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水稻剑蛋白亚基OsKTN80a参与微管动态的功能分析
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摘要
微管作为细胞骨架的一种,对于细胞的形态建成、细胞内的物质运输、细胞分裂、一些特殊结构的组成(如鞭毛、纤毛)和植物细胞壁的合成等重要的生命活动是必不可少的。微管具有高度动态的特性,原因之一是受到微管相关蛋白的调控。Katanin是一类可以切割微管的微管相关蛋白,由两个亚基组成,P60和P80,P60具有ATPase的活性和微管切割的活性;P80不具有微管切割的活性,其主要负责将P60导向中心体和调控P60的微管切割活性。P60本身参与了许多重要的生命活动,而P80亦在线虫减数分裂、人神经细胞发育、小鼠雄配子发生、低等生物的cytokinesis以及鞭毛发生中扮演重要角色。在拟南芥细胞中,KATANIN P60参与了细胞各向异性的延伸和根尖细胞的有丝分裂,并且作为生长素信号传导末端的效应子参与皮层微管排列。但是高等植物中P80的功能仍然不甚明了。
     在本研究中,我们利用水稻中的KATANIN P60(KTN60)作为诱饵蛋白淘洗水稻幼穗的细菌双杂交cDNA文库,获得了OsKTN80b,通过BLAST水稻蛋白数据库获得了另外两个同源蛋白(OsKTN80a和OsKTN80c),这3个同源蛋白均与人和小鼠的KATANIN P80具有极高的序列相似性。通过酵母双杂交验证、结构域分析和进化关联分析,证实这3个基因即为katanin p80在水稻中的同源基因。我们在水稻中过表达OsKTN80a (KTN80a-OX),导致水稻幼苗根的生长受阻。分析发现是由两个原因造成的:一是KTN80a-OX根尖细胞的延长障碍,主要是由于KTN80a过表达后导致伸长区细胞皮层微管的排列异常;二是KTN80a-OX根尖细胞的分裂滞留于有丝分裂后的细胞质分裂(cytokinesis)时期,可能是由于遗传物质的分离异常以及成膜体微管的形态和功能异常。上述两个原因可以解释为KTN80a抑制了KTN60的微管切割活性,从而导致微管的动态受到破坏。证据如下:1)KTN80a和KTN60相互作用;2)KTN80a-OX的根尖细胞变短和伸长区细胞皮层微管变得紊乱的表型均和KTN60的突变体(fra2)极为相似。3)当KTN80a过表达后,和野生型相比,其根尖的生长和伸长区的皮层微管更加耐受微管特异性的去稳定药物oryzalin的处理,表明KTN80a促使微管更稳定。
     我们讨论了KTN80a参与细胞延伸和细胞分裂的分子机制。从上述KTN80a的作用机制可以看出,控制KATANIN的活性对于水稻根尖细胞的伸长和细胞的分裂是必须的。
As a kind of cytoskeletons, microtubules are essential for cell morphogenesis, the intracellular transportation, cell division, composition of some novel cellular apparatus such as flagellum and cilia and the plant cell wall biosynthesis and so on. Microtubules are highly dynamic and one of the reasons is that it was regulated by microtubule associated proteins (MAPs). Katanin, a kind of MAP involved in microtubule severing, is composed of two subunits, one is P60which has ATPase and microtubule severing activity and the other is P80which has no microtubule severing activity but plays roles in targeting P60to the centrosome and regulating P60's microtubule severing activity. P60itself are involved in many significant life activities while P80also functions essentially in C. elegans meiosis, human neuronal development, the male gamete genesis in mouse and in cytokinesis and flagella biogenesis in lower order species. KATANIN P60is necessary for the anisotropic cell expansion and cell division in root tip and also acts as the effector in cortical microtubule orientation in auxin signaling pathway in Arabidopsis cell. When it comes to the other KATANIN subunit P80in higher plants, its function remain largely unknown.
