoin","year":"2000","unstructured":"Blanchoin, L. et al. Direct observation of dendritic actin filament networks nucleated by Arp2\/3 complex and WASP\/Scar proteins. Nature 404, 1007\u20131011 (2000).","journal-title":"Nature"},{"key":"BFsrep43161_CR47","doi-asserted-by":"crossref","first-page":"713","DOI":"10.1038\/nrm2026","volume":"7","author":"ED Goley","year":"2006","unstructured":"Goley, E. D. & Welch, M. D. The ARP2\/3 complex: an actin nucleator comes of age. Nat Rev Mol Cell Biol 7, 713\u2013726 (2006).","journal-title":"Nat Rev Mol Cell Biol"},{"key":"BFsrep43161_CR48","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1242\/jcs.107250","volume":"126","author":"D Breitsprecher","year":"2013","unstructured":"Breitsprecher, D. & Goode, B. L. Formins at a glance. J Cell Sci 126, 1\u20137 (2013).","journal-title":"J Cell Sci"},{"key":"BFsrep43161_CR49","doi-asserted-by":"crossref","first-page":"800","DOI":"10.1111\/j.1744-7909.2012.01160.x","volume":"54","author":"Y Zheng","year":"2012","unstructured":"Zheng, Y., Xin, H., Lin, J., Liu, C. M. & Huang, S. An Arabidopsis class II formin, AtFH19, nucleates actin assembly, binds to the barbed end of actin filaments, and antagonizes the effect of AtFH1 on actin dynamics. J Integr Plant Biol 54, 800\u2013813 (2012).","journal-title":"J Integr Plant Biol"},{"key":"BFsrep43161_CR50","doi-asserted-by":"crossref","first-page":"1002","DOI":"10.1111\/jipb.12089","volume":"55","author":"J Wang","year":"2013","unstructured":"Wang, J., Zhang, Y., Wu, J., Meng, L. & Ren, H. AtFH16, An Arabidopsis type II formin, binds and bundles both microfilaments and microtubules, and preferentially binds to microtubules. J Integr Plant Biol 55, 1002\u20131015 (2013).","journal-title":"J Integr Plant Biol"},{"key":"BFsrep43161_CR51","doi-asserted-by":"crossref","first-page":"900","DOI":"10.1016\/j.molp.2016.03.006","volume":"9","author":"S Zhang","year":"2016","unstructured":"Zhang, S. et al. A Processive Arabidopsis Formin Modulates Actin Filament Dynamics in Association with Profilin. Mol Plant 9, 900\u2013910 (2016).","journal-title":"Mol Plant"},{"key":"BFsrep43161_CR52","doi-asserted-by":"crossref","first-page":"1347","DOI":"10.1016\/S0960-9822(98)00015-3","volume":"8","author":"LM Machesky","year":"1998","unstructured":"Machesky, L. M. & Insall, R. H. Scar1 and the related Wiskott-Aldrich syndrome protein, WASP, regulate the actin cytoskeleton through the Arp2\/3 complex. Curr Biol 8, 1347\u20131356 (1998).","journal-title":"Curr Biol"},{"key":"BFsrep43161_CR53","doi-asserted-by":"crossref","first-page":"3739","DOI":"10.1073\/pnas.96.7.3739","volume":"96","author":"LM Machesky","year":"1999","unstructured":"Machesky, L. M. et al. Scar, a WASp-related protein, activates nucleation of actin filaments by the Arp2\/3 complex. Proc Natl Acad Sci USA 96, 3739\u20133744 (1999).","journal-title":"Proc Natl Acad Sci USA"},{"key":"BFsrep43161_CR54","doi-asserted-by":"crossref","first-page":"221","DOI":"10.1016\/S0092-8674(00)80732-1","volume":"97","author":"R Rohatgi","year":"1999","unstructured":"Rohatgi, R. et al. The interaction between N-WASP and the Arp2\/3 complex links Cdc42-dependent signals to actin assembly. Cell 97, 221\u2013231 (1999).","journal-title":"Cell"},{"key":"BFsrep43161_CR55","doi-asserted-by":"crossref","first-page":"4","DOI":"10.1016\/j.ceb.2005.12.003","volume":"18","author":"TE Stradal","year":"2006","unstructured":"Stradal, T. E. & Scita, G. Protein complexes regulating Arp2\/3-mediated actin assembly. Curr Opin Cell Biol 18, 4\u201310 (2006).","journal-title":"Curr Opin Cell Biol"},{"key":"BFsrep43161_CR56","doi-asserted-by":"crossref","first-page":"747","DOI":"10.1038\/nsmb796","volume":"11","author":"JR Peterson","year":"2004","unstructured":"Peterson, J. R. et