植物驱动蛋白: 从微管阵列到生理活动调控

doi:10.11983/CBB22007

摘要/Abstract

摘要： 驱动蛋白(kinesin)是以微管为轨道的分子马达, 其催化ATP水解为ADP, 将贮藏在ATP分子中的化学能高效地转化为机械能, 在细胞形态建成、细胞分裂、细胞运动、胞内物质运输和信号转导等多种生命活动中发挥重要作用。对植物驱动蛋白的研究落后于动物和真菌, 其原因不仅由于植物进化出独有的驱动蛋白家族, 而且其家族成员数量远多于动物驱动蛋白。该文主要总结了驱动蛋白在微管阵列动态组织, 包括周质微管和有丝分裂早前期微管带、纺锤体及成膜体中的角色和功能, 以及其对植物生理活动的调控作用。同时对重要经济作物大豆(Glycine max)中的驱动蛋白进行了系统分析、分类及功能预测, 发现大豆驱动蛋白数量庞大。结合公共数据库中大豆转录组数据, 对部分大豆驱动蛋白进行功能预测, 以期对大豆及其它作物驱动蛋白功能研究提供线索和启示。

关键词: 驱动蛋白, 微管阵列, 生理调控, 大豆

Abstract: Kinesins are molecular motors that move along microtubules tracks, catalyze the hydrolysis of ATP to ADP, convert the chemical energy stored in ATP molecules into a mechanical force efficiently, and play important roles in various life activities such as cell morphogenesis, cell division, cell movement, intracellular transport, and signal transduction. Plant kinesin research lag behind that of animals and fungi, not only because plants have evolved a unique kinesin family, but also have more members than that of animals. Here we summarize the most recent advancements made towards understanding kinesin functions in the dynamic organization of microtubule arrays, including cortical microtubules and mitotic pre-prophase band, spindle apparatus and phragmoplast, and the regulation of plant physiological activities. We also performed a systematic analysis, classification and functional prediction of kinesins in soybean, the important cash crop, and found that the numbers of soybean kinesins are largely expanded. Taking advantage of the soybean transcriptome data in the public database, the functions of some soybean kinesins were predicted, to provide some clues for the study of kinesin functions in soybean and other crops.

Key words: kinesin, microtubule arrays, physiological regulation, soybean

王韫慧, 王一帆, 蔺佳雨, 李金红, 姚士恩, 冯湘池, 曹振林, 王俊, 李美娜. 植物驱动蛋白: 从微管阵列到生理活动调控. 植物学报, 2022, 57(3): 358-374.

Yunhui Wang, Yifan Wang, Jiayu Lin, Jinhong Li, Shien Yao, Xiangchi Feng, Zhenlin Cao, Jun Wang, Meina Li. Plant Kinesin: from Microtubule Arrays to Physiological Regulation. Chinese Bulletin of Botany, 2022, 57(3): 358-374.

