start-ver=1.4 cd-journal=joma no-vol=18 cd-vols= no-issue=1 article-no= start-page=2281159 end-page= dt-received= dt-revised= dt-accepted= dt-pub-year=2023 dt-pub=20231115 dt-online= en-article= kn-article= en-subject= kn-subject= en-title= kn-title=Microtubule-associated proteins WDL5 and WDL6 play a critical role in pollen tube growth in Arabidopsis thaliana en-subtitle= kn-subtitle= en-abstract= kn-abstract=Morphological response of cells to environment involves concerted rearrangements of microtubules and actin microfilaments. A mutant of WAVE-DAMPENED2-LIKE5 (WDL5), which encodes an ethylene-regulated microtubule-associated protein belonging to the WVD2/WDL family in Arabidopsis thaliana, shows attenuation in the temporal root growth reduction in response to mechanical stress. We found that a T-DNA knockout of WDL6, the closest homolog of WDL5, oppositely shows an enhancement of the response. To know the functional relationship between WDL5 and WDL6, we attempted to generate the double mutant by crosses but failed in isolation. Close examination of gametophytes in plants that are homozygous for one and heterozygous for the other revealed that these plants produce pollen grains with a reduced rate of germination and tube growth. Reciprocal cross experiments of these plants with the wild type confirmed that the double mutation is not inherited paternally. These results suggest a critical and cooperative function of WDL5 and WDL6 in pollen tube growth. en-copyright= kn-copyright= en-aut-name=OkamotoTakashi en-aut-sei=Okamoto en-aut-mei=Takashi kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=1 ORCID= en-aut-name=MotoseHiroyasu en-aut-sei=Motose en-aut-mei=Hiroyasu kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=2 ORCID= en-aut-name=TakahashiTaku en-aut-sei=Takahashi en-aut-mei=Taku kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=3 ORCID= affil-num=1 en-affil=Department of Biological Science, Graduate School of Environmental, Life, Natural Science and Technology, Okayama University kn-affil= affil-num=2 en-affil=Department of Biological Science, Graduate School of Environmental, Life, Natural Science and Technology, Okayama University kn-affil= affil-num=3 en-affil=Department of Biological Science, Graduate School of Environmental, Life, Natural Science and Technology, Okayama University kn-affil= en-keyword=Arabidopsis kn-keyword=Arabidopsis en-keyword=pollen germination kn-keyword=pollen germination en-keyword=pollen tube growth kn-keyword=pollen tube growth en-keyword=the WVD2/WDL family kn-keyword=the WVD2/WDL family END start-ver=1.4 cd-journal=joma no-vol=596 cd-vols= no-issue=23 article-no= start-page=3005 end-page=3014 dt-received= dt-revised= dt-accepted= dt-pub-year=2022 dt-pub=20220812 dt-online= en-article= kn-article= en-subject= kn-subject= en-title= kn-title=Loss of function of an Arabidopsis homologue of JMJD6 suppresses the dwarf phenotype of acl5, a mutant defective in thermospermine biosynthesis en-subtitle= kn-subtitle= en-abstract= kn-abstract=In Arabidopsis thaliana, the ACL5 gene encodes thermospermine synthase and its mutant, acl5, exhibits a dwarf phenotype with excessive xylem formation. Studies of suppressor mutants of acl5 reveal the involvement of thermospermine in enhancing mRNA translation of the SAC51 gene family. We show here that a mutant, sac59, which partially suppresses the acl5 phenotype, has a point mutation in JMJ22 encoding a D6-class Jumonji C protein (JMJD6). A T-DNA insertion allele, jmj22-2, also partially suppressed the acl5 phenotype while mutants of its closest two homologs JMJ21 and JMJ20 had no such effects, suggesting a unique role for JMJ22 in plant development. We found that mRNAs of the SAC51 family are more stabilized in acl5 jmj22-2 than in acl5. en-copyright= kn-copyright= en-aut-name=MatsuoHirotoshi en-aut-sei=Matsuo en-aut-mei=Hirotoshi kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=1 ORCID= en-aut-name=FukushimaHiroko en-aut-sei=Fukushima en-aut-mei=Hiroko kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=2 ORCID= en-aut-name=KurokawaShinpei en-aut-sei=Kurokawa en-aut-mei=Shinpei kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=3 ORCID= en-aut-name=KawanoEri en-aut-sei=Kawano en-aut-mei=Eri kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=4 ORCID= en-aut-name=OkamotoTakashi en-aut-sei=Okamoto en-aut-mei=Takashi kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=5 ORCID= en-aut-name=MotoseHiroyasu en-aut-sei=Motose en-aut-mei=Hiroyasu kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=6 ORCID= en-aut-name=TakahashiTaku en-aut-sei=Takahashi en-aut-mei=Taku kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=7 ORCID= affil-num=1 en-affil=Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=2 en-affil=Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=3 en-affil=Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=4 en-affil=Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=5 en-affil=Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=6 en-affil=Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=7 en-affil=Graduate School of Natural Science and Technology, Okayama University kn-affil= en-keyword=Arabidopsis kn-keyword=Arabidopsis en-keyword=JMJD6 kn-keyword=JMJD6 en-keyword=mRNA stability kn-keyword=mRNA stability en-keyword=thermospermine kn-keyword=thermospermine en-keyword=xylem development kn-keyword=xylem development END start-ver=1.4 cd-journal=joma no-vol=2020 cd-vols= no-issue=18 article-no= start-page=2745 end-page=2753 dt-received= dt-revised= dt-accepted= dt-pub-year=2020 dt-pub=20200320 dt-online= en-article= kn-article= en-subject= kn-subject= en-title= kn-title=Chemical Synthesis