     In this study, we used rice KATANIN P60(KTN60) as the bait to screen the bacteria two hybrid cDNA library of rice immature panicle and obtained a KTN80b. Through BLAST the protein database of Oryza sativa using this protein, we found the other two homologous proteins (KTN80a and KTN80c) in rice. There are high sequence similarity between the3homologous proteins mentioned above and the KATANIN P80in human and mouse. Later the3homologs are demonstrated as katanin p80orthologs in rice after yeast two hybrid assay, the domain analysis and the phylogenic analysis. The overexpression of OsKTN80a in rice (KTN80a-OX) caused the delayed primary root growth of rice seedling and two reasons results in it:the first is that cell elongation dysfunction in root tip in KTN80a-OX which is thought as the disordered cortical microtubule arrangement in elongation zone after Ktn80a overexpression; the second is that cell division are stalled at cytokinesis which are mainly caused by the incomplete separation of genetic materials into the daughter cells and the abnormal phragmoplast morphology and function. The above two reasons could be explained that the KTN60microtubule severing activity are repressed by KTN80a and the event caused the broken microtubule dynamics. There are3evidence supporting the idea:1) KTN80a interacts with KTN60;2) The phenotype of the shorter cell in root tip and the disordered cortical microtubule array in elongation zone when KTN80a is overexpressed could perfectly mimic the case in KTN60mutant (fra2);3) Compared with the wild type, the root tip growth and cortical microtubules in elongation zone are more resistant to treatment of oryzalin, a kind of microtubule-special destabilizing drug, which implies that KTN80a stabilizes microtubules.
     We discuss the molecular mechanism of the role of KTN80a plays in cell expansion and in cell division. From the above mechanism of KTN80a function in, we can get the conclusion that controlling the KATANIN activity is essential for cell expansion and cell division in rice root tip.
引文
Adeline Bichet, et al. (2001),'BOTERO1 is required for normal orientation of cortical microtubules and anisotropic cell expansion in Arabidopsis', Plant J,25 (2),137-48.
    Ambrose, J. C. and Wasteneys, G. O. (2008),'CLASP modulates microtubule-cortex interaction during self-organization of acentrosomal microtubules', Mol Biol Cell,19 (11),4730-7.
    Ambrose, J. C., et al. (2007),'The Arabidopsis CLASP gene encodes a microtubule-associated protein involved in cell expansion and division', Plant Cell,19 (9),2763-75.
    Andrey, V. Korolev, et al. (2007),'AtMAP70-5, a divergent member of the MAP70 family of microtubule-associated proteins, is required for anisotropic cell growth in Arabidopsis', Journal of Cell Science.
    Azimzadeh, J., et al. (2008), Arabidopsis TONNEAU1 proteins are essential for preprophase band formation and interact with centrin', Plant Cell,20 (8),2146-59.
    Baas, P. W., Karabay, A., and Qiang, L. (2005),'Microtubules cut and run', Trends Cell Biol,15 (10),518-24.
    Benz, C., et al. (2012),'Cytokinesis in bloodstream stage Trypanosoma brucei requires a family of katanins and spastin', PLoS One,7 (1), e30367.
    Bo, Liu, et al. (1994),'y-Tubulin in Arabidopsis:Gene Sequence, Immunoblot, and Immunofluorescence Studies', Plant Cell.
    Bouquin, T., et al. (2003),'The Arabidopsis Iue1 mutant defines a katanin p60 ortholog involved in hormonal control of microtubule orientation during cell growth', J Cell Sci,116 (Pt 5),791-801.
    Brown, R. C. and Lemmon, B. E. (2006),'Polar organizers and girdling bands of microtubules are associated with y-tubulin and act in establishment of meiotic quadripolarity in the hepatic Aneura pinguis (Bryophyta)', Protoplasma,227 (2-4),77-85.
    Burk, D. H. (2002), Alteration of Oriented Deposition of Cellulose Microfibrils by Mutation of a Katanin-Like Microtubule-Severing Protein', Plant Cell, 14 (9),2145-60.
    Burk, D. H., et al. (2001),'A katanin-like protein regulates normal cell wall biosynthesis and cell elongation', Plant Cell,13 (4),807-27.
    Buschmann, H., et al. (2006),'Microtubule-associated AIR9 recognizes the cortical division site at preprophase and cell-plate insertion', CurrBiol, 16(19),1938-43.
    Camilleri, C. (2002), The Arabidopsis TONNEAU2 Gene Encodes a Putative Novel Protein Phosphatase 2A Regulatory Subunit Essential for the Control of the Cortical Cytoskeleton', Plant Cell,14 (4),833-45.
    Casanova, M., et al. (2009),'Microtubule-severing proteins are involved in flagellar length control and mitosis in Trypanosomatids', Mol Microbiol, 71 (6),1353-70.