导出引用管理器 EndNote|Ris|BibTeX

链接本文: https://www.chinbullbotany.com/CN/10.11983/CBB22007

https://www.chinbullbotany.com/CN/Y2022/V57/I3/358

图/表 5

参考文献 96

[1]	沈锦波, 姜里文 (2018). 中国科学家在植物细胞骨架研究领域取得突破性进展. 植物学报 53, 741-744. DOI
[2]	岳剑茹, 赫云建, 邱天麒, 郭南南, 韩雪萍, 王显玲 (2021). 植物微管骨架参与下胚轴伸长调节机制研究进展. 植物学报 56, 363-371.
[3]	Albertsen MC, Palmer RG (1979). A comparative light- and electron-microscopic study of microsporogenesis in male sterile (MS₁) and male fertile soybeans (Glycine max (L.) Merr.). Am J Bot 66, 253-265. DOI URL
[4]	Ambrose JC, Cyr R (2007). The kinesin ATK5 functions in early spindle assembly in Arabidopsis. Plant Cell 19, 226- 236. DOI URL
[5]	Ambrose JC, Li WX, Marcus A, Ma H, Cyr R (2005). A minus-end-directed kinesin with plus-end tracking protein activity is involved in spindle morphogenesis. Mol Biol Cell 16, 1584-1592.
[6]	Asada T, Kuriyama R, Shibaoka H (1997). TKRP125, a kinesin-related protein involved in the centrosome-independent organization of the cytokinetic apparatus in tobacco BY-2 cells. J Cell Sci 110, 179-189. DOI URL
[7]	Bannigan A, Scheible WR, Lukowitz W, Fagerstrom C, Wadsworth P, Somerville C, Baskin TI (2007). A conserved role for kinesin-5 in plant mitosis. J Cell Sci 120, 2819-2827. DOI PMID
[8]	Bieling P, Telley IA, Surrey T (2010). A minimal midzone protein module controls formation and length of antiparallel microtubule overlaps. Cell 142, 420-432. DOI PMID
[9]	Blancaflor EB (2013). Regulation of plant gravity sensing and signaling by the actin cytoskeleton. Am J Bot 100, 143-152. DOI PMID
[10]	Brady ST (1985). A novel brain ATPase with properties expected for the fast axonal transport motor. Nature 317, 73-75. DOI URL
[11]	Brim CA, Young MF (1971). Inheritance of a male-sterile character in soybeans. Crop Sci 11, 564-566. DOI URL
[12]	Buschmann H, Dols J, Kopischke S, Peña EJ, Andrade- Navarro MA, Heinlein M, Szymanski DB, Zachgo S, Doonan JH, Lloyd CW (2015). Arabidopsis KCBP interacts with AIR9 but stays in the cortical division zone throughout mitosis via its MyTH4-FERM domain. J Cell Sci 128, 2033-2046.
[13]	Chandrasekaran G, Tátrai P, Gergely F (2015). Hitting the brakes: targeting microtubule motors in cancer. Br J Cancer 113, 693-698. DOI URL
[14]	Chen CB, Marcus A, Li WX, Hu Y, Calzada JPV, Grossniklaus U, Cyr JR, Ma H (2002). The Arabidopsis ATK1 gene is required for spindle morphogenesis in male meiosis. Development 129, 2401-2409. DOI URL
[15]	Dawe RK, Lowry EG, Gent JI, Stitzer MC, Swentowsky KW, Higgins DM, Ross-Ibarra J, Wallace JG, Kanizay LB, Alabady M, Qiu WH, Tseng KF, Wang N, Gao Z, Birchler JA, Harkess AE, Hodges AL, Hiatt EN (2018). A kinesin-14 motor activates neocentromeres to promote meiotic drive in maize. Cell 173, 839-850. DOI
[16]	de Cuevas M, Tao T, Goldstein LSB (1992). Evidence that the stalk of Drosophila kinesin heavy chain is an α-helical coiled coil. J Cell Biol 116, 957-965. DOI PMID
[17]	de Keijzer J, Kieft H, Ketelaar T, Goshima G, Janson ME (2017). Shortening of microtubule overlap regions defines membrane delivery sites during plant cytokinesis. Curr Biol 27, 514-520. DOI PMID
[18]	Desai A, Verma S, Mitchison TJ, Walczak CE (1999). Kin I kinesins are microtubule-destabilizing enzymes. Cell 96, 69-78. DOI PMID
[19]	Duroc Y, Lemhemdi A, Larchevêque C, Hurel A, Cuacos M, Cromer L, Horlow C, Armstrong SJ, Chelysheva L, Mercier R (2014). The kinesin AtPSS1 promotes synapsis and is required for proper crossover distribution in meiosis. PLoS Genet 10, e1004674.
[20]	Eng RC, Wasteneys GO (2014). The microtubule plus-end tracking protein ARMADILLO-REPEAT KINESIN 1 promotes microtubule catastrophe in Arabidopsis. Plant Cell 26, 3372-3386. DOI URL