and Biological Effect on Xylem Formation of Xylemin and Its Analogues en-subtitle= kn-subtitle= en-abstract= kn-abstract=Xylemin (6 ) and its designed structural analogues 18 –23 , N ‐(4‐aminobutyl)alkylamines, were synthesized by 2‐nitrobenzenesulfonamide (Ns) strategy. Investigation of the improved synthesis of 20 –23 resulted in the development of one‐step synthesis of these analogues from the commercially available corresponding ketones. Biological assessment of the synthetic molecules elucidated that xylemin (6 ) and the analogue N ‐(4‐aminobutyl)cyclopentylamine (21 ) promoted the expression level of thermospermine synthase ACAULIS5 (ACL5 ) and enhanced xylem formation. In addition, xylemin (6 ) was found to significantly promote lateral root formation, whereas xylemin analogues 18 –23 including 21 did not. These results indicate that the analogue 21 has the potential as a novel inhibitor of thermospermine synthesis to work specifically in xylem differentiation. en-copyright= kn-copyright= en-aut-name=TakamuraHiroyoshi en-aut-sei=Takamura en-aut-mei=Hiroyoshi kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=1 ORCID= en-aut-name=MotoseHiroyasu en-aut-sei=Motose en-aut-mei=Hiroyasu kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=2 ORCID= en-aut-name=OtsuTaichi en-aut-sei=Otsu en-aut-mei=Taichi kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=3 ORCID= en-aut-name=ShinoharaShiori en-aut-sei=Shinohara en-aut-mei=Shiori kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=4 ORCID= en-aut-name=KounoRyugo en-aut-sei=Kouno en-aut-mei=Ryugo kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=5 ORCID= en-aut-name=KadotaIsao en-aut-sei=Kadota en-aut-mei=Isao kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=6 ORCID= en-aut-name=TakahashiTaku en-aut-sei=Takahashi en-aut-mei=Taku kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=7 ORCID= affil-num=1 en-affil=Department of Biological Science, Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=2 en-affil=Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=3 en-affil=Department of Chemistry, Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=4 en-affil=Department of Biological Science, Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=5 en-affil=Department of Biological Science, Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=6 en-affil=Department of Chemistry, Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=7 en-affil=Department of Biological Science, Graduate School of Natural Science and Technology, Okayama University kn-affil= en-keyword=Amines kn-keyword=Amines en-keyword=Biological activity kn-keyword=Biological activity en-keyword=Chemical synthesis kn-keyword=Chemical synthesis en-keyword=Reductive amination kn-keyword=Reductive amination en-keyword=Structure‐activity relationships kn-keyword=Structure‐activity relationships END start-ver=1.4 cd-journal=joma no-vol=57 cd-vols= no-issue=8 article-no= start-page=1583 end-page=1592 dt-received= dt-revised= dt-accepted= dt-pub-year=2016 dt-pub=201608 dt-online= en-article= kn-article= en-subject= kn-subject= en-title= kn-title=The SAC51 Family Plays a Central Role in Thermospermine Responses in Arabidopsis en-subtitle= kn-subtitle= en-abstract= kn-abstract= The acaulis5 (acl5) mutant of Arabidopsis thaliana is defective in the biosynthesis of thermospermine and shows a dwarf phenotype associated with excess xylem differentiation. SAC51 was identified from a dominant suppressor of acl5, sac51-d, and encodes a basic helix-loop-helix protein. The sac51-d mutant has a premature termination codon in an upstream open reading frame (uORF) that is conserved among all four members of the SAC51 family, SAC51 and SACL1-SACL3 This suggests that thermospermine cancels the inhibitory effect of the uORF in main ORF translation. Another suppressor, sac57-d, has a mutation in the conserved uORF of SACL3 To define further the function of the SAC51 family in the thermospermine response, we analyzed T-DNA insertion mutants of each gene. Although sacl1-1 may not be a null allele, the quadruple mutant showed a semi-dwarf phenotype but with an increased level of thermospermine and decreased sensitivity to exogenous thermospermine that normally represses xylem differentiation. The sac51-1 sacl3-1 double mutant was also insensitive to thermospermine. These results suggest that SAC51 and SACL3 play a key role in thermospermine-dependent negative control of thermospermine biosynthesis and xylem differentiation. Using 5' leader-GUS (β-glucuronidase) fusion constructs, however, we detected a significant enhancement of the GUS activity by thermospermine only in SAC51 and SACL1 constructs. Furthermore, while acl5-1 sac51-1 showed the acl5 dwarf phenotype, acl5-1 sacl3-1 exhibited an extremely tiny-plant phenotype. These results suggest a complex regulatory network for the thermospermine response in which SAC51 and SACL3 function in parallel pathways. en-copyright= kn-copyright= en-aut-name=CaiQingqing en-aut-sei=Cai en-aut-mei=Qingqing kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=1 ORCID= en-aut-name=FukushimaHiroko en-aut-sei=Fukushima en-aut-mei=Hiroko kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=2 ORCID= en-aut-name=YamamotoMai en-aut-sei=Yamamoto en-aut-mei=Mai kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=3 ORCID= en-aut-name=IshiiNami en-aut-sei=Ishii en-aut-mei=Nami kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=4 ORCID= en-aut-name=SakamotoTomoaki en-aut-sei=Sakamoto en-aut-mei=Tomoaki kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=5 ORCID= en-aut-name=KurataTetsuya en-aut-sei=Kurata en-aut-mei=Tetsuya kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=6 