    Chan, J., et al. (2009),'Arabidopsis cortical microtubules are initiated along, as well as branching from, existing microtubules', Plant Cell,21 (8), 2298-306.
    Chen, R., et al. (2007),'Rice UDP-glucose pyrophosphorylase1 is essential for pollen callose deposition and its cosuppression results in a new type of thermosensitive genie male sterility', Plant Cell,19 (3),847-61.
    Cox, G. A., et al. (2000), The mouse fidgetin gene defines a new role for AAA family proteins in mammalian development', Nat Genet,26 (2), 198-202.
    Crowell, E. F, et al. (2010),'Regulation of anisotropic cell expansion in higher plants', C R Biol,333 (4),320-4.
    Diaz-Valencia, J. D., et al. (2011),'Drosophila katanin-60 depolymerizes and severs at microtubule defects', Biophys J,100 (10),2440-9.
    Dymek, E. E., Lefebvre, P. A., and Smith, E. F. (2004),'PF15p is the chlamydomonas homologue of the Katanin p80 subunit and is required for assembly of flagellar central microtubules', Eukaryot Cell,3 (4), 870-9.
    Fache, V., et al. (2010),'Arabidopsis kinetochore fiber-associated MAP65-4 cross-links microtubules and promotes microtubule bundle elongation', Plant Cell,22 (11),3804-15.
    Francis J., McNally and Ronald D., Vale (1993),'Identification of katanin, an ATPase that severs and disassembles stable microtubules', Cell,75, 419-29.
    Franklin, Amie E. and Cande, W. Zacheus (1999),'Nuclear Organization and Chromosome Segregation', Plant Cell,11,523-34.
    Gutierrez, R., et al. (2009),'Arabidopsis cortical microtubules position cellulose synthase delivery to the plasma membrane and interact with cellulose synthase trafficking compartments', Nat Cell Biol,11 (7),797-806.
    Hamada, T. (2007),'Microtubule-associated proteins in higher plants', J Plant Res,120 (1),79-98.
    Hartman, J. J. (1999),'Microtubule Disassembly by ATP-Dependent Oligomerization of the AAA Enzyme Katanin', Science,286 (5440), 782-85.
    Hazan, J., et al. (1999),'Spastin, a new AAA protein, is altered in the most frequent form of autosomal dominant spastic paraplegia', Nat Genet,23 (3),296-303.
    Ho, C. M., et al. (2011a),'Interaction of antiparallel microtubules in the phragmoplast is mediated by the microtubule-associated protein MAP65-3 in Arabidopsis', Plant Cell,23 (8),2909-23.
    Ho, C. M., et al. (2011b),'Augmin plays a critical role in organizing the spindle and phragmoplast microtubule arrays in Arabidopsis', Plant Cell,23 (7), 2606-18.
    James J. Hartman, et al. (1998),'Katanin, a microtubule-severing protein, is a novel AAA ATPase that targets to the centrosome using a WD40-containing subunit', Cell,93,277-87.
    Job, Didier, Valiron, Odile, and Oakley, Berl (2003),'Microtubule nucleation', CurrOpinin Cell Biol,15 (1),111-17.
    Johnson, J. L., et al. (2009),'Levels of the ubiquitin ligase substrate adaptor MEL-26 are inversely correlated with MEI-1/katanin microtubule-severing activity during both meiosis and mitosis', Dev Biol, 330 (2),349-57.
    Juranic, M., et al. (2012),'Germline-specific MATH-BTB substrate adaptor MAB1 regulates spindle length and nuclei identity in maize', Plant Cell, 24 (12),4974-91.
    Komorisono, M., et al. (2005),'Analysis of the rice mutant dwarf and gladius leaf 1. Aberrant katanin-mediated microtubule organization causes up-regulation of gibberellin biosynthetic genes independently of gibberellin signaling', Plant Physiol,138 (4),1982-93.
    Kong, Z., et al. (2010a),'The-Tubulin Complex Protein GCP4 Is Required for Organizing Functional Microtubule Arrays in Arabidopsis thaliana', Plant Cell,22 (1),191-204.
    Kong, Z., et al. (2010b),'The{gamma}-tubulin complex protein GCP4 is required for organizing functional microtubule arrays in Arabidopsis thaliana1, Plant Cell,22 (1),191-204.