[21]	Fang XL, Sun XY, Yang XD, Li Q, Lin CJ, Xu J, Gong WJ, Wang YF, Liu L, Zhao LM, Liu BH, Qin J, Zhang MC, Zhang CB, Kong FJ, Li MN (2021). MS1 is essential for male fertility by regulating the microsporocyte cell plate expansion in soybean. Sci China Life Sci 64, 1533-1545. DOI URL
[22]	Frey N, Klotz J, Nick P (2009). Dynamic bridges—a calponin-domain kinesin from rice links actin filaments and microtubules in both cycling and non-cycling cells. Plant Cell Physiol 50, 1493-1506. DOI URL
[23]	Fu Y, Li H, Yang ZB (2002). The ROP2 GTPase controls the formation of cortical fine F-actin and the early phase of directional cell expansion during Arabidopsis organogenesis. Plant Cell 14, 777-794. DOI URL
[24]	Ganguly A, DeMott L, Zhu CM, McClosky DD, Anderson CT, Dixit R (2018). Importin-β directly regulates the motor activity and turnover of a Kinesin-4. Dev Cell 44, 642-651. DOI
[25]	Ganguly A, Zhu CM, Chen WZ, Dixit R (2020). FRA1 kinesin modulates the lateral stability of cortical microtubules through cellulose synthase-microtubule uncoupling proteins. Plant Cell 32, 2508-2524. DOI URL
[26]	Gicking AM, Wang P, Liu C, Mickolajczyk KJ, Guo LJ, Hancock WO, Qiu WH (2019). The orphan kinesin PAKRP2 achieves processive motility via a noncanonical stepping mechanism. Biophys J 116, 1270-1281. DOI
[27]	Hamada T (2014). Microtubule organization and microtubule-associated proteins in plant cells. Int Rev Cel Mol Biol 312, 1-52.
[28]	Hedden P, Phillips AL (2000). Gibberellin metabolism: new insights revealed by the genes. Trends Plant Sci 5, 523- 530. DOI PMID
[29]	Herrmann A, Livanos P, Zimmermann S, Berendzen K, Rohr L, Lipka E, Müller S (2021). KINESIN-12E regulates metaphase spindle flux and helps control spindle size in Arabidopsis. Plant Cell 33, 27-43.
[30]	Higgins DM, Nannas NJ, Dawe RK (2016). The maize divergent spindle-1 (dv1) gene encodes a Kinesin-14A motor protein required for meiotic spindle pole organization. Front Plant Sci 7, 1277. DOI PMID
[31]	Hiwatashi Y, Obara M, Sato Y, Fujita T, Murata T, Hasebe M (2008). Kinesins are indispensable for interdigitation of phragmoplast microtubules in the Moss Physcomitrella pa-tens. Plant Cell 20, 3094-3106. DOI PMID
[32]	Hiwatashi Y, Sato Y, Doonan JH (2014). Kinesins have a dual function in organizing microtubules during both tip growth and cytokinesis in Physcomitrella patens. Plant Cell 26, 1256-1266. DOI URL
[33]	Ho CMK, Hotta T, Guo FL, Roberson RW, Lee YRJ, Liu B (2011). Interaction of antiparallel microtubules in the phragmoplast is mediated by the microtubule-associated pro- tein MAP65-3 in Arabidopsis. Plant Cell 23, 2909-2923. DOI URL
[34]	Hu CK, Coughlin M, Field CM, Mitchison TJ (2011). KIF4 regulates midzone length during cytokinesis. Curr Biol 21, 815-824. DOI URL
[35]	Huang YC, Wang HH, Huang X, Wang Q, Wang JC, An D, Li JQ, Wang WQ, Wu YR (2019). Maize VKS1 regulates mitosis and cytokinesis during early endosperm development. Plant Cell 31, 1238-1256. DOI URL
[36]	Itoh R, Fujiwara M, Yoshida S (2001). Kinesin-related proteins with a mitochondrial targeting signal. Plant Physiol 127, 724-726. PMID
[37]	Iwata S, Morikawa M, Takei Y, Hirokawa N (2020). An activity-dependent local transport regulation via degradation and synthesis of KIF17 underlying cognitive flexibility. Sci Adv 6, eabc8355.
[38]	Jones MA, Raymond MJ, Smirnoff N (2006). Analysis of the root-hair morphogenesis transcriptome reveals the molecular identity of six genes with roles in root-hair development in Arabidopsis. Plant J 45, 83-100. PMID
[39]	Kadota A, Yamada N, Suetsugu N, Hirose M, Saito C, Shoda K, Ichikawa S, Kagawa T, Nakano A, Wada M (2009). Short actin-based mechanism for light-directed chloroplast movement in Arabidopsis. Proc Natl Acad Sci USA 106, 13106-13111. DOI URL