al. Chemical inhibition of N-WASP by stabilization of a native autoinhibited conformation. Nat Struct Mol Biol 11, 747\u2013755 (2004).","journal-title":"Nat Struct Mol Biol"},{"key":"BFsrep43161_CR57","doi-asserted-by":"crossref","first-page":"10624","DOI":"10.1073\/pnas.201393198","volume":"98","author":"JR Peterson","year":"2001","unstructured":"Peterson, J. R., Lokey, R. S., Mitchison, T. J. & Kirschner, M. W. A chemical inhibitor of N-WASP reveals a new mechanism for targeting protein interactions. Proc Natl Acad Sci U S A 98, 10624\u201310629 (2001).","journal-title":"Proc Natl Acad Sci U S A"},{"key":"BFsrep43161_CR58","doi-asserted-by":"crossref","first-page":"120","DOI":"10.1038\/nature10914","volume":"484","author":"T Metzger","year":"2012","unstructured":"Metzger, T. et al. MAP and kinesin-dependent nuclear positioning is required for skeletal muscle function. Nature 484, 120\u2013124 (2012).","journal-title":"Nature"},{"key":"BFsrep43161_CR59","doi-asserted-by":"crossref","first-page":"2733","DOI":"10.1091\/mbc.11.8.2733","volume":"11","author":"E Chytilova","year":"2000","unstructured":"Chytilova, E. et al. Nuclear dynamics in Arabidopsis thaliana. Mol Biol Cell 11, 2733\u20132741 (2000).","journal-title":"Mol Biol Cell"},{"key":"BFsrep43161_CR60","doi-asserted-by":"crossref","first-page":"2283","DOI":"10.1242\/jcs.111.16.2283","volume":"111","author":"S Reinsch","year":"1998","unstructured":"Reinsch, S. & Gonczy, P. Mechanisms of nuclear positioning. J Cell Sci 111 (Pt 16), 2283\u20132295 (1998).","journal-title":"J Cell Sci"},{"key":"BFsrep43161_CR61","doi-asserted-by":"crossref","first-page":"384","DOI":"10.1093\/pcp\/pcf045","volume":"43","author":"G Jedd","year":"2002","unstructured":"Jedd, G. & Chua, N. H. Visualization of peroxisomes in living plant cells reveals acto-myosin-dependent cytoplasmic streaming and peroxisome budding. Plant Cell Physiol 43, 384\u2013392 (2002).","journal-title":"Plant Cell Physiol"},{"key":"BFsrep43161_CR62","doi-asserted-by":"crossref","first-page":"331","DOI":"10.1093\/pcp\/pcf037","volume":"43","author":"S Mano","year":"2002","unstructured":"Mano, S. et al. Distribution and characterization of peroxisomes in Arabidopsis by visualization with GFP: dynamic morphology and actin-dependent movement. Plant Cell Physiol 43, 331\u2013341 (2002).","journal-title":"Plant Cell Physiol"},{"key":"BFsrep43161_CR63","doi-asserted-by":"crossref","first-page":"1127","DOI":"10.1104\/pp.121.4.1127","volume":"121","author":"A Nebenfuhr","year":"1999","unstructured":"Nebenfuhr, A. et al. Stop-and-go movements of plant Golgi stacks are mediated by the acto-myosin system. Plant Physiol 121, 1127\u20131142 (1999).","journal-title":"Plant Physiol"},{"key":"BFsrep43161_CR64","doi-asserted-by":"crossref","first-page":"1405","DOI":"10.1016\/j.cub.2004.06.062","volume":"14","author":"D El-Assal Sel","year":"2004","unstructured":"El-Assal Sel, D., Le, J., Basu, D., Mallery, E. L. & Szymanski, D. B. Arabidopsis GNARLED encodes a NAP125 homolog that positively regulates ARP2\/3. Curr Biol 14, 1405\u20131409 (2004).","journal-title":"Curr Biol"},{"key":"BFsrep43161_CR65","doi-asserted-by":"crossref","first-page":"40","DOI":"10.1111\/jipb.12293","volume":"57","author":"S Huang","year":"2015","unstructured":"Huang, S., Qu, X. & Zhang, R. Plant villins: versatile actin regulatory proteins. J Integr Plant Biol 57, 40\u201349 (2015).","journal-title":"J Integr Plant Biol"},{"key":"BFsrep43161_CR66","doi-asserted-by":"crossref","first-page":"1776","DOI":"10.1016\/j.cub.2013.07.035","volume":"23","author":"K Tamura","year":"2013","unstructured":"Tamura, K. et al. Myosin XI-i links the nuclear membrane to the cytoskeleton to control nuclear movement and shape in Arabidopsis. Curr Biol 23, 