    Korolev, A. V., et al. (2005),'Identification of a novel family of 70 kDa microtubule-associated proteins in Arabidopsis cells', Plant J,42 (4), 547-55.
    Kurz, T. (2002),'Cytoskeletal Regulation by the Nedd8 Ubiquitin-Like Protein Modification Pathway', Science,295 (5558),1294-98.
    Lee, Y. R., Li, Y, and Liu, B. (2007), Two Arabidopsis phragmoplast-associated kinesins play a critical role in cytokinesis during male gametogenesis', Plant Cell,19 (8),2595-605.
    Lin, D., et al. (2013),'Rho GTPase Signaling Activates Microtubule Severing to Promote Microtubule Ordering in Arabidopsis', Curr Biol,23 (4),290-7.
    Lloyd, C. and Chan, J. (2008),'The parallel lives of microtubules and cellulose microfibrils', Curr Opin Plant Biol,11 (6),641-6.
    Loughlin, R., et al. (2011),'Katanin contributes to interspecies spindle length scaling in Xenopus', Cell,147 (6),1397-407.
    Lucas, J. R., et al. (2011),'Microtubule-associated proteins MAP65-1 and MAP65-2 positively regulate axial cell growth in etiolated Arabidopsis hypocotyls', Plant Cell,23 (5),1889-903.
    Lyle, K., Kumar, P., and Wittmann, T. (2009a),'Snapshot:Microtubule Regulators I', Cell,136 (2),380,80 e1.-(2009b),'Snapshot:Microtubule regulators II', Cell,136 (3),566,66 e1.
    Martin Srayko, et al. (2000),'MEI-1/MEI-2 katanin-like microtubule severing activity is required for Caenorhabditis elegans meiosis', Genes & Development,14,1072-84.
    Mathieu, Erhardt1, et al. (2002a),'The plant Spc98p homologue colocalizes with y-tubulin at microtubule nucleation sites and is required for microtubule nucleation', Journal of Cell Science, (115),2423-31.
    Mathieu, Erhardt, Virginie, Stoppin-Mellet, and Sarah, Campagne (2002b), 'The plant Spc98p homologue colocalizes with y-tubulin at microtubule nucleation sites and is required for microtubule nucleation1, Journal of Cell Science.
    McClinton, R. S., Chandler, J. S., and Callis, J. (2001),'cDNA isolation, characterization, and protein intracellular localization of a katanin-like p60 subunit from Arabidopsis thaliana', Protoplasma,216 (3-4),181-90.
    McMichael, C. M. and Bednarek, S. Y. (2013),'Cytoskeletal and membrane dynamics during higher plant cytokinesis', New Phytol,197 (4), 1039-57.
    McNally, K., et al. (2006),'Katanin controls mitotic and meiotic spindle length1, J Cell Biol,175 (6),881-91.
    McNally, K. P., Bazirgan, O. A., and McNally, F. J. (2000), Two domains of p80 katanin regulate microtubule severing and spindle pole targeting by p60 katanin', J Cell Sci,113 (Pt 9),1623-33.
    McNally, K. P., Buster, D., and McNally, F. J. (2002),'Katanin-mediated microtubule severing can be regulated by multiple mechanisms', Cell Motil Cytoskeleton,53 (4),337-49.
    Mei, Y, et al. (2012), The Arabidopsis ARCP protein, CSI1, which is required for microtubule stability, is necessary for root and anther development', Plant Cell,24 (3),1066-80.
    Melanie Webb, et al. (2002),'Cell specification in the Arabidopsis root epidermis requires the activity of ECTOPIC ROOT HAIR3-a katanin-p60 protein', Development,129,123-31.
    Muller, S., et al. (2004), The plant microtubule-associated protein AtMAP65-3/PLE is essential for cytokinetic phragmoplast function', Curr Biol,14(5),412-7.
    Murata, T., et al. (2005),'Microtubule-dependent microtubule nucleation based on recruitment of gamma-tubulin in higher plants', Nat Cell Biol,7 (10), 961-8.
    Nakamura, M., Ehrhardt, D. W., and Hashimoto, T. (2010),'Microtubule and katanin-dependent dynamics of microtubule nucleation complexes in the acentrosomal Arabidopsis cortical array1, Nat Cell Biol,12 (11), 1064-70.
