【《微管的组成及动态性研究文献综述》2400字】

微管的组成及动态性研究文献综述1.1微管的组成及动态性微管是一种坚硬的非共价聚合物，它在细胞内形成了一个坚硬的网络，起到细胞骨架的作用。微管是由13根微管原丝所组成的空心管状结构，外直径为24nm，内直径12nmADDINEN.CITE<EndNote><Cite><Author>Hohmann</Author><Year>2019</Year><RecNum>125</RecNum><DisplayText>(HohmannandDehghani,2019)</DisplayText><record><rec-number>125</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1560931503">125</key><keyapp="ENWeb"db-id="">0</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Hohmann,T.</author><author>Dehghani,F.</author></authors></contributors><auth-address>InstituteofAnatomyandCellBiology,MartinLutherUniversityHalle-Wittenberg,GrosseSteinstrasse52,06108Halle(Saale),Germany.tim.hohmann@medizin.uni-halle.de. InstituteofAnatomyandCellBiology,MartinLutherUniversityHalle-Wittenberg,GrosseSteinstrasse52,06108Halle(Saale),Germany.faramarz.dehghani@medizin.uni-halle.de.</auth-address><titles><title>TheCytoskeleton-AComplexInteractingMeshwork</title><secondary-title>Cells</secondary-title></titles><periodical><full-title>Cells</full-title></periodical><volume>8</volume><number>4</number><keywords><keyword>actin</keyword><keyword>glioma</keyword><keyword>intermediatefilaments</keyword><keyword>microtubules</keyword><keyword>migration</keyword><keyword>motility</keyword><keyword>signaling</keyword></keywords><dates><year>2019</year><pub-dates><date>Apr18</date></pub-dates></dates><isbn>2073-4409(Print) 2073-4409(Linking)</isbn><accession-num>31003495</accession-num><urls><related-urls><url>/pubmed/31003495</url></related-urls></urls><custom2>PMC6523135</custom2><electronic-resource-num>10.3390/cells8040362</electronic-resource-num></record></Cite></EndNote>(HohmannandDehghani,2019)。每根原丝由α微管蛋白和β微管蛋白首尾相连形成的异源二聚体所构成。在正常的细胞内，微管蛋白主要是以αβ-微管蛋白二聚体的形式游离在细胞质中ADDINEN.CITE<EndNote><Cite><Author>Brouhard</Author><Year>2015</Year><RecNum>1236</RecNum><DisplayText>(Brouhard,2015)</DisplayText><record><rec-number>1236</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1615810866">1236</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Brouhard,G.J.</author></authors></contributors><auth-address>DepartmentofBiology,McGillUniversity,Montreal,QCH3A1B1,Canadagary.brouhard@mcgill.ca.</auth-address><titles><title>Dynamicinstability30yearslater:complexitiesinmicrotubulegrowthandcatastrophe</title><secondary-title>MolBiolCell</secondary-title></titles><periodical><full-title>MolBiolCell</full-title></periodical><pages>1207-10</pages><volume>26</volume><number>7</number><edition>2015/04/01</edition><keywords><keyword>Animals</keyword><keyword>GuanosineTriphosphate/metabolism</keyword><keyword>Microtubules/*metabolism</keyword><keyword>Tubulin/*metabolism</keyword></keywords><dates><year>2015</year><pub-dates><date>Apr1</date></pub-dates></dates><isbn>1939-4586(Electronic) 1059-1524(Linking)</isbn><accession-num>25823928</accession-num><urls><related-urls><url>/pubmed/25823928</url></related-urls></urls><custom2>PMC4454169</custom2><electronic-resource-num>10.1091/mbc.E13-10-0594</electronic-resource-num></record></Cite></EndNote>(Brouhard,2015)。α微管蛋白和β微管蛋白都含有GTP结合位点，但是稳定性不同。α微管蛋白的GTP结合位点（N位点）由于在蛋白空间构象原因，该位点的GTP不会被水解。β微管蛋白的GTP结合位点（E位点），可以参与αβ-微管蛋白二聚体的微管组装，在这个过程E位点的GTP会水解成GDP。在微管去组装过程中，β微管蛋白的E位点上的GDP会被GTP替代ADDINEN.CITE<EndNote><Cite><Author>Roostalu</Author><Year>2017</Year><RecNum>1239</RecNum><DisplayText>(RoostaluandSurrey,2017)</DisplayText><record><rec-number>1239</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1615811069">1239</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Roostalu,J.