ORCID= en-aut-name=MotoseHiroyasu en-aut-sei=Motose en-aut-mei=Hiroyasu kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=7 ORCID= en-aut-name=TakahashiTaku en-aut-sei=Takahashi en-aut-mei=Taku kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=8 ORCID= affil-num=1 en-affil=Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=2 en-affil=Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=3 en-affil=Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=4 en-affil=Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=5 en-affil=Graduate School of Biological Sciences, Nara Institute of Science and Technology kn-affil= affil-num=6 en-affil=Graduate School of Biological Sciences, Nara Institute of Science and Technology kn-affil= affil-num=7 en-affil=Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=8 en-affil=Graduate School of Natural Science and Technology, Okayama University kn-affil= en-keyword=Arabidopsis kn-keyword=Arabidopsis en-keyword=Thermospermine kn-keyword=Thermospermine en-keyword=Translation kn-keyword=Translation en-keyword=Xylem differentiation kn-keyword=Xylem differentiation en-keyword=uORF kn-keyword=uORF END start-ver=1.4 cd-journal=joma no-vol=7 cd-vols= no-issue= article-no= start-page=7826 end-page= dt-received= dt-revised= dt-accepted= dt-pub-year=2017 dt-pub=20170810 dt-online= en-article= kn-article= en-subject= kn-subject= en-title= kn-title=Directional cell expansion requires NIMA-related kinase 6 (NEK6)-mediated cortical microtubule destabilization; en-subtitle= kn-subtitle= en-abstract= kn-abstract=　Plant cortical microtubules align perpendicular to the growth axis to determine the direction of cell growth. However, it remains unclear how plant cells form well-organized cortical microtubule arrays in the absence of a centrosome. In this study, we investigated the functions of Arabidopsis NIMA-related kinase 6 (NEK6), which regulates microtubule organization during anisotropic cell expansion. Quantitative analysis of hypocotyl cell growth in the nek6-1 mutant demonstrated that NEK6 suppresses ectopic outgrowth and promotes cell elongation in different regions of the hypocotyl. Loss of NEK6 function led to excessive microtubule waving and distortion, implying that NEK6 suppresses the aberrant cortical microtubules. Live cell imaging showed that NEK6 localizes to the microtubule lattice and to the shrinking plus and minus ends of microtubules. In agreement with this observation, the induced overexpression of NEK6 reduced and disorganized cortical microtubules and suppressed cell elongation. Furthermore, we identified five phosphorylation sites in β-tubulin that serve as substrates for NEK6 in vitro. Alanine substitution of the phosphorylation site Thr166 promoted incorporation of mutant β-tubulin into microtubules. Taken together, these results suggest that NEK6 promotes directional cell growth through phosphorylation of β-tubulin and the resulting destabilization of cortical microtubules. en-copyright= kn-copyright= en-aut-name=TakataniShogo en-aut-sei=Takatani en-aut-mei=Shogo kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=1 ORCID= en-aut-name=OzawaShinichiro en-aut-sei=Ozawa en-aut-mei=Shinichiro kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=2 ORCID= en-aut-name=YagiNoriyoshi en-aut-sei=Yagi en-aut-mei=Noriyoshi kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=3 ORCID= en-aut-name=HottaTakashi en-aut-sei=Hotta en-aut-mei=Takashi kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=4 ORCID= en-aut-name=HashimotoTakashi en-aut-sei=Hashimoto en-aut-mei=Takashi kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=5 ORCID= en-aut-name=TakahashiYuichiro en-aut-sei=Takahashi en-aut-mei=Yuichiro kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=6 ORCID= en-aut-name=TakahashiTaku en-aut-sei=Takahashi en-aut-mei=Taku kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=7 ORCID= en-aut-name=MotoseHiroyasu en-aut-sei=Motose en-aut-mei=Hiroyasu kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=8 ORCID= affil-num=1 en-affil=Department of Biological Science, Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=2 en-affil=Department of Biological Science, Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=3 en-affil=Graduate School of Biological Science, Nara Institute of Science and Technology kn-affil= affil-num=4 en-affil=Graduate School of Biological Science, Nara Institute of Science and Technology kn-affil= affil-num=5 en-affil=Graduate School of Biological Science, Nara Institute of Science and Technology kn-affil= affil-num=6 en-affil=Graduate School of Natural Science and Technology/Faculty of Science, Okayama University kn-affil= affil-num=7 en-affil=Department of Biological Science, Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=8 en-affil=Department of Biological Science, Graduate School of Natural Science and Technology, Okayama University kn-affil= en-keyword=Cell growth kn-keyword=Cell growth en-keyword=Microtubules kn-keyword=Microtubules en-keyword=Plant cytoskeleton kn-keyword=Plant cytoskeleton END start-ver=1.4 cd-journal=joma no-vol=6 cd-vols= no-issue= article-no= start-page=21487 end-page= dt-received= dt-revised= dt-accepted= dt-pub-year=2016 dt-pub=20160216 dt-online= en-article= kn-article= en-subject= kn-subject= en-title= kn-title=Chemical control of xylem differentiation by thermospermine, xylemin and auxin en-subtitle= kn-subtitle= en-abstract= kn-abstract= The xylem conducts water and minerals from the root to the shoot and provides mechanical strength to the plant body. The vascular precursor cells of the procambium differentiate to form continuous vascular strands, from which xylem and phloem cells are