    O'Donnell, L, et al. (2012),'An essential role for katanin p80 and microtubule severing in male gamete production', PLoS Genet,8 (5), e1002698.
    Panteris, E. and Adamakis, I. D. (2012),'Aberrant microtubule organization in dividing root cells of p60-katanin mutants', Plant Signal Behav,7 (1), 16-8.
    Panteris, E., et al. (2011),'A role for katanin in plant cell division:microtubule organization in dividing root cells of fra2 and lue1Arabidopsis thaliana mutants', Cytoskeleton,68 (7),401-13.
    Paradez, A., Wright, A., and Ehrhardt, D. W. (2006),'Microtubule cortical array organization and plant cell morphogenesis', Curr Opin Plant Biol,9 (6), 571-8.
    Paredez, A. R., Somerville, C. R., and Ehrhardt, D. W. (2006),'Visualization of cellulose synthase demonstrates functional association with microtubules', Science,312 (5779),1491-5.
    Pellettieri, Jason, et al. (2003),'Coordinate Activation of Maternal Protein Degradation during the Egg-to-Embryo Transition in C. elegans', Dev Cell,5(3),451-62.
    Perrin, Robyn M., et al. (2007),'WVD2 is a novel microtubule-associated protein in Arabidopsis thaliana', The Plant Journal,49 (6),961-71.
    Qiang, L, et al. (2006), Tau protects microtubules in the axon from severing by katanin', J Neurosci,26 (12),3120-9.
    Rebecca, Heald and Eva, Nogales (2002),'Microtubule dynamics', J Cell Sci, (115),3-4.
    Roll-Mecak, A. and Vale, R. D. (2008),'Structural basis of microtubule severing by the hereditary spastic paraplegia protein spastin', Nature,451 (7176), 363-7.
    Roll-Mecak, A. and McNally, F. J. (2010),'Microtubule-severing enzymes', Curr Opin Cell Biol,22 (1),96-103.
    Sedbrook, J. C. and Kaloriti, D. (2008),'Microtubules, MAPs and plant directional cell expansion', Trends Plant Sci,13 (6),303-10.
    Sedbrook, J. C., et al. (2004), The Arabidopsis sku6/spiral1 gene encodes a plus end-localized microtubule-interacting protein involved in directional cell expansion', Plant Cell,16 (6),1506-20.
    Seltzer, V, et al. (2007),'Arabidopsis GCP2 and GCP3 are part of a soluble gamma-tubulin complex and have nuclear envelope targeting domains', Plant J,52(2),322-31.
    Sharma, N., et al. (2007),'Katanin regulates dynamics of microtubules and biogenesis of motile cilia', J Cell Biol, 178 (6),1065-79.
    Sharp, D. J. and Ross, J. L. (2012),'Microtubule-severing enzymes at the cutting edge', J Cell Sci, 125 (Pt 11),2561-9.
    Shimamura, M., et al. (2004),'Gamma-tubulin in basal land plants: characterization, localization, and implication in the evolution of acentriolar microtubule organizing centers', Plant Cell,16 (1),45-59.
    Soga, Kouichi, et al. (2010),'1-Aminocyclopropane-1-carboxylic acid (ACC)-induced reorientation of cortical microtubules is accompanied by a transient increase in the transcript levels of y-tubulin complex and katanin genes in azuki bean epicotyls', J Plant Physiol,167 (14), 1165-71.
    Stitzel, M. L, Pellettieri, J., and Seydoux, G. (2006), The C. elegans DYRK Kinase MBK-2 Marks Oocyte Proteins for Degradation in Response to Meiotic Maturation', Curr Biol,16 (1),56-62.
    Stoppin-Mellet, V., Gaillard, J., and Vantard, M. (2002),'Functional evidence for in vitro microtubule severing by the plant katanin homologue', Biochem J,365 (Pt2),337-42.- (2006),'Katanin's severing activity favors bundling of cortical microtubules in plants', Plant J,46 (6),1009-17.
    Stoppin-Mellet, V., et al. (2007),'Arabidopsis katanin binds microtubules using a multimeric microtubule-binding domain', Plant Physiol Biochem,45 (12),867-77.
    Stragene (2004),'BacterioMatch? II Two-Hybrid System Library Construction Kit INSTRUCTION MANUAL'.