</author><author>Surrey,T.</author></authors></contributors><auth-address>TheFrancisCrickInstitute,1MidlandRoad,LondonNW11AT,UK.</auth-address><titles><title>Microtubulenucleation:beyondthetemplate</title><secondary-title>NatRevMolCellBiol</secondary-title></titles><periodical><full-title>NatRevMolCellBiol</full-title></periodical><pages>702-710</pages><volume>18</volume><number>11</number><edition>2017/08/24</edition><keywords><keyword>Animals</keyword><keyword>Humans</keyword><keyword>Microtubules/*metabolism</keyword><keyword>Tubulin/*metabolism</keyword></keywords><dates><year>2017</year><pub-dates><date>Nov</date></pub-dates></dates><isbn>1471-0080(Electronic) 1471-0072(Linking)</isbn><accession-num>28831203</accession-num><urls><related-urls><url>/pubmed/28831203</url></related-urls></urls><electronic-resource-num>10.1038/nrm.2017.75</electronic-resource-num></record></Cite></EndNote>(RoostaluandSurrey,2017)。图1.15微管的组成和动态不稳定性α-微管蛋白和β微管蛋白组成异源二聚体，二聚体在收尾相连形成一个空心管状结构。在微管的正端形成GFP-cap结构，微管变长保护微管不回缩。如果微管正端GFP-cap消失，则会发生微管回缩ADDINEN.CITE<EndNote><Cite><Author>Hohmann</Author><Year>2019</Year><RecNum>125</RecNum><DisplayText>(HohmannandDehghani,2019)</DisplayText><record><rec-number>125</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1560931503">125</key><keyapp="ENWeb"db-id="">0</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Hohmann,T.</author><author>Dehghani,F.</author></authors></contributors><auth-address>InstituteofAnatomyandCellBiology,MartinLutherUniversityHalle-Wittenberg,GrosseSteinstrasse52,06108Halle(Saale),Germany.tim.hohmann@medizin.uni-halle.de. InstituteofAnatomyandCellBiology,MartinLutherUniversityHalle-Wittenberg,GrosseSteinstrasse52,06108Halle(Saale),Germany.faramarz.dehghani@medizin.uni-halle.de.</auth-address><titles><title>TheCytoskeleton-AComplexInteractingMeshwork</title><secondary-title>Cells</secondary-title></titles><periodical><full-title>Cells</full-title></periodical><volume>8</volume><number>4</number><keywords><keyword>actin</keyword><keyword>glioma</keyword><keyword>intermediatefilaments</keyword><keyword>microtubules</keyword><keyword>migration</keyword><keyword>motility</keyword><keyword>signaling</keyword></keywords><dates><year>2019</year><pub-dates><date>Apr18</date></pub-dates></dates><isbn>2073-4409(Print) 2073-4409(Linking)</isbn><accession-num>31003495</accession-num><urls><related-urls><url>/pubmed/31003495</url></related-urls></urls><custom2>PMC6523135</custom2><electronic-resource-num>10.3390/cells8040362</electronic-resource-num></record></Cite></EndNote>(HohmannandDehghani,2019)。在电子显微镜下，可以观察到微管的13根原丝是同向平行排布的，相邻的原丝之间存在大约1nm的交错。每根原丝上的αβ-微管蛋白二聚体都是规律排列，因此每根原丝的头尾都是不同的微管蛋白，从而使微管的两端呈现不同的极性ADDINEN.CITE<EndNote><Cite><Author>Howard</Author><Year>2007</Year><RecNum>1242</RecNum><DisplayText>(HowardandHyman,2007)</DisplayText><record><rec-number>1242</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1615811182">1242</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Howard,J.</author><author>Hyman,A.A.