[40]	Kasahara M, Kagawa T, Oikawa K, Suetsugu N, Miyao M, Wada M (2002). Chloroplast avoidance movement reduces photodamage in plants. Nature 420, 829-832. DOI URL
[41]	Kong ZS, Ioki M, Braybrook S, Li SD, Ye ZH, Lee YRJ, Hotta T, Chang A, Tian J, Wang GD, Liu B (2015). Kinesin-4 functions in vesicular transport on cortical micro- tubules and regulates cell wall mechanics during cell elongation in plants. Mol Plant 8, 1011-1023. DOI URL
[42]	Lane J, Allan V (1998). Microtubule-based membrane movement. Biochim Biophys Acta 1376, 27-55. PMID
[43]	Lawrence CJ, Dawe RK, Christie KR, Cleveland DW, Dawson SC, Endow SA, Goldstein LSB, Goodson HV, Hirokawa N, Howard J, Malmberg RL, McIntosh JR, Miki H, Mitchison TJ, Okada Y, Reddy ASN, Saxton WM, Schliwa M, Scholey JM, Vale RD, Walczak CE, Wordeman L (2004). A standardized kinesin nomenclature. J Cell Biol 167, 19-22. DOI PMID
[44]	Lee YRJ, Giang HM, Liu B (2001). A novel plant kinesin-related protein specifically associates with the phragmoplast organelles. Plant Cell 13, 2427-2439. DOI PMID
[45]	Lee YRJ, Li Y, Liu B (2007). Two Arabidopsis phragmoplast-associated kinesins play a critical role in cytokinesis during male gametogenesis. Plant Cell 19, 2595-2605. DOI URL
[46]	Lee YRJ, Liu B (2000). Identification of a phragmoplast- associated kinesin-related protein in higher plants. Curr Biol 10, 797-800. DOI PMID
[47]	Lee YRJ, Liu B (2004). Cytoskeletal motors in Arabidopsis. Sixty-one kinesins and seventeen myosins. Plant Physiol 136, 3877-3883. DOI URL
[48]	Lee YRJ, Qiu WH, Liu B (2015). Kinesin motors in plants: from subcellular dynamics to motility regulation. Curr Opin Plant Biol 28, 120-126. DOI URL
[49]	Li J, Jia JF, Qian Q, Yun YY, Zhang C, Xiao J, Du C, Luo W, Zou GX, Chen ML, Huang YQ, Feng YQ, Cheng ZK, Yuan M, Chong K (2011). Mutation of rice BC12/GDD1, which encodes a kinesin-like protein that binds to a GA biosynthesis gene promoter, leads to dwarfism with impaired cell elongation. Plant Cell 23, 628-640. DOI URL
[50]	Li YF, Deng ZG, Kamisugi Y, Chen ZR, Wang JJ, Han X, Wei YX, He H, Terzaghi W, Cove DJ, Cuming AC, Chen HD (2021). A minus-end directed kinesin motor directs gravitropism in Physcomitrella patens. Nat Commun 12, 4470. DOI URL
[51]	Malcos JL, Cyr RJ (2011). An ungrouped plant kinesin accumulates at the preprophase band in a cell cycle- dependent manner. Cytoskeleton 68, 247-258. DOI URL
[52]	Marcus AI, Li W, Ma H, Cyr RJ (2003). A kinesin mutant with an atypical bipolar spindle undergoes normal mitosis. Mol Biol Cell 14, 1717-1726. DOI PMID
[53]	McFarlane HE, Döring A, Persson S (2014). The cell biology of cellulose synthesis. Annu Rev Plant Biol 65, 69-94. DOI PMID
[54]	Miki H, Okada Y, Hirokawa N (2005). Analysis of the kinesin superfamily: insights into structure and function. Trends Cell Biol 15, 467-476. DOI URL
[55]	Mineyuki Y (1999). The preprophase band of microtubules: its function as a cytokinetic apparatus in higher plants. Int Rev Cytol 187, 1-49.
[56]	Moschou PN, Gutierrez-Beltran E, Bozhkov PV, Smertenko A (2016). Separase promotes microtubule polymerization by activating CENP-E-related kinesin Kin7. Dev Cell 37, 350-361. DOI URL
[57]	Moschou PN, Smertenko AP, Minina EA, Fukada K, Savenkov EI, Robert S, Hussey PJ, Bozhkov PV (2013). The caspase-related protease separase (EXTRA SPINDLE POLES) regulates cell polarity and cytokinesis in Arabidopsis. Plant Cell 25, 2171-2186. DOI URL
[58]	Müller S, Han SC, Smith LG (2006). Two kinesins are involved in the spatial control of cytokinesis in Arabidopsis thaliana. Curr Biol 16, 888-894. DOI URL
[59]	Myers SM, Collins I (2016). Recent findings and future directions for interpolar mitotic kinesin inhibitors in cancer therapy. Future Med Chem 8, 463-489. DOI URL