    Sudo, H. and Maru, Y. (2007),'LAPSER1 is a putative cytokinetic tumor suppressor that shows the same centrosome and midbody subcellular localization pattern as p80 katanin', FASEB J,21 (9),2086-100.-(2008),'LAPSER1/LZTS2:a pluripotent tumor suppressor linked to the inhibition of katanin-mediated microtubule severing', Hum Mol Genet, 17 (16),2524-40.
    Sudo, H. and Baas, P. W. (2010),'Acetylation of microtubules influences their sensitivity to severing by katanin in neurons and fibroblasts', J Neurosci, 30(21),7215-26.
    Sugimoto, K., Williamson, R. E., and Wasteneys, G. O. (2000),'New techniques enable comparative analysis of microtubule orientation, wall texture, and growth rate in intact roots of Arabidopsis', Plant Physiol, 124(4),1493-506.
    Tanaka, Ichiro (1993),'Development of Male Gametes in Flowering Plants', J Plant Research, (106),55-63.
    Uyttewaal, M., et al. (2012),'Mechanical stress acts via katanin to amplify differences in growth rate between adjacent cells in Arabidopsis', Cell, 149 (2),439-51.
    Wang, X., et al. (2012),'Arabidopsis microtubule destabilizing protein40 is involved in brassinosteroid regulation of hypocotyl elongation', Plant Cell,24(10),4012-25.
    Wang, X., et al. (2007),'Arabidopsis MICROTUBULE-ASSOCIATED PROTEIN 18 functions in directional cell growth by destabilizing cortical microtubules', Plant Cell, 19 (3),877-89.
    Whitehead, E., Heald, R., and Wilbur, J. D. (2013),'N-terminal phosphorylation of p60 katanin directly regulates microtubule severing', J Mol Biol, 425 (2),214-21.
    Whittington, A. T., et al. (2001),'MOR1 is essential for organizing cortical microtubules in plants', Nature, 411 (6837),610-3.
    Wiese, C. and Zheng, Y. (2006),'Microtubule nucleation:gamma-tubulin and beyond', J Cell Sci, 119 (Pt 20),4143-53.
    Wightman, R. and Turner, S. (2010),'Trafficking of the plant cellulose synthase complex', Plant Physiol, 153 (2),427-32.
    Yao-Guang Liu, et al. (1995),'Efiicient isolation and mapping of Arabidopsis thaliana T-DNA insert junctions by thermal assymmetric interlaced PCR', Plant J, 8(3),457-7.
    Young, Robin E. and Bisgrove, Sherryl R. (2011),'Microtubule Plus End-Tracking Proteins and Their Activities in Plants', Plant Cytoskeleton, 95-117.
    Yu, W., et al. (2005),'Regulation of microtubule severing by katanin subunits during neuronal development', J Neurosci,25 (23),5573-83.
    Zeng, C. J., Lee, Y. R., and Liu, B. (2009), The WD40 repeat protein NEDD1 functions in microtubule organization during cell division in Arabidopsis thaliana', Plant Cell,21 (4),1129-40.
    Zhang, D., et al. (2007), Three microtubule severing enzymes contribute to the "Pacman-flux" machinery that moves chromosomes', J Cell Biol,177 (2), 231-42.
    Zhang, D., et al. (2011),'Drosophila katanin is a microtubule depolymerase that regulates cortical-microtubule plus-end interactions and cell migration', Nat Cell Biol,13 (4),361-70.
    Zhang, D. H., Wadsworth, P., and Hepler, P. K. (1992),'Modulation of anaphase spindle microtubule structure in stamen hair cells of Tradescantia by calcium and related agents', J Cell Sci,102,79-89.
    胡晶(2010),‘水稻BphiOO8a基因介导水稻抗褐飞虱机理研究’,武汉大学博士学位论文
    胡骏(2009),‘红莲型水稻细胞质雄性不育育性恢复机理的研究’,武汉尢学博士学位论文
    李阳(2010),‘籼稻高效遗传转化技术和愈伤组织的质量生化鉴定及表达分析’,武汉大学博士学位论文
    刘盖(2011),‘红莲型细胞质雄性不育水稻线粒体蛋白质组学分析’,武汉才学博士学位论文
    朱蕾(2008),‘拟南芥微管结合蛋白MAP18对微管动态调控机理的研究’,中国农业大学博士学位论文

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