</author></authors></contributors><auth-address>MPI-CBGPfotenhauerstr,10801307,Dresden,Germany.howard@mpi-cbg.de</auth-address><titles><title>Microtubulepolymerasesanddepolymerases</title><secondary-title>CurrOpinCellBiol</secondary-title></titles><periodical><full-title>CurrOpinCellBiol</full-title></periodical><pages>31-5</pages><volume>19</volume><number>1</number><edition>2006/12/23</edition><keywords><keyword>Animals</keyword><keyword>Kinesin/*physiology</keyword><keyword>Kinetochores/physiology</keyword><keyword>Microtubule-AssociatedProteins/*physiology</keyword><keyword>Microtubules/*physiology</keyword><keyword>SaccharomycescerevisiaeProteins/physiology</keyword><keyword>XenopusProteins/physiology</keyword></keywords><dates><year>2007</year><pub-dates><date>Feb</date></pub-dates></dates><isbn>0955-0674(Print) 0955-0674(Linking)</isbn><accession-num>17184986</accession-num><urls><related-urls><url>/pubmed/17184986</url></related-urls></urls><electronic-resource-num>10.1016/j.ceb.2006.12.009</electronic-resource-num></record></Cite></EndNote>(HowardandHyman,2007)。我们将微管组装较快的一端称为微管正端（plusend），另一端称为负端（minusend）。同时，由于α和β-微管蛋白的C端含有酸性氨基酸，因此，微管表面带有较强负电。微管蛋白在动物进化过程中相对保守，酵母和人的微管蛋白在氨基酸序列上的重复度达到75%ADDINEN.CITEADDINEN.CITE.DATA(Tietal.,2018)。α-微管蛋白含有450个氨基酸，β-微管蛋白含有455个氨基酸，二者的氨基酸序列相似性只有45%，但两者的三维结构非常相似，每个单体都由三个长度相似的结构域和次级结构域构成：N端结构域、中间域和C端结构域。微管在体内是呈现高度动态的，微管处于不断的伸长和回缩之间，这种聚合、解聚合的状态称为微管的动态不稳定性（dynamicinstability）ADDINEN.CITEADDINEN.CITE.DATA(Bowne-Andersonetal.,2013;HowardandHyman,2007)。微管这一特性首次在1972年被报道，Weisenberg在体外实验中，加入在含有钙螯合剂和GTP的缓冲液中微管蛋白可逆自组装，自此开始微管的动力学特征被描述。微管的动力学主要集中在体外实验中ADDINEN.CITE<EndNote><Cite><Author>Olmsted</Author><Year>1974</Year><RecNum>1251</RecNum><DisplayText>(Olmstedetal.,1974)</DisplayText><record><rec-number>1251</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1615812240">1251</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Olmsted,J.B.</author><author>Marcum,J.M.</author><author>Johnson,K.A.</author><author>Allen,C.</author><author>Borisy,G.G.</author></authors></contributors><titles><title>Microtubleassembly:somepossibleregulatorymechanisms</title><secondary-title>JSupramolStruct</secondary-title></titles><periodical><full-title>JSupramolStruct</full-title></periodical><pages>429-50</pages><volume>2</volume><number>2-4</number><edition>1974/01/01</edition><keywords><keyword>Animals</keyword><keyword>Brain/*physiology</keyword><keyword>Calcium/pharmacology</keyword><keyword>Chlamydomonas</keyword><keyword>Chromatography,Affinity</keyword><keyword>Colchicine/pharmacology</keyword><keyword>Electrophoresis,PolyacrylamideGel</keyword><keyword>GuanosineTriphosphate/pharmacology</keyword><keyword>Hydrogen-IonConcentration</keyword><keyword>Kinetics</keyword><keyword>MacromolecularSubstances</keyword><keyword>Magnesium/pharmacology</keyword><keyword>Microscopy,Electron</keyword><keyword>Microtubules/drugeffects/*physiology/ultrastructure</keyword><keyword>NerveTissueProteins/*metabolism</keyword><keyword>OsmolarConcentration</keyword><keyword>Swine</keyword><keyword>Temperature</keyword><keyword>Thermodynamics</keyword><keyword>Ultracentrifugation</keyword><keyword>Viscosity</keyword></keywords><dates><year>1974</year></dates><isbn>0091-7419(Print) 0091-7419(Linking)</isbn><accession-num>4612256</accession-num><urls><related-urls><url>/pubmed/4612256</url></related-urls></urls><electronic-resource-num>10.1002/jss.400020230</electronic-resource-num></record></Cite></EndNote>(Olmstedetal.,1974)。在体外试管中，当携带GFP的αβ-微管蛋白二聚体浓度较高时，微管末端的延伸速度较快。β微管蛋白上的GTP被水解为GDP，这个过程中会产生一定的延迟效应，从而在微管的末端形成了GTP状态的微管蛋白区域，称为GTPcap。