[60]	Nakamura M, Toyota M, Tasaka M, Morita MT (2011). An Arabidopsis E3 ligase, SHOOT GRAVITROPISM 9, modu- lates the interaction between statoliths and F-actin in gravity sensing. Plant Cell 23, 1830-1848. DOI URL
[61]	Nebenführ A, Dixit R (2018). Kinesins and myosins: molecular motors that coordinate cellular functions in plants. Annu Rev Plant Biol 69, 329-361. DOI PMID
[62]	Ni CZ, Wang HQ, Xu T, Qu Z, Liu GQ (2005). AtKP1, a kinesin-like protein, mainly localizes to mitochondria in Arabidopsis thaliana. Cell Res 15, 725-733. DOI URL
[63]	Nishihama R, Ishikawa M, Araki S, Soyano T, Asada T, Machida Y (2001). The NPK1 mitogen-activated protein kinase kinase kinase is a regulator of cell-plate formation in plant cytokinesis. Genes Dev 15, 352-363. DOI URL
[64]	Nishihama R, Soyano T, Ishikawa M, Araki S, Tanaka H, Asada T, Irie K, Ito M, Terada M, Banno H, Yamazaki Y, Machida Y (2002). Expansion of the cell plate in plant cytokinesis requires a kinesin-like protein/MAPKKK complex. Cell 109, 87-99. DOI PMID
[65]	Oda Y, Fukuda H (2013). Rho of plant Gtpase signaling regulates the behavior of Arabidopsis kinesin-13A to establish secondary cell wall patterns. Plant Cell 25, 4439- 4450. DOI URL
[66]	Oh SA, Allen T, Kim GJ, Sidorova A, Borg M, Park SK, Twell D (2012). Arabidopsis fused kinase and the Kinesin-12 subfamily constitute a signaling module required for phragmoplast expansion. Plant J 72, 308-319. DOI URL
[67]	Oh SA, Bourdon V, Das 'Pal M, Dickinson H, Twell D (2008). Arabidopsis kinesins HINKEL and TETRASPORE act redundantly to control cell plate expansion during cytokinesis in the male gametophyte. Mol Plant 1, 794-799. DOI URL
[68]	Olszewski N, Sun TP, Gubler F (2002). Gibberellin signaling: biosynthesis, catabolism, and response pathways. Plant Cell 14, S61-S80. DOI URL
[69]	Pan RQ, Lee YRJ, Liu B (2004). Localization of two homologous Arabidopsis kinesin-related proteins in the phragmoplast. Planta 220, 156-164. DOI URL
[70]	Preuss ML, Kovar DR, Lee YRJ, Staiger CJ, Delmer DP, Liu B (2004). A plant-specific kinesin binds to actin microfilaments and interacts with cortical microtubules in cotton fibers. Plant Physiol 136, 3945-3955. DOI PMID
[71]	Rath O, Kozielski F (2012). Kinesins and cancer. Nat Rev Cancer 12, 527-539. DOI URL
[72]	Sasabe M, Machida Y (2012). Regulation of organization and function of microtubules by the mitogen-activated protein kinase cascade during plant cytokinesis. Cytoskeleton 69, 913-918. DOI URL
[73]	Song Y, Li G, Nowak J, Zhang XQ, Xu DB, Yang XJ, Huang GQ, Liang WQ, Yang LT, Wang CH, Bulone V, Nikoloski Z, Hu JP, Persson S, Zhang DB (2019). The rice actin-binding protein RMD regulates light-dependent shoot gravitropism. Plant Physiol 181, 630-644.
[74]	Strompen G, El Kasmi F, Richter S, Lukowitz W, Assaad FF, Jürgens G, Mayer U (2002). The Arabidopsis HINKEL gene encodes a kinesin-related protein involved in cytokinesis and is expressed in a cell cycle-dependent manner. Curr Biol 12, 153-158. DOI PMID
[75]	Suetsugu N, Yamada N, Kagawa T, Yonekura H, Uyeda TQP, Kadota A, Wada M (2010). Two kinesin-like proteins mediate actin-based chloroplast movement in Arabidopsis thaliana. Proc Natl Acad Sci USA 107, 8860-8865. DOI URL
[76]	Tian J, Han LB, Feng ZD, Wang GD, Liu WW, Ma YP, Yu YJ, Kong ZS (2015). Orchestration of microtubules and the actin cytoskeleton in trichome cell shape determination by a plant-unique kinesin. eLife 4, e09351. DOI URL
[77]	Tseng KF, Wang P, Lee YRJ, Bowen J, Gicking AM, Guo LJ, Liu B, Qiu WH (2018). The preprophase band-associated Kinesin-14 OsKCH2 is a processive minus-end- directed microtubule motor. Nat Commun 9, 1067. DOI URL
[78]	Vale RD (2003). The molecular motor toolbox for intracellular transport. Cell 112, 467-480. DOI URL