generated in the proper spatiotemporal pattern. Procambium formation and xylem differentiation are directed by auxin. In angiosperms, thermospermine, a structural isomer of spermine, suppresses xylem differentiation by limiting auxin signalling. However, the process of auxin-inducible xylem differentiation has not been fully elucidated and remains difficult to manipulate. Here, we found that an antagonist of spermidine can act as an inhibitor of thermospermine biosynthesis and results in excessive xylem differentiation, which is a phenocopy of a thermospermine-deficient mutant acaulis5 in Arabidopsis thaliana. We named this compound xylemin owing to its xylem-inducing effect. Application of a combination of xylemin and thermospermine to wild-type seedlings negates the effect of xylemin, whereas co-treatment with xylemin and a synthetic proauxin, which undergoes hydrolysis to release active auxin, has a synergistic inductive effect on xylem differentiation. Thus, xylemin may serve as a useful transformative chemical tool not only for the study of thermospermine function in various plant species but also for the control of xylem induction and woody biomass production. en-copyright= kn-copyright= en-aut-name=YoshimotoKaori en-aut-sei=Yoshimoto en-aut-mei=Kaori kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=1 ORCID= en-aut-name=TakamuraHiroyoshi en-aut-sei=Takamura en-aut-mei=Hiroyoshi kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=2 ORCID= en-aut-name=KadotaIsao en-aut-sei=Kadota en-aut-mei=Isao kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=3 ORCID= en-aut-name=MotoseHiroyasu en-aut-sei=Motose en-aut-mei=Hiroyasu kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=4 ORCID= en-aut-name=TakahashiTaku en-aut-sei=Takahashi en-aut-mei=Taku kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=5 ORCID= affil-num=1 en-affil=Department of Biological Science, Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=2 en-affil=Department of Chemistry, Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=3 en-affil=Department of Chemistry, Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=4 en-affil=Department of Biological Science, Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=5 en-affil=Department of Biological Science, Graduate School of Natural Science and Technology, Okayama University kn-affil= END start-ver=1.4 cd-journal=joma no-vol=6 cd-vols= no-issue= article-no= start-page=21487 end-page= dt-received= dt-revised= dt-accepted= dt-pub-year=2016 dt-pub=20160216 dt-online= en-article= kn-article= en-subject= kn-subject= en-title= kn-title=Chemical control of xylem differentiation by thermospermine, xylemin, and auxin en-subtitle= kn-subtitle= en-abstract= kn-abstract=The xylem conducts water and minerals from the root to the shoot and provides mechanical strength to the plant body. The vascular precursor cells of the procambium differentiate to form continuous vascular strands, from which xylem and phloem cells are generated in the proper spatiotemporal pattern. Procambium formation and xylem differentiation are directed by auxin. In angiosperms, thermospermine, a structural isomer of spermine, suppresses xylem differentiation by limiting auxin signalling. However, the process of auxin-inducible xylem differentiation has not been fully elucidated and remains difficult to manipulate. Here, we found that an antagonist of spermidine can act as an inhibitor of thermospermine biosynthesis and results in excessive xylem differentiation, which is a phenocopy of a thermospermine-deficient mutant acaulis5 in Arabidopsis thaliana. We named this compound xylemin owing to its xylem-inducing effect. Application of a combination of xylemin and thermospermine to wild-type seedlings negates the effect of xylemin, whereas co-treatment with xylemin and a synthetic proauxin, which undergoes hydrolysis to release active auxin, has a synergistic inductive effect on xylem differentiation. Thus, xylemin may serve as a useful transformative chemical tool not only for the study of thermospermine function in various plant species but also for the control of xylem induction and woody biomass production. en-copyright= kn-copyright= en-aut-name=YoshimotoKaori en-aut-sei=Yoshimoto en-aut-mei=Kaori kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=1 ORCID= en-aut-name=TakamuraHiroyoshi en-aut-sei=Takamura en-aut-mei=Hiroyoshi kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=2 ORCID= en-aut-name=KadotaIsao en-aut-sei=Kadota en-aut-mei=Isao kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=3 ORCID= en-aut-name=MotoseHiroyasu en-aut-sei=Motose en-aut-mei=Hiroyasu kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=4 ORCID= en-aut-name=TakahashiTaku en-aut-sei=Takahashi en-aut-mei=Taku kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=5 ORCID= affil-num=1 en-affil= kn-affil=Department of Biological Science, Graduate School of Natural Science and Technology, Okayama University affil-num=2 en-affil= kn-affil=Department of Chemistry, Graduate School of Natural Science and Technology, Okayama University affil-num=3 en-affil= kn-affil=Department of Chemistry, Graduate School of Natural Science and Technology, Okayama University affil-num=4 en-affil=Department of Biological Science, Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=5 en-affil= kn-affil=Department of Biological Science, Graduate School of Natural Science and Technology, Okayama University END start-ver=1.4 cd-journal=joma no-vol=128 cd-vols= no-issue=6 article-no= start-page=875 end-page=891 dt-received= dt-revised= dt-accepted= dt-pub-year=2015 dt-pub=201511 dt-online= en-article= kn-article= en-subject= kn-subject= en-title= kn-title=Structure, function, and