当GTPcap消失后，微管就会发生崩塌ADDINEN.CITE<EndNote><Cite><Author>Hill</Author><Year>1984</Year><RecNum>1246</RecNum><DisplayText>(HillandChen,1984)</DisplayText><record><rec-number>1246</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1615812019">1246</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Hill,T.L.</author><author>Chen,Y.</author></authors></contributors><titles><title>PhasechangesattheendofamicrotubulewithaGTPcap</title><secondary-title>ProcNatlAcadSciUSA</secondary-title></titles><periodical><full-title>ProcNatlAcadSciUSA</full-title></periodical><pages>5772-6</pages><volume>81</volume><number>18</number><edition>1984/09/01</edition><keywords><keyword>Animals</keyword><keyword>GuanosineTriphosphate/*metabolism</keyword><keyword>Kinetics</keyword><keyword>Mathematics</keyword><keyword>Microtubules/*ultrastructure</keyword><keyword>*Models,Biological</keyword><keyword>ProteinBinding</keyword><keyword>Tubulin/*metabolism</keyword></keywords><dates><year>1984</year><pub-dates><date>Sep</date></pub-dates></dates><isbn>0027-8424(Print) 0027-8424(Linking)</isbn><accession-num>6592585</accession-num><urls><related-urls><url>/pubmed/6592585</url></related-urls></urls><custom2>PMC391793</custom2><electronic-resource-num>10.1073/pnas.81.18.5772</electronic-resource-num></record></Cite></EndNote>(HillandChen,1984)。1.2非中心体微管在分化的细胞中，微管采用特定的空间排列，执行不同的功能。早期电子显微镜显示了不同的亚细胞部位含有的微管聚集中心，这个部位被称为微管组织中心（microtubuleorganizingcenter,MTOC）ADDINEN.CITE<EndNote><Cite><Author>Sanchez</Author><Year>2017</Year><RecNum>1223</RecNum><DisplayText>(SanchezandFeldman,2017b)</DisplayText><record><rec-number>1223</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1615809745">1223</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Sanchez,A.D.</author><author>Feldman,J.L.</author></authors></contributors><auth-address>DepartmentofBiology,StanfordUniversity,371SerraMall,Stanford,CA94305,USA. DepartmentofBiology,StanfordUniversity,371SerraMall,Stanford,CA94305,USA.Electronicaddress:feldmanj@.</auth-address><titles><title>Microtubule-organizingcenters:fromthecentrosometonon-centrosomalsites</title><secondary-title>CurrOpinCellBiol</secondary-title></titles><periodical><full-title>CurrOpinCellBiol</full-title></periodical><pages>93-101</pages><volume>44</volume><edition>2016/09/27</edition><keywords><keyword>Animals</keyword><keyword>CellDifferentiation</keyword><keyword>Centrosome/metabolism</keyword><keyword>Cytoskeleton</keyword><keyword>Humans</keyword><keyword>Microtubule-OrganizingCenter/*chemistry/*metabolism</keyword><keyword>Microtubules/metabolism</keyword><keyword>Mitosis</keyword></keywords><dates><year>2017</year><pub-dates><date>Feb</date></pub-dates></dates><isbn>1879-0410(Electronic) 0955-0674(Linking)</isbn><accession-num>27666167</accession-num><urls><related-urls><url>/pubmed/27666167</url></related-urls></urls><custom2>PMC5362366</custom2><electronic-resource-num>10.1016/j.ceb.2016.09.003</electronic-resource-num></record></Cite></EndNote>(SanchezandFeldman,2017b)。自从50年前MTOC被发现以来，人们的研究主要集中在中心体上。除了中心体外，体内还存在很多其它的结构，也能起到微管组织中心的作用，例如纤毛和鞭毛的基部，高尔基体前哨的反面囊膜区域也具有微管组装的能力等。中心体是由两个直径为0.2μm，长度0.4μm的桶状中心粒所组成，两个中心粒呈现L直角型排列ADDINEN.CITE<EndNote><Cite><Author>Doxsey</Author><Year>2005</Year><RecNum>1230</RecNum><DisplayText>(Doxseyetal.,2005)</DisplayText><record><rec-number>1230</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1615810121">1230</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Doxsey,S.