[79]	Vale RD, Fletterick RJ (1997). The design plan of kinesin motors. Annu Rev Cell Dev Biol 13, 745-777. PMID
[80]	Vale RD, Reese TS, Sheetz MP (1985). Identification of a novel force-generating protein, kinesin, involved in microtubule-based motility. Cell 42, 39-50. DOI PMID
[81]	Walker KL, Müller S, Moss D, Ehrhardt DW, Smith LG (2007). Arabidopsis TANGLED identifies the division plane throughout mitosis and cytokinesis. Curr Biol 17, 1827- 1836. DOI PMID
[82]	Wang HQ, Liu RJ, Wang JW, Wang P, Shen PWY, Liu GQ (2014). The Arabidopsis kinesin gene AtKin-1 plays a role in the nuclear division process during megagametogenesis. Plant Cell Rep 33, 819-828. DOI URL
[83]	Wick SM, Seagull RW, Osborn M, Weber K, Gunning BES (1981). Immunofluorescence microscopy of organized mic- rotubule arrays in structurally stabilized meristematic plant cells. J Cell Biol 89, 685-690. PMID
[84]	Xu XM, Zhao Q, Rodrigo-Peiris T, Brkljacic J, He CS, Müller S, Meier I (2008). RanGAP1 is a continuous marker of the Arabidopsis cell division plane. Proc Natl Acad Sci USA 105, 18637-18642. DOI URL
[85]	Yamada M, Tanaka-Takiguchi Y, Hayashi M, Nishina M, Goshima G (2017). Multiple Kinesin-14 family members drive microtubule minus end-directed transport in plant cells. J Cell Biol 216, 1705-1714. DOI PMID
[86]	Yang CY, Spielman M, Coles JP, Li Y, Ghelani S, Bourdon V, Brown RC, Lemmon BE, Scott RJ, Dickinson HG (2003). TETRASPORE encodes a kinesin required for male meiotic cytokinesis in Arabidopsis. Plant J 34, 229- 240. DOI PMID
[87]	Yang GH, Gao P, Zhang H, Huang SJ, Zheng ZL (2007). A mutation in MRH2 kinesin enhances the root hair tip growth defect caused by constitutively activated ROP2 small GTPase in Arabidopsis. PLoS One 2, e1074. DOI URL
[88]	Yang XY, Chen ZW, Xu T, Qu Z, Pan XD, Qin XH, Ren DT, Liu GQ (2011). Arabidopsis kinesin KP1 specifically interacts with VDAC3, a mitochondrial protein, and regulates respiration during seed germination at low temperature. Plant Cell 23, 1093-1106. DOI URL
[89]	Yin XL, Takei Y, Kido MA, Hirokawa N (2011). Molecular motor KIF17 is fundamental for memory and learning via differential support of synaptic NR2A/2B levels. Neuron 70, 310-325. DOI URL
[90]	Yuan M, Shaw PJ, Warn RM, Lloyd CW (1994). Dynamic reorientation of cortical microtubules, from transverse to longitudinal, in living plant cells. Proc Natl Acad Sci USA 91, 6050-6053. DOI URL
[91]	Zhang M, Zhang BC, Qian Q, Yu YC, Li R, Zhang JW, Liu XL, Zeng DL, Li JY, Zhou YH (2010). Brittle Culm 12, a dual-targeting Kinesin-4 protein, controls cell-cycle progression and wall properties in rice. Plant J 63, 312-328. DOI URL
[92]	Zhong RQ, Burk DH, Morrison III WH, Ye ZH (2002). A kinesin-like protein is essential for oriented deposition of cellulose microfibrils and cell wall strength. Plant Cell 14, 3101-3117. DOI URL
[93]	Zhou SR, Wang Y, Li WC, Zhao ZG, Ren YL, Wang Y, Gu SH, Lin QB, Wang D, Jiang L, Su N, Zhang X, Liu LL, Cheng ZJ, Lei CL, Wang JL, Guo XP, Wu FQ, Ikehashi H, Wang HY, Wan JM (2011). Pollen Semi-Sterility 1 encodes a kinesin-1-like protein important for male meiosis, anther dehiscence, and fertility in rice. Plant Cell 23, 111-129. DOI URL
[94]	Zhu CM, Dixit R (2012). Functions of the Arabidopsis kinesin superfamily of microtubule-based motor proteins. Protoplasma 249, 887-899. DOI URL
[95]	Zhu CM, Ganguly A, Baskin TI, McClosky DD, Anderson CT, Foster C, Meunier KA, Okamoto R, Berg H, Dixit R (2015). The Fragile Fiber 1 kinesin contributes to cortical microtubule-mediated trafficking of cell wall components. Plant Physiol 167, 780-792. DOI
[96]	Zou JJ, Zheng ZY, Xue S, Li HH, Wang YR, Le J (2016). The role of Arabidopsis Actin-Related Protein 3 in amyloplast sedimentation and polar auxin transport in root gravitropism. J Exp Bot 67, 5325-5337. DOI URL

