文稿教程成果chapter

Chapter 细胞骨§6.1细胞骨架的概Thecytoskeletonisthecomplexnetwork（复杂网络）ofproteinfilaments（蛋白纤维）thatextendthroughouttheeukaryoticcell（真核细Ahighlydynamicstructure（动态结构）thatreorganizescontinuouslyasthecellchangesshape,divides,andrespondstoi interactwithenvironmentdirectmovementsintermediatefilament（中等纤维nuclearmatrix（核基质nuclearlamina（核纤层肌动肌动蛋白微中间纤two-strandedhelicallong,hollowcylindersattachedtoamicrotubuleorganizingcenter(MTOC)Intermediateforma中等纤肌动蛋白丝质网络末中间纤§6.2.1≥5%oftotalActinmoleculeα-invarioustypesofmusclecellsβ-andγ-innonmusclecells

肌动蛋白负正动态结

Ca2+,lowNa+,ATPandMg2+andhighNa+,

G-肌动蛋白的结负负正正解聚作聚解聚作聚合incorporatedinto蛋白质结合，使肌蛋白不能与ADP结合肌动白丝的

肌动蛋白结合点上结合了其他蛋能够聚 不能聚§6.2.2microfilament-associatedprotein（MAP,微丝结合蛋白）肌动蛋白纤维的三种排列方式应力纤

伪细胞皮收缩 胶状网

紧密平行α-α-辅肌动蛋 Lightgreen—α-helicalmotifsDarkgreen—β-sheetmotifs肌球蛋白:依赖于微丝的分子马肌肌球蛋白肌动蛋白结合位ATP结合位肌球蛋500个氨基肌球蛋白-氨氨基末绕线式α螺轻羧基颈肌球蛋白-II肌球蛋白轻 肌动蛋白结合位磷酸

自组

15-20肌球蛋白非活化状态：轻链未活化状态：轻链磷肌球蛋白-肌球蛋白-肌球蛋白-质质整合蛋细胞结合到胞外基细胞胞外基 纤连蛋粘着连肌动蛋α-辅肌动蛋封端蛋

纽蛋桩蛋一种与细胞粘着有关纤连蛋白受体（整合蛋白纤连蛋MAPs蛋白质功 举 形状，大小和分子质量上的比

与肌动蛋互作用的加强微

肌动原肌球蛋

负 正

亚单微丝

毛缘蛋α-辅肌动蛋将微丝相连形成滑动微

肌球蛋白-微丝结合到质膜上的位点血影蛋固定肌动固定肌动蛋单微丝结合蛋白间的相互作细丝蛋 相连成网肌动蛋原肌球蛋

收缩性肌动蛋白微绒无定型组织，深 肌动蛋白丝质侧臂（肌白-1，钙调蛋白横向连接（绒毛蛋白，毛缘蛋白细收缩培养纤维原应力§6.2.3肌alsocalledmusclefibersbythefusionofmanyseparatecellsthenucleiareretainedinthislargecellandliejustbeneaththeplasmathebulkofthecytoplasmismadeupof肌纤细胞肌原纤 Severalcentimeterslong肌小 暗 亮A I

细纤维（肌动蛋白 contractileunitofmyofibrilZdisc（Z盘ateitherend entsiteforactinMline（M线thelocationofspecificproteinsthatlink filamentstoeachotherLightband（明带containonlyactinDarkband（暗带locationof 双股α-

氨基末轻 肌球蛋白 aggregatetogetherbymeansoftheirtailHeadsof projectingtotheBarezoneinthecenterofthefilament tirely肌小肌肌球蛋白粗丝在肌小Z盘是肌动蛋白丝的正肌动蛋白丝的负TheplusendofeachactinfilamentisembeddedintheZdiscTheminusendpointstowardthemyosin肌肉收amyosinheadlackingaboundnucleotideislockedtightlyontoanactinfilament ATPbindstothemyosinchangetheconformationoftheactin-bindingreducestheaffinityoftheheadforactinandallowsittomovealongthefilamenthydrolysisofATPADPandPiremaintightlyboundtotheweakbindingofmyosinheadtoanewsiteontheactincausesthereleaseofPiforce-generatingchangeinheadregainsitsoriginalconformation,losesmyosinheadmovedtoanewpositionontheactin,isagainlockedtightlytotheactin肌动蛋白负 正结 肌球蛋白部松ATP头部竖产生动位点形成弱的结Ca2+肌原纤Ca质膜上内陷的横向电压敏感