</author><author>Zimmerman,W.</author><author>Mikule,K.</author></authors></contributors><auth-address>UnivMassachusetts,SchMed,ProgramMolMed,Worcester,MA01605USA HarvardUniv,DeptMol&CellularBiol,Cambridge,MA02138USA</auth-address><titles><title>Chromosomesegregationandaneuploidyseries:Centrosomecontrolofthecell</title><secondary-title>TrendsinCellBiology</secondary-title><alt-title>TrendsCellBiol</alt-title></titles><periodical><full-title>TrendsinCellBiology</full-title><abbr-1>TrendsCellBiol</abbr-1></periodical><alt-periodical><full-title>TrendsinCellBiology</full-title><abbr-1>TrendsCellBiol</abbr-1></alt-periodical><pages>303-311</pages><volume>15</volume><number>6</number><keywords><keyword>cycleprogression</keyword><keyword>s-phase</keyword><keyword>gamma-tubulin</keyword><keyword>scaffoldproteins</keyword><keyword>vertebratecells</keyword><keyword>DNA-replication</keyword><keyword>polokinase</keyword><keyword>cytokinesis</keyword><keyword>duplication</keyword><keyword>localization</keyword></keywords><dates><year>2005</year><pub-dates><date>Jun</date></pub-dates></dates><isbn>0962-8924</isbn><accession-num>WOS:000230053400004</accession-num><urls><related-urls><url><GotoISI>://WOS:000230053400004</url></related-urls></urls><electronic-resource-num>10.1016/j.tcb.2005.04.008</electronic-resource-num><language>English</language></record></Cite></EndNote>(Doxseyetal.,2005)。每个中心粒包含9组三联体微管。电镜下观察，在中心粒的周围还包含了其它电子致密物质(PCM)，这些蛋白共同完成了招募和微管成核作用，微管从这个区域延伸出去ADDINEN.CITE<EndNote><Cite><Author>Ito</Author><Year>2018</Year><RecNum>1233</RecNum><DisplayText>(ItoandBettencourt-Dias,2018)</DisplayText><record><rec-number>1233</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1615810517">1233</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Ito,D.</author><author>Bettencourt-Dias,M.</author></authors></contributors><auth-address>InstitutoGulbenkiandeCiencia,RuadaQuintaGrande6,2780-156Oeiras,Portugal.dito@igc.gulbenkian.pt. InstitutoGulbenkiandeCiencia,RuadaQuintaGrande6,2780-156Oeiras,Portugal.mdias@igc.gulbenkian.pt.</auth-address><titles><title>CentrosomeRemodellinginEvolution</title><secondary-title>Cells</secondary-title></titles><periodical><full-title>Cells</full-title></periodical><volume>7</volume><number>7</number><edition>2018/07/11</edition><keywords><keyword>Pcm</keyword><keyword>Spb</keyword><keyword>centriole</keyword><keyword>centrosome</keyword><keyword>evolution</keyword><keyword>spindlepolebody</keyword></keywords><dates><year>2018</year><pub-dates><date>Jul6</date></pub-dates></dates><isbn>2073-4409(Print) 2073-4409(Linking)</isbn><accession-num>29986477</accession-num><urls><related-urls><url>/pubmed/29986477</url></related-urls></urls><custom2>PMC6070874</custom2><electronic-resource-num>10.3390/cells7070071</electronic-resource-num></record></Cite></EndNote>(ItoandBettencourt-Dias,2018)。从中心体延伸出去的微管多数为正端朝外的方式，并且具有高度的动态性ADDINEN.CITE<EndNote><Cite><Author>Sanchez</Author><Year>2017</Year><RecNum>480</RecNum><DisplayText>(SanchezandFeldman,2017a)</DisplayText><record><rec-number>480</rec-number><foreign-keys><keyapp="EN"db-id="tsvwxvpfktsw99e9wrax99f3pwwa5rdvsasw"timestamp="1614079690">480</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Sanchez,ArianaD.</author><author>Feldman,JessicaL.