1776\u20131781 (2013).","journal-title":"Curr Biol"},{"key":"BFsrep43161_CR67","doi-asserted-by":"crossref","first-page":"290","DOI":"10.1007\/s00438-004-1052-2","volume":"272","author":"I Zimmermann","year":"2004","unstructured":"Zimmermann, I., Saedler, R., Mutondo, M. & Hulskamp, M. The Arabidopsis GNARLED gene encodes the NAP125 homolog and controls several actin-based cell shape changes. Mol Genet Genomics 272, 290\u2013296 (2004).","journal-title":"Mol Genet Genomics"},{"key":"BFsrep43161_CR68","doi-asserted-by":"crossref","first-page":"237","DOI":"10.1038\/nrm2867","volume":"11","author":"KG Campellone","year":"2010","unstructured":"Campellone, K. G. & Welch, M. D. A nucleator arms race: cellular control of actin assembly. Nat Rev Mol Cell Biol 11, 237\u2013251 (2010).","journal-title":"Nat Rev Mol Cell Biol"},{"key":"BFsrep43161_CR69","doi-asserted-by":"crossref","first-page":"536","DOI":"10.1016\/j.chom.2009.10.011","volume":"6","author":"MP Dodding","year":"2009","unstructured":"Dodding, M. P. & Way, M. Nck- and N-WASP-dependent actin-based motility is conserved in divergent vertebrate poxviruses. Cell Host Microbe 6, 536\u2013550 (2009).","journal-title":"Cell Host Microbe"},{"key":"BFsrep43161_CR70","doi-asserted-by":"crossref","first-page":"850","DOI":"10.1093\/embo-reports\/kve197","volume":"2","author":"S Lommel","year":"2001","unstructured":"Lommel, S. et al. Actin pedestal formation by enteropathogenic Escherichia coli and intracellular motility of Shigella flexneri are abolished in N-WASP-defective cells. EMBO Rep 2, 850\u2013857 (2001).","journal-title":"EMBO Rep"},{"key":"BFsrep43161_CR71","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1038\/nrm3492","volume":"14","author":"JD Rotty","year":"2013","unstructured":"Rotty, J. D., Wu, C. & Bear, J. E. New insights into the regulation and cellular functions of the ARP2\/3 complex. Nat Rev Mol Cell Biol 14, 7\u201312 (2013).","journal-title":"Nat Rev Mol Cell Biol"},{"key":"BFsrep43161_CR72","doi-asserted-by":"crossref","first-page":"897","DOI":"10.1038\/ncb1001-897","volume":"3","author":"SB Snapper","year":"2001","unstructured":"Snapper, S. B. et al. N-WASP deficiency reveals distinct pathways for cell surface projections and microbial actin-based motility. Nat Cell Biol 3, 897\u2013904 (2001).","journal-title":"Nat Cell Biol"},{"key":"BFsrep43161_CR73","doi-asserted-by":"crossref","first-page":"87","DOI":"10.1038\/nature07773","volume":"458","author":"I Weisswange","year":"2009","unstructured":"Weisswange, I., Newsome, T. P., Schleich, S. & Way, M. The rate of N-WASP exchange limits the extent of ARP2\/3-complex-dependent actin-based motility. Nature 458, 87\u201391 (2009).","journal-title":"Nature"},{"key":"BFsrep43161_CR74","doi-asserted-by":"crossref","first-page":"220","DOI":"10.1016\/j.tcb.2008.03.001","volume":"18","author":"A Lambrechts","year":"2008","unstructured":"Lambrechts, A., Gevaert, K., Cossart, P., Vandekerckhove, J. & Van Troys, M. Listeria comet tails: the actin-based motility machinery at work. Trends Cell Biol 18, 220\u2013227 (2008).","journal-title":"Trends Cell Biol"},{"key":"BFsrep43161_CR75","doi-asserted-by":"crossref","first-page":"265","DOI":"10.1038\/385265a0","volume":"385","author":"MD Welch","year":"1997","unstructured":"Welch, M. D., Iwamatsu, A. & Mitchison, T. J. Actin polymerization is induced by Arp2\/3 protein complex at the surface of Listeria monocytogenes. Nature 385, 265\u2013269 (1997).","journal-title":"Nature"},{"key":"BFsrep43161_CR76","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1083\/jcb.127.1.107","volume":"127","author":"LM Machesky","year":"1994","unstructured":"Machesky, L. M., Atkinson, S. J., Ampe, C., Vandekerckhove, J. & Pollard, T. D. Purification of a cortical complex