evolution of plant NIMA-related kinases: implication for phosphorylation-dependent microtubule regulation en-subtitle= kn-subtitle= en-abstract= kn-abstract=icrotubules are highly dynamic structures that control the spatiotemporal pattern of cell growth and division. Microtubule dynamics are regulated by reversible protein phosphorylation involving both protein kinases and phosphatases. Never in mitosis A (NIMA)-related kinases (NEKs) are a family of serine/threonine kinases that regulate microtubule-related mitotic events in fungi and animal cells (e.g. centrosome separation and spindle formation). Although plants contain multiple members of the NEK family, their functions remain elusive. Recent studies revealed that NEK6 of Arabidopsis thaliana regulates cell expansion and morphogenesis through β-tubulin phosphorylation and microtubule destabilization. In addition, plant NEK members participate in organ development and stress responses. The present phylogenetic analysis indicates that plant NEK genes are diverged from a single NEK6-like gene, which may share a common ancestor with other kinases involved in the control of microtubule organization. On the contrary, another mitotic kinase, polo-like kinase, might have been lost during the evolution of land plants. We propose that plant NEK members have acquired novel functions to regulate cell growth, microtubule organization, and stress responses. en-copyright= kn-copyright= en-aut-name=TakataniShogo en-aut-sei=Takatani en-aut-mei=Shogo kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=1 ORCID= en-aut-name=OtaniKento en-aut-sei=Otani en-aut-mei=Kento kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=2 ORCID= en-aut-name=KanazawaMai en-aut-sei=Kanazawa en-aut-mei=Mai kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=3 ORCID= en-aut-name=TakahashiTaku en-aut-sei=Takahashi en-aut-mei=Taku kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=4 ORCID= en-aut-name=MotoseHiroyasu en-aut-sei=Motose en-aut-mei=Hiroyasu kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=5 ORCID= affil-num=1 en-affil= kn-affil=Division of Bioscience, Graduate School of Natural Science and Technology, Okayama University affil-num=2 en-affil= kn-affil=Division of Bioscience, Graduate School of Natural Science and Technology, Okayama University affil-num=3 en-affil= kn-affil=Department of Biology, Faculty of Science, Okayama University affil-num=4 en-affil=Division of Bioscience, Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=5 en-affil=Department of Biology, Faculty of Science, Okayama University kn-affil= en-keyword=NIMA-related kinase kn-keyword=NIMA-related kinase en-keyword=Phosphorylation kn-keyword=Phosphorylation en-keyword=Tubulin kn-keyword=Tubulin en-keyword=Microtubule kn-keyword=Microtubule en-keyword=Cell expansion kn-keyword=Cell expansion en-keyword=Cell division kn-keyword=Cell division END start-ver=1.4 cd-journal=joma no-vol=5 cd-vols= no-issue= article-no= start-page=11364 end-page= dt-received= dt-revised= dt-accepted= dt-pub-year=2015 dt-pub=20150612 dt-online= en-article= kn-article= en-subject= kn-subject= en-title= kn-title=Abscisic acid induces ectopic outgrowth in epidermal cells through cortical microtubule reorganization in Arabidopsis thaliana en-subtitle= kn-subtitle= en-abstract= kn-abstract=Abscisic acid (ABA) regulates seed maturation, germination and various stress responses in plants. The roles of ABA in cellular growth and morphogenesis, however, remain to be explored. Here, we report that ABA induces the ectopic outgrowth of epidermal cells in Arabidopsis thaliana. Seedlings of A. thaliana germinated and grown in the presence of ABA developed ectopic protrusions in the epidermal cells of hypocotyls, petioles and cotyledons. One protrusion was formed in the middle of each epidermal cell. In the hypocotyl epidermis, two types of cell files are arranged alternately into non-stoma cell files and stoma cell files, ectopic protrusions being restricted to the non-stoma cell files. This suggests the presence of a difference in the degree of sensitivity to ABA or in the capacity of cells to form protrusions between the two cell files. The ectopic outgrowth was suppressed in ABA insensitive mutants, whereas it was enhanced in ABA hypersensitive mutants. Interestingly, ABA-induced ectopic outgrowth was also suppressed in mutants in which microtubule organization was compromised. Furthermore, cortical microtubules were disorganized and depolymerized by the ABA treatment. These results suggest that ABA signaling induces ectopic outgrowth in epidermal cells through microtubule reorganization. en-copyright= kn-copyright= en-aut-name=TakataniShogo en-aut-sei=Takatani en-aut-mei=Shogo kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=1 ORCID= en-aut-name=HirayamaTakashi en-aut-sei=Hirayama en-aut-mei=Takashi kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=2 ORCID= en-aut-name=HashimotoTakashi en-aut-sei=Hashimoto en-aut-mei=Takashi kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=3 ORCID= en-aut-name=TakahashiTaku en-aut-sei=Takahashi en-aut-mei=Taku kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=4 ORCID= en-aut-name=MotoseHiroyasu en-aut-sei=Motose en-aut-mei=Hiroyasu kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=5 ORCID= affil-num=1 en-affil= kn-affil=Department of Biological Science, Graduate School of Natural Science and Technology, Okayama University affil-num=2 en-affil= kn-affil=Institute of Plant Science and Resources, Okayama University affil-num=3 en-affil= kn-affil=Graduate School of Biological Science, Nara Institute of Science and Technology affil-num=4 en-affil= kn-affil=Department of