</author></authors></contributors><titles><title>Microtubule-organizingcenters:fromthecentrosometonon-centrosomalsites</title><secondary-title>CurrentOpinioninCellBiology</secondary-title></titles><periodical><full-title>CurrentOpinioninCellBiology</full-title><abbr-1>Curr.Opin.CellBiol.</abbr-1></periodical><pages>93-101</pages><volume>44</volume><dates><year>2017</year><pub-dates><date>Feb</date></pub-dates></dates><isbn>0955-0674</isbn><accession-num>WOS:000400022400014</accession-num><urls><related-urls><url><GotoISI>://WOS:000400022400014</url></related-urls></urls><electronic-resource-num>10.1016/j.ceb.2016.09.003</electronic-resource-num></record></Cite></EndNote>(SanchezandFeldman,2017a)。然而在动物体内，还存在很多微管负端没有锚定到中心体上，它们不形成星状的骨架网络，这类微管称为非中心体微管ADDINEN.CITE<EndNote><Cite><Author>Sanchez</Author><Year>2017</Year><RecNum>1223</RecNum><DisplayText>(SanchezandFeldman,2017b)</DisplayText><record><rec-number>1223</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1615809745">1223</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Sanchez,A.D.</author><author>Feldman,J.L.</author></authors></contributors><auth-address>DepartmentofBiology,StanfordUniversity,371SerraMall,Stanford,CA94305,USA. DepartmentofBiology,StanfordUniversity,371SerraMall,Stanford,CA94305,USA.Electronicaddress:feldmanj@.</auth-address><titles><title>Microtubule-organizingcenters:fromthecentrosometonon-centrosomalsites</title><secondary-title>CurrOpinCellBiol</secondary-title></titles><periodical><full-title>CurrOpinCellBiol</full-title></periodical><pages>93-101</pages><volume>44</volume><edition>2016/09/27</edition><keywords><keyword>Animals</keyword><keyword>CellDifferentiation</keyword><keyword>Centrosome/metabolism</keyword><keyword>Cytoskeleton</keyword><keyword>Humans</keyword><keyword>Microtubule-OrganizingCenter/*chemistry/*metabolism</keyword><keyword>Microtubules/metabolism</keyword><keyword>Mitosis</keyword></keywords><dates><year>2017</year><pub-dates><date>Feb</date></pub-dates></dates><isbn>1879-0410(Electronic) 0955-0674(Linking)</isbn><accession-num>27666167</accession-num><urls><related-urls><url>/pubmed/27666167</url></related-urls></urls><custom2>PMC5362366</custom2><electronic-resource-num>10.1016/j.ceb.2016.09.003</electronic-resource-num></record></Cite></EndNote>(SanchezandFeldman,2017b)。例如神经元中的微管就呈现不同的排列方式，在轴突中，微管正端朝向顶端，在树突中，微管呈极性混合排列，而在果蝇和线虫的树突中，神经元中微管负端朝向顶端ADDINEN.CITEADDINEN.CITE.DATA(Craftetal.,2015;Maniaretal.,2012;Zhengetal.,2008)。在果蝇的IV型树突类神经元中，细胞器高尔基体前哨结构具有ncMTOCs（non-centrosomalmicrotubule-organizingcenters）的功能，在实验中，不同的高尔基体前哨结构可以调节部分树突中微管的成核，但是高尔基体复合物不直接参与成核位点ADDINEN.CITEADDINEN.CITE.DATA(Nguyenetal.,2014)。在高等植物细胞内，由于没有中心体结构及类似结构，但是细胞内微管在局部却呈现高度规则排列，这种微管排列的方式是植物正常发育所必需的ADDINEN.CITEADDINEN.CITE.DATA(Murataetal.,2005)。非中心体微管负端的稳定通常有几种方式，例如基于高尔基体或细胞质中的微管负端稳定蛋白（例如Camsap/Nezha/Patronin家族蛋白）的机制。在哺乳细胞中，微管负端稳定蛋白主要分为CAMSAP1、CAMSAP2和NezhaADDINEN.CITEADDINEN.CITE.DATA(Bainesetal.,2009)。在果蝇体内，CAMSAP3的同源蛋白ssp4/Patronin蛋白可以诱导在细胞分裂间期有丝分裂短纺锤体的微管的分离ADDINEN.CITEADDINEN.CITE.DATA(Goshimaetal.,2007)。Patronin蛋白可以抑制kinesin13对微管的解聚作用，从而达到稳定微管末端。因此，CAMSAP/Patronin被认为是稳定微管模板所必须的蛋白ADDINEN.CITE<EndNote><Cite><Author>Stone</Author><Year>2008</Year><RecNum>1072</RecNum><DisplayText>(Stoneetal.,2008)</DisplayText><record><rec-number>1072</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1615085126">1072</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Stone,M.C.</author><author>Roegiers,F.