蛋白家族	物种	基因名	基因登录号	细胞学功能	生理学功能	参考文献
KIN1	拟南芥	AtKin-1/AtPSS1	AT3G63480	减数分裂调控纺锤体形态	育性控制	Wang et al., 2014
	水稻	OsPSS1	Os08g0117000	减数分裂调控纺锤体形态	育性控制	Zhou et al., 2011
KIN4	拟南芥	AtKinesin-4A/FRA1	AT5G47820		合成细胞壁	Zhu et al., 2015
	水稻	BC2/GDD1	Os09g0114500		合成细胞壁; 调控细胞延伸	Li et al., 2011
	苔藓	Kin4-Ia	Pp3c25_8700	抑制成膜体中间区的微管生长		de Keijzer et al., 2017
	苔藓	Kin4-Ic	Pp3c10_9510	抑制成膜体中间区的微管生长		de Keijzer et al., 2017
KIN5	拟南芥	RSW7/AtKRP125c	AT2g28620	有丝分裂间期交联微管, 前中期稳定纺锤体两极间微管阵列		Bannigan et al., 2007
KIN7	拟南芥	AtNACK1/HINKEL	AT1G18370	有丝分裂及减数分裂II末期促进成膜体解聚	育性控制	Strompen et al., 2002
	拟南芥	STUD/TES/AtNACK2	AT3G43210		育性控制	Yang et al., 2003
	烟草	NACK1	LOC107780313		育性控制	Nishihama et al., 2002
	烟草	NACK2	LOC107775653		育性控制	Nishihama et al., 2002
	大豆	GmNACK2	Glyma.13G114200		育性控制	Fang et al., 2021
蛋白家族	物种	基因名	基因登录号	细胞学功能	生理学功能		参考文献
	拟南芥	Kinesin7	AT3g12020	有丝分裂间期促进微管组装			Moschou et al., 2016
	拟南芥	MKRP1	AT1G21730		与线粒体共定位		Itoh et al., 2001
	拟南芥	MKRP2	AT4G39050		与线粒体共定位		Itoh et al., 2001
KIN10	拟南芥	AtPAKRP2	AT4g14330	组装细胞板			Lee et al., 2001; Gicking et al., 2019
	苔藓	KINID1a	AB434497	交联成膜体反向平行微管阵列			Hiwatashi et al., 2008, 2014
	苔藓	KINID1b	AB434498	交联成膜体反向平行微管阵列			Hiwatashi et al., 2008, 2014
KIN12	拟南芥	PAKRP1/Kinesin-12A	AT4G14150	稳定成膜体反向平行微管阵列			Lee and Liu, 2000; Pan et al., 2004
	拟南芥	PAKRP1L/Kinesin-12B	AT3G23670	稳定成膜体反向平行微管阵列			Lee and Liu, 2000; Pan et al., 2004
	拟南芥	POK1	AT3G17360	标识周质分裂位点			Müller et al., 2006
	拟南芥	POK2	AT3G19050	标识周质分裂位点			Müller et al., 2006
	拟南芥	Kinesin-12E	AT3G44050	有丝分裂组装纺锤体			Herrmann et al., 2021
KIN13	拟南芥	AtKinesin-13A/AtMACK	AT3G16630	有丝分裂间期解聚微管			Oda and Fukuda, 2013
KIN14	拟南芥	KATA/ATK1	AT4G21270	有丝分裂向纺锤体两极拉动微管			Chen et al., 2002; Marcus et al., 2003
	拟南芥	ATK5	AT4G05190	有丝分裂纺锤体中间区交联微管			Ambrose and Cyr, 2007
	拟南芥	ATKP1	AT3G44730	沿微管运输微丝	与线粒体共定位		Yang et al., 2011
	水稻	OsKCH1	Os12g0547500				Frey et al., 2009
	水稻	OsKCH2	未公布				Tseng et al., 2018
	拟南芥	KAC1	AT5G10470		介导光驱动叶绿体		Suetsugu et al., 2010
	拟南芥	KAC2	AT5G65460		介导光驱动叶绿体		Suetsugu et al., 2010
	棉花	GhKCH1	AY695833				Preuss et al., 2004
	棉花	GhKCH2	EF432568				Xu et al., 2009
	拟南芥	KCBP/ZWI	AT5G65930	有丝分裂间期交联微管微丝, 后期将成膜体外围微管拉向中间区			Tian et al., 2015
	玉米	VSK1	Zm00001d018624	有丝分裂组装纺锤体			Huang et al., 2019
	苔藓	KINDR	KX759199- KX759203	激活着丝粒			Dawe et al., 2018
	苔藓	GTRC	Pp3c2_9150	调控微丝极性	参与重力感受		Li et al., 2021
KINU	拟南芥	ARK1/AtKINUc	AT3G54870	有丝分裂间期促进微管灾变			Yang et al., 2007; Eng and Wasteneys, 2014
	拟南芥	ARK3/AtKINUa	AT1G12430	有丝分裂促进早前期微管带的形成与稳定			Malcos and Cyr, 2011