Biological Science, Graduate School of Natural Science and Technology, Okayama University affil-num=5 en-affil= kn-affil=自然科学研究科 en-keyword=Cell growth kn-keyword=Cell growth en-keyword=Plant cytoskeleton kn-keyword=Plant cytoskeleton END start-ver=1.4 cd-journal=joma no-vol=7 cd-vols= no-issue=12 article-no= start-page=1552 end-page=1555 dt-received= dt-revised= dt-accepted= dt-pub-year=2012 dt-pub=201212 dt-online= en-article= kn-article= en-subject= kn-subject= en-title= kn-title=NIMA-related kinases regulate directional cell growth and organ development through microtubule function in Arabidopsis thaliana. en-subtitle= kn-subtitle= en-abstract= kn-abstract=NIMA-related kinase 6 (NEK6) regulates cellular expansion and morphogenesis through microtubule organizaiton in Arabidopsis thaliana. Loss-of-function mutations in NEK6 (nek6/ibo1) cause ectopic outgrowth and microtubule disorganization in epidermal cells. We recently found that NEK6 forms homodimers and heterodimers with NEK4 and NEK5 to destabilize cortical microtubules possibly by direct binding to microtubules and the β-tubulin phosphorylation. Here, we identified a new allele of NEK6 and further analyzed the morphological phenotypes of nek6/ibo1 mutants, along with alleles of nek4 and nek5 mutants. Phenotypic analysis demonstrated that NEK6 is required for the directional growth of roots and hypocotyls, petiole elongation, cell file formation, and trichome morphogenesis. In addition, nek4, nek5, and nek6/ibo1 mutants were hypersensitive to microtubule inhibitors such as propyzamide and taxol. These results suggest that plant NEKs function in directional cell growth and organ development through the regulation of microtubule organization. en-copyright= kn-copyright= en-aut-name=MotoseHiroyasu en-aut-sei=Motose en-aut-mei=Hiroyasu kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=1 ORCID= en-aut-name=TakataniShogo en-aut-sei=Takatani en-aut-mei=Shogo kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=2 ORCID= en-aut-name=IkedaTatsuya en-aut-sei=Ikeda en-aut-mei=Tatsuya kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=3 ORCID= en-aut-name=TakahashiTaku en-aut-sei=Takahashi en-aut-mei=Taku kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=4 ORCID= affil-num=1 en-affil= kn-affil=Division of Biological Sciences; Graduate School of Natural Science and Technology; Okayama University affil-num=2 en-affil= kn-affil=Division of Biological Sciences; Graduate School of Natural Science and Technology; Okayama University affil-num=3 en-affil= kn-affil=Division of Biological Sciences; Graduate School of Natural Science and Technology; Okayama University affil-num=4 en-affil= kn-affil=Division of Biological Sciences; Graduate School of Natural Science and Technology; Okayama University en-keyword=Arabidopsis kn-keyword=Arabidopsis en-keyword=NIMA-related kinase kn-keyword=NIMA-related kinase en-keyword=directional growth kn-keyword=directional growth en-keyword=epidermis kn-keyword=epidermis en-keyword=microtubule kn-keyword=microtubule en-keyword=tubulin kn-keyword=tubulin END start-ver=1.4 cd-journal=joma no-vol=5 cd-vols= no-issue=11 article-no= start-page=2854 end-page=2859 dt-received= dt-revised= dt-accepted= dt-pub-year=2013 dt-pub=201305 dt-online= en-article= kn-article= en-subject= kn-subject= en-title= kn-title=Determination of polyamines in Arabidopsis thaliana by capillary electrophoresis using salicylaldehyde-5-sulfonate as a derivatizing reagent en-subtitle= kn-subtitle= en-abstract= kn-abstract=Herein, we report a novel method for the determination of polyamines in a sample extracted from Arabidopsis thaliana by capillary electrophoresis (CE) using salicylaldehyde-5-sulfonate (SAS) as a derivatizing reagent. An aldehyde group of SAS forms a Schiff base with amino groups of aliphatic polyamines, resulting in an anionic species with an absorption band in the ultraviolet region. The derivatization method was straightforward since the derivatives were formed by mixing a sample with the derivatizing reagent at a neutral pH. In addition, the negative charges induced by SAS led to a high resolution with a short analysis time. This method permitted the separation of five polyamines, which play important roles in plants. However, further improvement in sensitivity was needed for the determination of the polyamines in plant samples. Therefore, the CE method was coupled with solid-phase extraction (SPE) using an ion-pairing formation with sodium dodecyl benzene sulfonate. The SPE method improved the concentration limits of detection to sub-μM levels, which corresponded with a 10-fold enhancement. The calibration curves for cadaverine, putrescine, and spermidine were linear with concentrations that ranged from 1 to 20 μM and correlation coefficients (R2) were greater than 0.998. The proposed method was applied to the determination of spermidine in a plant sample, Arabidopsis thaliana. en-copyright= kn-copyright= en-aut-name=InoueGenki en-aut-sei=Inoue en-aut-mei=Genki kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=1 ORCID= en-aut-name=KanetaTakashi en-aut-sei=Kaneta en-aut-mei=Takashi kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=2 ORCID= en-aut-name=TakayanagiToshio en-aut-sei=Takayanagi en-aut-mei=Toshio kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=3 ORCID= en-aut-name=KakehiJunichi en-aut-sei=Kakehi en-aut-mei=Junichi kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=4 ORCID= en-aut-name=MotoseHiroyasu en-aut-sei=Motose en-aut-mei=Hiroyasu kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=5 ORCID= en-aut-name=TakahashiTaku en-aut-sei=Takahashi en-aut-mei=Taku kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=6 