</author><author>Rolls,M.M.</author></authors></contributors><auth-address>DepartmentofBiochemistryandMolecularBiology,ThePennsylvaniaStateUniversity,UniversityPark,PA16802,USA.</auth-address><titles><title>MicrotubuleshaveoppositeorientationinaxonsanddendritesofDrosophilaneurons</title><secondary-title>MolBiolCell</secondary-title></titles><periodical><full-title>MolBiolCell</full-title></periodical><pages>4122-9</pages><volume>19</volume><number>10</number><edition>2008/08/01</edition><keywords><keyword>Animals</keyword><keyword>Animals,GeneticallyModified</keyword><keyword>Axons/*metabolism</keyword><keyword>Cytoskeleton/metabolism</keyword><keyword>Dendrites/*metabolism</keyword><keyword>Drosophilamelanogaster/*metabolism</keyword><keyword>Endosomes/metabolism</keyword><keyword>GreenFluorescentProteins/metabolism</keyword><keyword>Heterozygote</keyword><keyword>LuminescentProteins/metabolism</keyword><keyword>Microtubules/*metabolism</keyword><keyword>Models,Biological</keyword><keyword>Neurons/*metabolism</keyword><keyword>Transgenes</keyword></keywords><dates><year>2008</year><pub-dates><date>Oct</date></pub-dates></dates><isbn>1939-4586(Electronic) 1059-1524(Linking)</isbn><accession-num>18667536</accession-num><urls><related-urls><url>/pubmed/18667536</url></related-urls></urls><custom2>PMC2555934</custom2><electronic-resource-num>10.1091/mbc.E07-10-1079</electronic-resource-num></record></Cite></EndNote>(Stoneetal.,2008)。在哺乳动物上皮细胞中分析，CAMSAP2和CAMSAP3位于非中心体微管的负端，并在组织中起到相互协调的作用ADDINEN.CITE<EndNote><Cite><Author>Tanaka</Author><Year>2012</Year><RecNum>1203</RecNum><DisplayText>(Tanakaetal.,2012)</DisplayText><record><rec-number>1203</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1615808253">1203</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Tanaka,N.</author><author>Meng,W.X.</author><author>Nagae,S.</author><author>Takeichi,M.</author></authors></contributors><auth-address>RIKENCtrDevBiol,Kobe,Hyogo6500047,Japan KyotoUniv,GradSchBiostudies,Kyoto6068502,Japan ChineseAcadSci,InstGenet&DevBiol,StateKeyLabMolDevBiol,Beijing100101,PeoplesRChina</auth-address><titles><title>Nezha/CAMSAP3andCAMSAP2cooperateinepithelial-specificorganizationofnoncentrosomalmicrotubules</title><secondary-title>ProceedingsoftheNationalAcademyofSciencesoftheUnitedStatesofAmerica</secondary-title><alt-title>PNatlAcadSciUSA</alt-title></titles><periodical><full-title>ProceedingsoftheNationalAcademyofSciencesoftheUnitedStatesofAmerica</full-title></periodical><pages>20029-20034</pages><volume>109</volume><number>49</number><keywords><keyword>gamma-tubulin</keyword><keyword>eb1</keyword><keyword>cells</keyword><keyword>nucleation</keyword><keyword>tracking</keyword><keyword>network</keyword><keyword>ends</keyword></keywords><dates><year>2012</year><pub-dates><date>Dec4</date></pub-dates></dates><isbn>0027-8424</isbn><accession-num>WOS:000312347200040</accession-num><urls><related-urls><url><GotoISI>://WOS:000312347200040</url></related-urls></urls><electronic-resource-num>10.1073/pnas.1218017109</electronic-resource-num><language>English</language></record></Cite></EndNote>(Tanakaetal.,2012)。近年的研究表明，非中心体微管在多种细胞的形态发生、极性建立和胞内物质运输等过程中发挥了重要的作用。但是到目前为止，我们对非中心体微管的理解还不是很深入。因此，在神经细胞中或者免疫细胞探索非中心体微管的结构及其细胞生物学具有重要意义，近年来已经成为微管细胞骨架生物学的研究前沿。参考文献Adly,N.,Alhashem,A.,Ammari,A.,andAlkuraya,F.S.(2014).CiliarygenesTBC1D32/C6orf170andSCLT1aremu