去极化的T－微管细胞肌质网

Ca离 通肌钙蛋白和肌球蛋白调节骨骼肌收缩时的肌钙蛋白复合肌动蛋

原肌球蛋

原肌球蛋白阻碍球蛋白接合

DefinitiveandpositedHCMSeidmanetal.,CircRes2011,108:743-HollowΦ24onlyineucaryoticSpidle（纺锤体Basalbody（基粒Centrosome（中心粒Cilia（纤毛）,flagemmae（鞭毛Axon（轴突§6.3.1Tubulin（微管蛋白Heterodimer(αbasicunitofmicrotubuleα-tubulin:450aa,50KDβ-tubulin:455aaacidicC-αβ αβ αβmicrotubuleiscomposedof13原纤微管蛋白分装Basalbody

间期细

纤毛Minusend(-)inPlusend(+)neartheplasma

期细纺神经细中心GTP-containingsubunitsareforcedintoalinearconformationcurvedconformationTubulinconcentration:GTPhydrolysisafterpolymerizationdestabilizes微管蛋白二可交换的直原纤

GTPGTP水解改变亚单位构并减弱聚合物之间的弯曲的原包含GDP-微管蛋白二解 的微管区域较不稳GDP-GTP生 降Microtubule-associatedproteins(MAPs,微管相关蛋白)BindingtoModifymicrotubules’Microtubulemotorproteins（微管动力蛋白正 负驱动蛋 动力蛋Kinesin（驱动蛋白Dynein（动力蛋白 enttomicrotubulesinanATP-dependentfashion otorproteinhastwoheavychainsandseveralsmalllightchainsEachheavychainformsaglobularheadthatinteractswithmicrotubulesMotorproteinwalksATP-dependentlyalongmicrotubulesMotorproteinmovealong（动力蛋白沿着微管移动货货物分受驱动动力蛋负正动力蛋微受货物动力KinesinmovetowardtheplusendofaDyneinmovetowardtheminusVesicletransportintwo驱动蛋白沿微管运动的步行模§6.3.2cilia（纤毛）andArrangementofmicrotubulesinaciliumor

ninespecialdoubletmicrotubulesarearrangedinaringaroundapairofsinglemicrotubulessinglecompletemicrotubule(13doubletonecompletemicrotubule(13subunits)onepartialmicrotubule(11subunits)Dyneindrivesthemovementsofcilia马达蛋白介导的纺锤体的行§6.3.3centrioles（中心粒）andbasalbody（基粒）Cylindricalstructure（柱状结构~0.2μmwideand0.4μmninegroupsofthreemicrotubules,fusedintotripletsformthewalloftheadjacenttripletsarelinkedatintervalsalongtheirlengthDuplicationof thetwomembersofapairThenadaughtercentrioleisformedperpendiculartoeachoriginalcentrioleAnimmaturecentriolecontainsaninefoldsymmetricarrayofsinglemicrotubules icrotubuleactsasatemplatefortheassemblyofthetripletmicrotubuleofmaturecentrioles.Anewlyreplicatedpairof §6.4intermediatefilaments§6.4.1cytosolicIfs（细胞质Monomer（单体C-terminalglobulartailcenterelongatedrodDimer（二聚体Tetramer（四聚体side-by-sideinastaggeredfashionwiththeirN-andC-terminalpointinginopposite(antiparallel)vianon-covalentRopelikeIF（绳状end-to-endandside-by-sidevianon-covalentbondingα螺旋区双股螺旋的二聚双股螺旋二聚体交错两个四聚体组合在PropertiesanddistributionofTissuespecific（组织特异Evol