ORCID= affil-num=1 en-affil= kn-affil=Department of Chemistry, Division of Earth, Life, and Molecular Sciences, Graduate School of Natural Science and Technology, Okayama University affil-num=2 en-affil= kn-affil=Department of Chemistry, Division of Earth, Life, and Molecular Sciences, Graduate School of Natural Science and Technology, Okayama University affil-num=3 en-affil= kn-affil=Department of Life System, Institute of Technology and Science, The University of Tokushima affil-num=4 en-affil= kn-affil=Department of Biological Science, Division of Earth, Life, and Molecular Sciences, Graduate School of Natural Science and Technology, Okayama University affil-num=5 en-affil= kn-affil=Department of Biological Science, Division of Earth, Life, and Molecular Sciences, Graduate School of Natural Science and Technology, Okayama University affil-num=6 en-affil= kn-affil=Department of Biological Science, Division of Earth, Life, and Molecular Sciences, Graduate School of Natural Science and Technology, Okayama University END start-ver=1.4 cd-journal=joma no-vol=53 cd-vols= no-issue=4 article-no= start-page=635 end-page=645 dt-received= dt-revised= dt-accepted= dt-pub-year=2012 dt-pub=201204 dt-online= en-article= kn-article= en-subject= kn-subject= en-title= kn-title=A chemical biology approach reveals an opposite action between thermospermine and auxin in xylem development in Arabidopsis thaliana en-subtitle= kn-subtitle= en-abstract= kn-abstract=Thermospermine, a structural isomer of spermine, is produced through the action of ACAULIS5 (ACL5) and suppresses xylem differentiation in Arabidopsis thaliana. To elucidate the molecular basis of the function of thermospermine, we screened chemical libraries for compounds that can modulate xylem differentiation in the acl5 mutant, which is deficient in thermospermine and shows a severe dwarf phenotype associated with excessive proliferation of xylem vessels. We found that the isooctyl ester of a synthetic auxin, 2,4-D, remarkably enhanced xylem vessel differentiation in acl5 seedlings. 2,4-D, 2,4-D analogs and IAA analogs, including 4-chloro IAA (4-Cl-IAA) and IAA ethyl ester, also enhanced xylem vessel formation, while IAA alone had little or no obvious effect on xylem differentiation. These effects of auxin analogs were observed only in the acl5 mutant but not in the wild type, and were suppressed by the anti-auxin, p-chlorophenoxyisobutyric acid (PCIB) and alpha-(phenyl ethyl-2-one)-IAA (PEO-IAA), and also by thermospermine. Furthermore, the suppressor of acaulis51-d (sac51-d) mutation, which causes SAC51 overexpression in the absence of thermospermine and suppresses the dwarf phenotype of acl5, also suppressed the effect of auxin analogs in acl5. These results suggest that the auxin signaling that promotes xylem differentiation is normally limited by SAC51-mediated thermospermine signaling but can be continually stimulated by exogenous auxin analogs in the absence of thermospermine. The opposite action between thermospermine and auxin may fine-tune the timing and spatial pattern of xylem differentiation. en-copyright= kn-copyright= en-aut-name=YoshimotoKaori en-aut-sei=Yoshimoto en-aut-mei=Kaori kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=1 ORCID= en-aut-name=NoutoshiYoshiteru en-aut-sei=Noutoshi en-aut-mei=Yoshiteru kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=2 ORCID= en-aut-name=HayashiKen-ichiro en-aut-sei=Hayashi en-aut-mei=Ken-ichiro kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=3 ORCID= en-aut-name=ShirasuKen en-aut-sei=Shirasu en-aut-mei=Ken kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=4 ORCID= en-aut-name=TakahashiTaku en-aut-sei=Takahashi en-aut-mei=Taku kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=5 ORCID= en-aut-name=MotoseHiroyasu en-aut-sei=Motose en-aut-mei=Hiroyasu kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=6 ORCID= affil-num=1 en-affil=Division of Bioscience, Graduate School of Natural Science and Technology, Okayama University kn-affil= affil-num=2 en-affil= kn-affil= affil-num=3 en-affil= kn-affil=Okayama Univ Sci affil-num=4 en-affil= kn-affil=RIKEN affil-num=5 en-affil= kn-affil=Okayama Univ affil-num=6 en-affil= kn-affil=Okayama Univ en-keyword=ACAULIS5 kn-keyword=ACAULIS5 en-keyword=Arabidopsis thaliana kn-keyword=Arabidopsis thaliana en-keyword=Auxin kn-keyword=Auxin en-keyword=2.4-D kn-keyword=2.4-D en-keyword=Thermospermine kn-keyword=Thermospermine en-keyword=Xylem differentiation kn-keyword=Xylem differentiation END start-ver=1.4 cd-journal=joma no-vol=7 cd-vols= no-issue=8 article-no= start-page=937 end-page=939 dt-received= dt-revised= dt-accepted= dt-pub-year=2012 dt-pub=201208 dt-online= en-article= kn-article= en-subject= kn-subject= en-title= kn-title=Thermospermine suppresses auxin-inducible xylem differentiation in Arabidopsis thaliana en-subtitle= kn-subtitle= en-abstract= kn-abstract=Thermospermine, a structural isomer of spermine, is synthesized by a thermospermine synthase designated ACAULIS5 (ACL5). Thermospermine-deficient acl5 mutant of Arabidopsis thaliana shows severe dwarfism and excessive xylem differentiation. By screening for compounds that affect xylem differentiation in the acl5 mutant, we identified auxin analogs that remarkably enhanced xylem vessel differentiation in the acl5 mutant but not in the wild type. The xylem-inducing effect of auxin analogs was clearly suppressed by thermospermine, indicating that auxin-inducible xylem differentiation is normally limited by thermospermine. Here, we further characterized xylem-inducing effect of auxin analogs in various organs. Auxin analogs promoted protoxylem differentiation in roots and cotyledons in the acl5 