蛋白家族	物种	基因名	基因登录号	细胞学功能	生理学功能	参考文献
KIN1	拟南芥	AtKin-1/AtPSS1	AT3G63480	减数分裂调控纺锤体形态	育性控制	Wang et al., 2014
	水稻	OsPSS1	Os08g0117000	减数分裂调控纺锤体形态	育性控制	Zhou et al., 2011
KIN4	拟南芥	AtKinesin-4A/FRA1	AT5G47820		合成细胞壁	Zhu et al., 2015
	水稻	BC2/GDD1	Os09g0114500		合成细胞壁; 调控细胞延伸	Li et al., 2011
	苔藓	Kin4-Ia	Pp3c25_8700	抑制成膜体中间区的微管生长		de Keijzer et al., 2017
	苔藓	Kin4-Ic	Pp3c10_9510	抑制成膜体中间区的微管生长		de Keijzer et al., 2017
KIN5	拟南芥	RSW7/AtKRP125c	AT2g28620	有丝分裂间期交联微管, 前中期稳定纺锤体两极间微管阵列		Bannigan et al., 2007
KIN7	拟南芥	AtNACK1/HINKEL	AT1G18370	有丝分裂及减数分裂II末期促进成膜体解聚	育性控制	Strompen et al., 2002
	拟南芥	STUD/TES/AtNACK2	AT3G43210		育性控制	Yang et al., 2003
	烟草	NACK1	LOC107780313		育性控制	Nishihama et al., 2002
	烟草	NACK2	LOC107775653		育性控制	Nishihama et al., 2002
	大豆	GmNACK2	Glyma.13G114200		育性控制	Fang et al., 2021
蛋白家族	物种	基因名	基因登录号	细胞学功能	生理学功能		参考文献
	拟南芥	Kinesin7	AT3g12020	有丝分裂间期促进微管组装			Moschou et al., 2016
	拟南芥	MKRP1	AT1G21730		与线粒体共定位		Itoh et al., 2001
	拟南芥	MKRP2	AT4G39050		与线粒体共定位		Itoh et al., 2001
KIN10	拟南芥	AtPAKRP2	AT4g14330	组装细胞板			Lee et al., 2001; Gicking et al., 2019
	苔藓	KINID1a	AB434497	交联成膜体反向平行微管阵列			Hiwatashi et al., 2008, 2014
	苔藓	KINID1b	AB434498	交联成膜体反向平行微管阵列			Hiwatashi et al., 2008, 2014
KIN12	拟南芥	PAKRP1/Kinesin-12A	AT4G14150	稳定成膜体反向平行微管阵列			Lee and Liu, 2000; Pan et al., 2004
	拟南芥	PAKRP1L/Kinesin-12B	AT3G23670	稳定成膜体反向平行微管阵列			Lee and Liu, 2000; Pan et al., 2004
	拟南芥	POK1	AT3G17360	标识周质分裂位点			Müller et al., 2006
	拟南芥	POK2	AT3G19050	标识周质分裂位点			Müller et al., 2006
	拟南芥	Kinesin-12E	AT3G44050	有丝分裂组装纺锤体			Herrmann et al., 2021
KIN13	拟南芥	AtKinesin-13A/AtMACK	AT3G16630	有丝分裂间期解聚微管			Oda and Fukuda, 2013
KIN14	拟南芥	KATA/ATK1	AT4G21270	有丝分裂向纺锤体两极拉动微管			Chen et al., 2002; Marcus et al., 2003
	拟南芥	ATK5	AT4G05190	有丝分裂纺锤体中间区交联微管			Ambrose and Cyr, 2007
	拟南芥	ATKP1	AT3G44730	沿微管运输微丝	与线粒体共定位		Yang et al., 2011
	水稻	OsKCH1	Os12g0547500				Frey et al., 2009
	水稻	OsKCH2	未公布				Tseng et al., 2018
	拟南芥	KAC1	AT5G10470		介导光驱动叶绿体		Suetsugu et al., 2010
	拟南芥	KAC2	AT5G65460		介导光驱动叶绿体		Suetsugu et al., 2010
	棉花	GhKCH1	AY695833				Preuss et al., 2004
	棉花	GhKCH2	EF432568				Xu et al., 2009
	拟南芥	KCBP/ZWI	AT5G65930	有丝分裂间期交联微管微丝, 后期将成膜体外围微管拉向中间区			Tian et al., 2015
	玉米	VSK1	Zm00001d018624	有丝分裂组装纺锤体			Huang et al., 2019
	苔藓	KINDR	KX759199- KX759203	激活着丝粒			Dawe et al., 2018
	苔藓	GTRC	Pp3c2_9150	调控微丝极性	参与重力感受		Li et al., 2021
KINU	拟南芥	ARK1/AtKINUc	AT3G54870	有丝分裂间期促进微管灾变			Yang et al., 2007; Eng and Wasteneys, 2014
	拟南芥	ARK3/AtKINUa	AT1G12430	有丝分裂促进早前期微管带的形成与稳定			Malcos and Cyr, 2011