containing two unconventional actins from Acanthamoeba by affinity chromatography on profilin-agarose. J Cell Biol 127, 107\u2013115 (1994).","journal-title":"J Cell Biol"},{"key":"BFsrep43161_CR77","doi-asserted-by":"crossref","first-page":"117","DOI":"10.1016\/S0955-0674(99)80014-3","volume":"11","author":"LM Machesky","year":"1999","unstructured":"Machesky, L. M. & Gould, K. L. The Arp2\/3 complex: a multifunctional actin organizer. Curr Opin Cell Biol 11, 117\u2013121 (1999).","journal-title":"Curr Opin Cell Biol"},{"key":"BFsrep43161_CR78","doi-asserted-by":"crossref","first-page":"196","DOI":"10.1016\/j.tplants.2004.02.004","volume":"9","author":"MK Kandasamy","year":"2004","unstructured":"Kandasamy, M. K., Deal, R. B., McKinney, E. C. & Meagher, R. B. Plant actin-related proteins. Trends Plant Sci 9, 196\u2013202 (2004).","journal-title":"Trends Plant Sci"},{"key":"BFsrep43161_CR79","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1007\/s00709-003-0004-8","volume":"222","author":"K Van Gestel","year":"2003","unstructured":"Van Gestel, K. et al. Immunological evidence for the presence of plant homologues of the actin- related protein Arp3 in tobacco and maize: subcellular localization to actin-enriched pit fields and emerging root hairs. Protoplasma 222, 45\u201352 (2003).","journal-title":"Protoplasma"},{"key":"BFsrep43161_CR80","doi-asserted-by":"crossref","first-page":"603","DOI":"10.1093\/jxb\/ern307","volume":"60","author":"J Maisch","year":"2009","unstructured":"Maisch, J., Fiserova, J., Fischer, L. & Nick, P. Tobacco Arp3 is localized to actin-nucleating sites in vivo. J Exp Bot 60, 603\u2013614 (2009).","journal-title":"J Exp Bot"},{"key":"BFsrep43161_CR81","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1007\/s00709-006-0146-6","volume":"227","author":"J Fiserova","year":"2006","unstructured":"Fiserova, J., Schwarzerova, K., Petrasek, J. & Opatrny, Z. ARP2 and ARP3 are localized to sites of actin filament nucleation in tobacco BY-2 cells. Protoplasma 227, 119\u2013128 (2006).","journal-title":"Protoplasma"},{"key":"BFsrep43161_CR82","doi-asserted-by":"crossref","first-page":"997","DOI":"10.1104\/pp.010906","volume":"128","author":"EC McKinney","year":"2002","unstructured":"McKinney, E. C., Kandasamy, M. K. & Meagher, R. B. Arabidopsis contains ancient classes of differentially expressed actin-related protein genes. Plant Physiol 128, 997\u20131007 (2002).","journal-title":"Plant Physiol"},{"key":"BFsrep43161_CR83","doi-asserted-by":"crossref","first-page":"3137","DOI":"10.1242\/dev.00549","volume":"130","author":"J Mathur","year":"2003","unstructured":"Mathur, J. et al. Arabidopsis CROOKED encodes for the smallest subunit of the ARP2\/3 complex and controls cell shape by region specific fine F-actin formation. Development 130, 3137\u20133146 (2003).","journal-title":"Development"},{"key":"BFsrep43161_CR84","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1016\/j.pbi.2004.11.004","volume":"8","author":"DB Szymanski","year":"2005","unstructured":"Szymanski, D. B. Breaking the WAVE complex: the point of Arabidopsis trichomes. Curr Opin Plant Biol 8, 103\u2013112 (2005).","journal-title":"Curr Opin Plant Biol"},{"key":"BFsrep43161_CR85","doi-asserted-by":"crossref","first-page":"954","DOI":"10.1038\/nrm1765","volume":"6","author":"MJ Deeks","year":"2005","unstructured":"Deeks, M. J. & Hussey, P. J. Arp2\/3 and SCAR: plants move to the fore. Nat Rev Mol Cell Biol 6, 954\u2013964 (2005).","journal-title":"Nat Rev Mol Cell Biol"},{"key":"BFsrep43161_CR86","doi-asserted-by":"crossref","first-page":"377","DOI":"10.1002\/bies.20206","volume":"27","author":"J Mathur","year":"2005","unstructured":"Mathur, J. The ARP2\/3 complex: giving plant cells a leading edge. Bioessays 27, 377\u2013387 (2005).","journal-title":"Bioessays"},{"key":"BFsrep43161_CR87","doi-asserted-by":"crossref","first-page":"2327","DOI":"10.1105\/tpc.105.033266","volume":"17","author":"PA Harries","year":"2005","unstructured":"Harries, P. A., Pan, A. & Quatrano, R. S. Actin-related protein2\/3 complex component ARPC1 is required for proper cell morphogenesis and polarized cell growth in Physcomitrella patens. Plant Cell 17, 2327\u20132339 (2005).","journal-title":"Plant Cell"},{"key":"BFsrep43161_CR88","doi-asserted-by":"crossref","first-page":"502","DOI":"10.1105\/tpc.104.027987","volume":"17","author":"D Basu","year":"2005","unstructured":"Basu, D. et al. DISTORTED3\/SCAR2 is a putative arabidopsis WAVE complex subunit that activates the Arp2\/3 complex and is required for epidermal morphogenesis. Plant Cell 17, 502\u2013524 (2005).","journal-title":"Plant Cell"},{"key":"BFsrep43161_CR89","doi-asserted-by":"crossref","first-page":"1341","DOI":"10.1016\/S0960-9822(03)00493-7","volume":"13","author":"J Le","year":"2003","unstructured":"Le, J., El-Assal Sel, D., Basu, D., Saad, M. E. & Szymanski, D. B. Requirements for Arabidopsis ATARP2 and ATARP3 during epidermal development. Curr Biol 13, 1341\u20131347 (2003).","journal-title":"Curr Biol"},{"key":"BFsrep43161_CR90","doi-asserted-by":"crossref","first-page":"4345","DOI":"10.1242\/dev.01307","volume":"131","author":"D Basu","year":"2004","unstructured":"Basu, D., El-Assal Sel, D., Le, J., Mallery, E. L. & Szymanski, D. B. Interchangeable functions of Arabidopsis PIROGI and the human WAVE complex subunit SRA1 during leaf epidermal development. Development 131, 4345\u20134355 (2004).","journal-title":"Development"},{"key":"BFsrep43161_CR91","doi-asserted-by":"crossref","first-page":"526","DOI":"10.1111\/j.1365-313X.2004.02065.x","volume":"38","author":"S El-Din El-Assal","year":"2004","unstructured":"El-Din El-Assal, S., Le, J., Basu, D., Mallery, E. L. & Szymanski, D. B. DISTORTED2 encodes an ARPC2 subunit of the putative Arabidopsis ARP2\/3 complex. Plant J 38, 526\u2013538 (2004).","journal-title":"Plant J"},{"key":"BFsrep43161_CR92","doi-asserted-by":"crossref","first-page":"2092","DOI":"10.1139\/b75-234","volume":"53","author":"F Constabel","year":"1975","unstructured":"Constabel, F., Dudits, D., Gamborg, O. L. & Kao, K. N. Nuclear fusion in intergeneric heterokaryons. Can J Botany 53, 2092\u20132095 (1975).","journal-title":"Can J Botany"},{"key":"BFsrep43161_CR93","doi-asserted-by":"crossref","first-page":"6","DOI":"10.1007\/BF00340171","volume":"214","author":"Y Hayashi","year":"1988","unstructured":"Hayashi, Y., Kyozuka, J. & Shimamoto, K. Hybrids of rice (Oryza sativa L.) and wild Oryza species obtained by cell fusion. Mol Gen Genet 214, 6\u201310 (1988).","journal-title":"Mol Gen Genet"}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/srep43161","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/srep43161.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/srep43161.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,12,23]],"date-time":"2022-12-23T19:46:56Z","timestamp":1671824816000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/srep43161"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2017,2,22]]},"references-count":93,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2017,3,9]]}},"alternative-id":["BFsrep43161"],"URL":"https:\/\/doi.org\/10.1038\/srep43161","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2017,2,22]]},"assertion":[{"value":"4 October 2016","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"20 January 2017","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"22 February 2017","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing financial interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"43161"}}