mutant. Our results indicate that the opposite action between thermospermine and auxin in xylem differentiation is common in different organs and also suggest that thermospermine might be required for the suppression of protoxylem differentiation. en-copyright= kn-copyright= en-aut-name=YoshimotoKaori en-aut-sei=Yoshimoto en-aut-mei=Kaori kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=1 ORCID= en-aut-name=NoutoshiYoshiteru en-aut-sei=Noutoshi en-aut-mei=Yoshiteru kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=2 ORCID= en-aut-name=HayashiKen-ichiro en-aut-sei=Hayashi en-aut-mei=Ken-ichiro kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=3 ORCID= en-aut-name=ShirasuKen en-aut-sei=Shirasu en-aut-mei=Ken kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=4 ORCID= en-aut-name=TakahashiTaku en-aut-sei=Takahashi en-aut-mei=Taku kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=5 ORCID= en-aut-name=MotoseHiroyasu en-aut-sei=Motose en-aut-mei=Hiroyasu kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=6 ORCID= affil-num=1 en-affil= kn-affil=Division of Bioscience; Graduate School of Natural Science and Technology; Okayama University affil-num=2 en-affil= kn-affil=Research Core for Interdisciplinary Sciences; Okayama University affil-num=3 en-affil= kn-affil=Department of Biochemistry; Okayama University of Science affil-num=4 en-affil= kn-affil=Plant Science Center; RIKEN affil-num=5 en-affil= kn-affil=Division of Bioscience; Graduate School of Natural Science and Technology; Okayama University affil-num=6 en-affil= kn-affil=Division of Bioscience; Graduate School of Natural Science and Technology; Okayama University en-keyword=2,4-D kn-keyword=2,4-D en-keyword=ACAULIS5 kn-keyword=ACAULIS5 en-keyword=auxin kn-keyword=auxin en-keyword=chemical biology kn-keyword=chemical biology en-keyword=thermospermine kn-keyword=thermospermine en-keyword=xylem differentiation kn-keyword=xylem differentiation END start-ver=1.4 cd-journal=joma no-vol=67 cd-vols= no-issue=6 article-no= start-page=993 end-page=1005 dt-received= dt-revised= dt-accepted= dt-pub-year=2011 dt-pub=201109 dt-online= en-article= kn-article= en-subject= kn-subject= en-title= kn-title=NIMA-related kinases 6, 4, and 5 interact with each other to regulate microtubule organization during epidermal cell expansion in Arabidopsis thaliana en-subtitle= kn-subtitle= en-abstract= kn-abstract=NimA-related kinase 6 (NEK6) has been implicated in microtubule regulation to suppress the ectopic outgrowth of epidermal cells; however, its molecular functions remain to be elucidated. Here, we analyze the function of NEK6 and other members of the NEK family with regard to epidermal cell expansion and cortical microtubule organization. The functional NEK6-green fluorescent protein fusion localizes to cortical microtubules, predominantly in particles that exhibit dynamic movement along microtubules. The kinase-dead mutant of NEK6 (ibo1-1) exhibits a disturbance of the cortical microtubule array at the site of ectopic protrusions in epidermal cells. Pharmacological studies with microtubule inhibitors and quantitative analysis of microtubule dynamics indicate excessive stabilization of cortical microtubules in ibo1/nek6 mutants. In addition, NEK6 directly binds to microtubules in vitro and phosphorylates beta-tubulin. NEK6 interacts and co-localizes with NEK4 and NEK5 in a transient expression assay. The ibo1-3 mutation markedly reduces the interaction between NEK6 and NEK4 and increases the interaction between NEK6 and NEK5. NEK4 and NEK5 are required for the ibo1/nek6 ectopic outgrowth phenotype in epidermal cells. These results demonstrate that NEK6 homodimerizes and forms heterodimers with NEK4 and NEK5 to regulate cortical microtubule organization possibly through the phosphorylation of beta-tubulins. en-copyright= kn-copyright= en-aut-name=MotoseHiroyasu en-aut-sei=Motose en-aut-mei=Hiroyasu kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=1 ORCID= en-aut-name=HamadaTakahiro en-aut-sei=Hamada en-aut-mei=Takahiro kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=2 ORCID= en-aut-name=YoshimotoKaori en-aut-sei=Yoshimoto en-aut-mei=Kaori kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=3 ORCID= en-aut-name=MurataTakashi en-aut-sei=Murata en-aut-mei=Takashi kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=4 ORCID= en-aut-name=HasebeMitsuyasu en-aut-sei=Hasebe en-aut-mei=Mitsuyasu kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=5 ORCID= en-aut-name=WatanabeYuichiro en-aut-sei=Watanabe en-aut-mei=Yuichiro kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=6 ORCID= en-aut-name=HashimotoTakashi en-aut-sei=Hashimoto en-aut-mei=Takashi kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=7 ORCID= en-aut-name=SakaiTatsuya en-aut-sei=Sakai en-aut-mei=Tatsuya kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=8 ORCID= en-aut-name=TakahashiTaku en-aut-sei=Takahashi en-aut-mei=Taku kn-aut-name= kn-aut-sei= kn-aut-mei= aut-affil-num=9 ORCID= affil-num=1 en-affil= kn-affil=Okayama Univ affil-num=2 en-affil= kn-affil=Univ Massachusetts affil-num=3 en-affil= kn-affil=Okayama Univ affil-num=4 en-affil= kn-affil=Natl Inst Basic Biol affil-num=5 en-affil= kn-affil=Natl Inst Basic Biol affil-num=6 en-affil= kn-affil=Univ Tokyo affil-num=7 en-affil= kn-affil=Nara Inst Sci & Technol affil-num=8 en-affil= kn-affil=Niigata Univ affil-num=9 en-affil= kn-affil=Okayama Univ en-keyword=NimA-related kinase kn-keyword=NimA-related kinase en-keyword=cell expansion kn-keyword=cell expansion en-keyword=microtubule kn-keyword=microtubule en-keyword=epidermal cell kn-keyword=epidermal cell en-keyword=Arabidopsis thaliana kn-keyword=Arabidopsis thaliana END